New carnivorous sponges from the Great Barrier Reef, Queensland, Australia collected by ROV from the RV FALKOR

John N . A . Hooper

This research presents three new species of carnivorous sponges from the family Cladorhizidae from the Great Barrier reef, in Queensland, Australia: Abyssocladia falkor sp. nov., Abyssocladia jeanvaceleti sp. nov. and Axoniderma wanda sp. nov. They were collected by roV during the expedition Fk200802-Seamounts, Canyons & reefs of the Coral Sea Cruise on the rV Falkor from the Schmidt ocean Institute. In addition, the roV collection of two complete specimens enabled the redescription of two other Australian species of carnivorous sponge (Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a and Asbestopluma (Asbestopluma) maxisigma Ekins et al., 2020a), previously known from the East coast of Australia based on incomplete specimens.

Zootaxa 5293 (3): 435–471 https://www.mapress.com/zt/ Copyright © 2023 Magnolia Press ISSN 1175-5326 (print edition) Article ZOOTAXA ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.5293.3.2 http://zoobank.org/urn:lsid:zoobank.org:pub:FE67E8C2-AFE5-491C-B673-2ECE82FA4D87 New carnivorous sponges from the Great Barrier Reef, Queensland, Australia collected by ROV from the RV FALKOR MErrICk EkINS1,2,3* & JohN N. A. hooPEr1,3,4 1 Queensland Museum, PO Box 3300, South Brisbane 4101, Brisbane, Queensland, Australia School of Biological Sciences, University of Queensland, St Lucia, Queensland, 4072 Australia 3 Griffith Institute for Drug Discovery, Griffith University, Brisbane 4111, Queensland, Australia 4� john.hooper@qm.qld.gov.au; https://orcid.org/0000-0003-1722-5954 * Corresponding author. � m.ekins@qm.qld.gov.au; https://orcid.org/0000-0002-4825-462X 2 Abstract This research presents three new species of carnivorous sponges from the family Cladorhizidae from the Great Barrier reef, in Queensland, Australia: Abyssocladia falkor sp. nov., Abyssocladia jeanvaceleti sp. nov. and Axoniderma wanda sp. nov. They were collected by roV during the expedition Fk200802—Seamounts, Canyons & reefs of the Coral Sea Cruise on the rV Falkor from the Schmidt ocean Institute. In addition, the roV collection of two complete specimens enabled the redescription of two other Australian species of carnivorous sponge (Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a and Asbestopluma (Asbestopluma) maxisigma Ekins et al., 2020a), previously known from the East coast of Australia based on incomplete specimens. Key words: Cladorhizidae, Abyssocladia, Asbestopluma, Axoniderma, Cladorhiza, Chondrocladia, Cladorhizidae, Great Barrier reef, Coral Sea, Schmidt ocean Institute, CSIro, Carnivorous Sponges, Exaptation Introduction Since the discovery of carnivory in sponges (Vacelet & Boury-Esnault 1995), there has been an explosion in the discovery and description of new species. This has been largely aided by the development of remote operated Vehicles (roVs) capable of reaching bathyal and abyssal depths in which these species commonly live. roVs allow the visualization of the entire animal in situ and before these delicate minute sponges become inadvertently damaged during collection as is often the case using bottom trawling collecting gear. Indeed, in this present study the roV photography and delicate suctioning collection methodology enabled complete specimens to be collected. In addition to discovering three new species of carnivorous sponges, two other previously known species (Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a and Asbestopluma (Asbestopluma) maxisigma Ekins et al., 2020a) were collected by the rV Falkor expedition and are redescribed here based on fully intact specimens. These species were originally collected using bottom trawl gear by the rV Investigator Cruise along the East Coast of Australia in 2017 and their original descriptions were based on incomplete specimens (Ekins et al. 2020a). The three new species described here from the Great Barrier reef are Abyssocladia falkor sp. nov., Abyssocladia jeanvaceleti sp. nov. and Axoniderma wanda sp. nov. Abyssocladia falkor sp. nov. is named in honour of the research vessel that enabled the discovery of these species, and Abyssocladia jeanvaceleti sp. nov. named in honour of the scientist who first discovered carnivorous sponges and has described many new species from the Pacific and elsewhere. Prior to the discovery of the exaptation in sponge spicules as they evolved into a carnivorous feeding strategy from a filter feeder, there were excellent descriptions of the Cladorhizidae sponges, particularly the impressive works of ridley & Dendy (1886, 1887), and all of the crew involved in the Challenger Expedition (1872–1876) to bring these tiny delicate creatures to the surface and to produce such accurate descriptions and illustrations. The new species described in this present work were collected by the scientific expedition Fk200802— Seamounts, Canyons & reefs of the Coral Sea Cruise on the rV Falkor from the Schmidt ocean Institute and Accepted by M. Klautau: 22 Jan. 2023; published: 23 May 2023 435 represent only a small number of dives made by the roV SuBastian that were specifically deployed to scan the depths for these sponges (Fig. 1). Previous collections of cladorhizids from Queensland include Lycopodina coralseaensis Ekins et al., 2021 from the rV Investigator cruise IN2019_V04, Abyssocladia desmophora (hooper & Lévi, 1989) discovered during the CIDArIS expeditions, and Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a from the rV Investigator IN2017_V03 Cruise. The addition of these three new species in the present work brings the known fauna of Cladorhizidae in Australian waters to 29. All the carnivorous sponges in Australian waters were previously compared in Table 1 of Ekins et al. (2021). here we report on an additional three new species of sponges discovered from the Queensland, Australia, an area surveyed using an roV to photograph and harvest the marine biota from previously unexplored mesophotic to bathyal depths. Materials and methods The material was collected by the roV SuBastian aboard the Schmidt ocean Institute research vessel the rV Falkor during the Fk200802—Seamounts, Canyons & reefs of the Coral Sea Cruise (Fig. 1). The specimens were collected by suction and stored in seawater, and then immediately transferred to 70 % ethanol. FIGURE 1. Map of the cruise showing the collection locations and depths of the species described. 436 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr A video of the roV collection of these specimens is available at this link: roV SuBastian Dive 399 (Pt A)— Southern Small Detached reef, Australia—Fk200930—Bing video Morphological analysis. Scanning Electron Microscope spicule preparations were made by dissolving the tissue in 12.5% sodium hypochlorite to remove the soft tissue, and neutralized in distilled water, rinsed again in distilled water, rinsed twice in 70% ethanol, and then finally rinsed twice in 98% ethanol and air dried. SEM preparations were sputter coated in gold to improve resolution. The scanning electron micrograph photos and measurements were made using a hitachi TM-1000 SEM and plates assembled in Adobe Photoshop. The spicule size ranges are from those spicules that were the largest and smallest of all spicules from all specimens. Specimens in the laboratory were photographed using a Canon EoS 7D. Specimens were examined under an olympus SZ60 dissection microscope and photographed with a Tucsen 3.0 camera. Results Genus Chondrocladia Thomson, 1873 Chondrocladia Thomson, 1873: 188 Crinorhiza Schmidt, 1880: 83 Meliiderma ridley & Dendy, 1887: 102 not Neocladia koltun, 1970: 193; Vacelet et al. 2009: 59. Diagnosis. Cladorhizidae with anchorate isochelae (from Lee et al. 2012). Type species: Chondrocladia virgata Thomson, 1873 (by monotypy). Subgenus Chondrocladia Thomson, 1873 Diagnosis: Chondrocladia without a layer of special spicules (subtrochirhabds or trochirhabds), lacking special rostriform (snoutlike) subtylostyles in filaments or terminal balls, and without planar vanes formed of evenly spaced upright branches (from Lee et al., 2012). Type species: Chondrocladia virgata Thomson, 1873 (by monotypy). Remarks. The genus Chondrocladia was summarised in Table 6 of Lopes et al. (2011), and updated to include additional information and recently described species in Table 10 in Ekins et al. (2020a). Chondrocladia (Chondrocladia) zygainadentonis Ekins, Erpenbeck & Hooper, 2020 Figures 2–3 Material examined: holotype QM G337557, off Gladstone, Coral Sea, Queensland, Australia, Station 12, 23.6311944 – -23.65900, 154.659694 – 154.643806, 1770–1761 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_V03, Sample 128-111, 13/vi/2017. other Material: QM G339304, ribbon reef 5, Canyon 8, Great Barrier reef Queensland, Australia, 15.36606511, 145.8662834, 1526.89 m, Site: S0378, Sample: 55, roV SuBastian, Coll. Jeremy horowitz on RV FALKOR, cruise Fk200802, 14/8/2020 Distribution: This species is presently known only from the Coral Sea and Great Barrier reef, off the Northeast coast of Queensland, bathyal depth. Description: Growth form: An erect delicate single-axis arborescent sponge resembling a tree consisting of a cylindrical stem with five columns of branches (Figs. 2 A, B). The sponge including the roots is 12 cm in height, the branches are up to 4 cm in length and the roots are also up to 4 cm in length. The stem base is short, only 12 mm in length and 2 mm in width. Each branch contains four alternate filaments (Fig. 2 C). The branches are between 0.5 and 1 mm in thickness. The filaments are up to 2.5 mm long and between 80 and 270 µm wide. Colour: white in situ, on deck and preserved in ethanol. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 437 FIGURE 2. Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a. A. the specimen QM G339304 underwater in vivo. B. Specimen QM G339304 fixed. C. Branch with filaments under a dissection microscope. D. Branch with filaments under scanning electron microscope. E. Filament with the ectosomal skeleton of tridentate unguiferate isochelae. F. root like processes, showing the bundles of styles, with no specialised ectosomal layer. Ectosomal skeleton: The ectosomal skeleton consists of a thin membrane containing chelae (Fig. 2 E). Choanosomal skeleton: The choanosomal skeleton consists of bundles of mycalostyles longitudinally arranged in the axis of the stem. The stem also contains the subtylostyles and rare thin subtylostyles. The filaments consist of longitudinally arranged subtylostyles, with radial arrangement for support against the stem. The roots consist of the same combination of styles bundled together as in the stem, but also include a smaller blunt style (Fig. 2 F). 438 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr FIGURE 3. Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a. QM G339304. A. Tridentate unguiferate isochelae. B. Large unguiferate isochelae. C. Mycalostyle. D. Close up of the mycalostyle ends in C. E. Subtylostyles. F. Close up of the subtylostyle ends in E. G. Styles in the root like appendages. h. Close up of the ends of the styles in G. I. Very thin supportive styles. J. Close up of the ends of the styles in I. k. Thin subtylostyles. L. Sigmancistras. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 439 Megascleres: Larger mycalostyles with tapering ends and a blunt point (872-(1109)-1280 x 14.9–(23.9)–36.0 µm, n=46) (Fig. 3 C, D). Subtylostyles with slightly swollen bases and tapering points (459–(558)–669 x 8.4– (12.2)–7.51 µm, n=34) (Figs. 3 E, F). rare thin subtylostyles (479–(763)–1060 x 3.0–(4.4)–6.3 µm, n=5) (Figs. 3 I, J). In the roots are blunt styles (149–(256)–371 x 1.0–(3.8)–7.1 µm, n=8) (Figs. 3 G, h). Microscleres: Abundant small tridentate unguiferate isochelae with equal sized alae (25.5– (33.9)–42.9 x 2.1– (3.2)–4.5 µm, n=41) (Fig. 3 A). uncommon large unguiferate isochelae, with variable dentation (often 3), 44.1– (53.9)–67.9 x 3.3–(5.7)–7.1 µm, n=16 (Fig. 3 B). rare thin sigmas 23.1–(27.1)–31.1 x 0.9–(1.0)–1.2 µm, n=2 (Fig. 3 k). rare sigmancistras 24.7–(26.5)–27.7 x 1.9–(2.5)–3.3 µm, n=4 (Fig. 3 L). Remarks: Despite the appearance of the holotype resembling the gross morphology of Cladorhiza abyssicola Sars, 1872, C. (C.). zygainadentonis has isochelae, and it also lacks sigmas and sigmancistras. The presence of isochelae indicates it clearly belongs in Chondrocladia, as illustrated in the figures and the diagnosis of hestetun et al. (2016a) and Ekins et al. (2020b). This species is Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a, with the unique unguiferate anchorate isochelae. The only other Chondrocladia (Chondrocladia) with some sort of arborescent morphology is Chondrocladia (Chondrocladia) dichotoma Lévi, 1964; but that species has quadridentate unguiferate isochelae. The redescription of C. (C.). zygainadentonis includes the addition of the styles in the root like appendages. Genus Abyssocladia Lévi, 1964 Abyssocladia Lévi, 1964: 78 Diagnosis. Cladorhizidae most often pedunculate, carrying a disciform or flabelliform body with a radial architecture, in other cases pinnate or branching. Microscleres are a combination of abyssochelae, cleistochelae, arcuate chelae and/or sigmancistras, but not placochelae (from hestetun et al. 2016a). Type species. Abyssocladia bruuni Lévi, 1964 (by monotypy). Remarks. Currently there are 38 accepted species assigned to Abyssocladia (de Voogd et al. 2022), with only five previously recorded from the Australian EEZ i.e. A. annae Ekins, Erpenbeck & hooper, 2020a, A. desmophora (hooper & Lévi, 1989), A. escheri Ekins, Erpenbeck & hooper, 2020a, A. gliscofila Ekins, Erpenbeck & hooper, 2020a and A. oxyasters Ekins, Erpenbeck, Goudie & hooper, 2020b. Three new species of Abyssocladia were also recently reported from New Caledonia (Vacelet 2020). They are A. kanaconi Vacelet, 2020, A. microstrongylata Vacelet, 2020 and A. mucronata Vacelet, 2020 (the latter now accepted as Echinostylinos gorgonopsis Lévi, 1993 (Vacelet & kelly 2022). Abyssocladia falkor sp. nov. Figures 4–5. Tables 1, 3 urn:lsid:zoobank.org:act:B58AB630-A45C-4DCB-AE83-8BB776394CAF Material Examined: holotype: QM G339306, ribbon reef 5, Canyon 8, Great Barrier reef, Queensland, Australia, -15.35981103, 145.862097, 1822.14 m, Site: S0378, Sample: 54, roV SuBastian, Coll. Merrick Ekins remotely in Brisbane via live satellite feed directing the roV pilots onboard the RV FALKOR on the Great Barrier reef and Jeremy horowitz on RV FALKOR, cruise Fk200802. Paratype: QM G339307, same collection details as QM G339306. Etymology: falkor, a noun in apposition, named in honour of the Schmidt ocean Institutes research vessel the rV Falkor. Distribution: This species is currently known from the Coral Sea in Queensland, Australia, bathyal depth. Description: Growth form: The holotype consists of a pedunculated sponge, with a short stem and a disc shaped body with filaments radiating out in a single plane from the disc margin. (Figs. 4 A–C). The paratype has the entire stem and roots as well, which were lost from the holotype during collection. The holotype has a body diameter of 7.8 mm, and 1.3 mm thick. The filaments are 2.3 mm long and 0.1 mm in width. The paratype has a body with a 6.7 mm diameter and 1.6 mm width, the stem and root are 40 mm in length, the filaments are up to 4.0 mm in length. 440 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr Colour: Cream in situ, on deck and in ethanol. Ectosomal skeleton: Thin membranous layer of the filaments is encrusted with small sigmancistras and abyssochelae. The sponge body and stem are encrusted with the predominately spherical microstrongyles, but also include the abyssochelae. Immediately under the encrusting layer are the sinuous styles. The ectosomal skeleton of the roots is a thin membranous layer with occasional abyssochelae, sigmas and sigmancistras. Endosomal skeleton: The axis of the stem consists of longitudinally arranged mycalostyles, styles and sinuous styles (Figs. 5 D, F, J). The main body consists of the radiating bundles of subtylostyles that form the filaments (Figs. 5 h, I). within the body between the bundles, the styles form a disorganised almost cross hatching skeleton. The roots are composed of the large mycalostyles, styles, sinuous styles, occasional subtylostyles and rarely strongyles (Fig. 4 G). The filaments axis is composed of subtylostyles only (Fig. 4 D). FIGURE 4. Abyssocladia falkor sp. nov. A. The holotype and paratype underwater in vivo. B. Paratype QM G339307 fixed. C. holotype QM G339306 on deck. D. Filaments emerging from the edge of the sponge body. E. Surface of the sponge body. F. Stem. G. root like appendages. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 441 FIGURE 5. Abyssocladia falkor sp. nov. A. Abyssochelae. B. Large thin sigma. C. Intermediate sigma. D. Large mycalostyles. E. Ends of the mycalostyle in D. F. Styles. G. Ends of styles in F. h. Subtylostyles. I. Ends of subtylostyle in h. J. Sinuous styles. k. Ends of styles in J. L. Strongyles. M. Ends of the strongyles in L. N. Small sigmancistras. o. Spherical microstrongyles. P. Dildo-shaped tylomicrostrongyle. 442 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr Megascleres: Mycalostyles are long and generally straight, sometimes with a slight curve, and have a rounded point (Figs. 5 D, E). Styles usually straight, sometimes with a slight curve, thickest in the centre also with a blunt point, sometimes with a very faint tyle swelling (Figs. 5 F, G). They are separated from the mycalostyles often by their location in the body but could well be a continuum of a single category of megasclere. The sinuous styles occur around the mycalostyles that make up the stem, under the dermal microstrongyles (Figs. 5 J, k). Subtylostyles are long and straight, with a sharp tip and compose the filaments (Figs. 5 h, I). There are also very rare strongyles present from the roots (Figs. 5 L, M). Microscleres: Abyssochelae isochelae with curved shaft, and opposing alae touching (Fig. 5 A). There are small sigmas and small sigmancistras that usually occur on the exterior surfaces (Figs. 5 C, N, respectively). There are also long thin sigmas that only occur on the sponge body, these were rare and only occurred on the paratype and so may not be native (Fig. 5 B). The predominately spherical to subspherical shaped microstrongyles are microspined (Figs. 4 E, F; 5 o). however, they are also dumbbell-shaped and rarely dildo—shaped (Fig. 5 P). The microstrongyles have either a rounded end (tylomicrostrongyle) or a sharp point (tylomicrostyles). Remarks: This species is most closely related to the Japanese species A. natsushimae Ise & Vacelet, 2010. however, this new species has a much smaller and circular body size. The new species also lacks the two size classes of abyssochelae, has subtylostyles, and has only one category of sigmancistras and two categories of sigmas. The microstrongyles of this new species are predominately spherical and lacks the long microstrongyles of A. natsushimae. This species is different from the diminutive species A. bruuni Lévi, 1964, which has the very large abyssochelae. Abyssocladia bruuni was also recorded by koltun (1970), but with two size classes of abyssochelae. Abyssocladia bruuni also lack microstrongyles. This new species also has similar abyssochelae and a rough morphological similarity to A. annae Ekins et al., 2020a and another closely related species A. fryerae hestetun et al., 2019, both of which have this ridged abyssochelae as well as subtylostyles and sigmancistras. TABLE 1. Comparison of spicule measurements smallest–(average)–largest (µm) for Abyssocladia falkor sp. nov. Specimen Mycalostyles oxetote styles Subtylostyles Sinuous styles Strongyles G339306 813–(1020)–1450 x 11.4–(17.4)– 30.7, n=32 365–(408)–583 x 9.4–(12.4)–14.7, n=12 276–(511)–762 x 4.7–(8.6)–11.7, n=13 256–(367)–540 x 7.2–(10.0)–13.5, n=25 _ G339307 715–(811)–1020 x 10.0–(13.4)–25.9, n=9 281–(408)–621 x 6.4–(9.6)–14.3, n=45 192–(426)–830 x 3.2–(7.9)–12.3, n=39 222–(375)–497 x 4.9–(9.6)–12.9, n=23 220–(355)–490 x 5.5–(10.1)–14.7, n=2 Continued. Specimen Abyssochelae (length x spine width) Long thin sigmas Sigmas Sigmancistras Spherical microstrongyles Dildo/Tylo– microstrongyles G339306 39.6–(44.9)– 51.9 x 3,7– (5.6)–8.6, n=28 _ 20.5–(26.1)– 32.9 x 1.1–(2.1)–3.1), n=32 8.0–(9.2)–10.9 x 0.5–(0.9)– 1.4, n=32 8.8–(14.0)–23 x 6.9–(9.7)–12.5, n=37 9.0–(24.4)–60 x 3.9–(8.5)–12.4, n=18 G339307 27.5–(40.7)– 52.8 x 1.9– (5.1)–6.9, n=38 50.6–(58.2)– 67.2 x 0.8– (1.1)–1.5, n=5 16.8–(24.8)– 33.1 x 1.6– (2.1)–2.6, n=20 6.9–(8.8)–10.4 x 0.7–(1.1)– 1.8, n=22 8.4–(14.4)–26.0 x 5.8–(8.7)– 12.9, n=32 17.9–(43.3)– 73.3 x 2.8– (4.8)–7.8, n=27 Abyssocladia jeanvaceleti sp. nov. Figures 6–7, Tables 2–3 urn:lsid:zoobank.org:act:0D481EB3-F3F9-4B0F-9D9B-DC57A25D8E42 Material Examined: holotype: QM G339390, Small Detached Plunge Pool, Great Barrier reef, Queensland, Australia, -12.5341061, 143.8600016, 1082.83 m, Site: S0399, Sample: 141, roV SuBastian, Coll. Mardie McNeil and rob Beaman on RV FALKOR, cruise Fk200802. 18/X/2020 NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 443 Paratypes: QM G335997, QM G339387, QM G338728, same collection details as QM G339390. other Material: QM G339391 same collection details as QM G339390, sample 141C. Etymology: Named in honour of Jean (Janus) Vacelet who as both a gentleman and a scholar has described many of the South Pacific carnivorous sponges, and proved the existence of carnivory in sponges. Distribution: This species is currently known from the Great Barrier reef in Queensland, Australia, bathyal depth. Description: Growth form: The holotype consists of a pedunculated sponge, with a short stem and a vertically orientated disc shaped body with filaments radiating out in a single plane from the disc margin. (Figs. 6 A–C). The circular sponge bodies were 6 to 9 mm in diameter, and 0.4 to 0.6 mm thick. There were up to 50 filaments surrounding the body. The preserved filaments were 4.3 to 9.3 mm long and 0.1 mm in width, and highly contracted and or destroyed during collections, compared to the underwater images which show the filaments as up to one and a half times the width of the body (Figs. 6 A, B). The stem lengths were 14 to 22 mm long and 0.4 mm in width. All the specimens had spherical structures within the sponge body, which are most likely to be spermatocytes or oocytes. Some of the specimens such as G338728 were still attached via a holdfast to the underlying rock strata. TABLE 2. Comparison of spicule measurements smallest–(average)–largest (µm) for Abyssocladia jeanvaceleti sp. nov. Specimens Mycalostyles (Filaments) Styles (Filaments) Styles L (Stem) Styles M (Stem) QM G339390 876–(1118)–1480 x 11.9–(15.9)–23.2, n=28 364–(519)–793 x 5.0–(7.7)–12.3, n=30 1060–(1243)–1470 x 18.3–(24.7)–29.8, n=20 387–(673)–962 x 19.1–(26.8)–35.1, n=17 QM G335997 971–(1114)–1240 x 12.6–(14.5)–15.2, n=5 413–(462)–477 x 6.4–(6.9)–7.4, n=13 621–(1006)–1620 x 12.6–(18.9)–24.5, n=9 QM G339387 738–(1223)–1710 x 11.9–(16.8)–24.8, n=45 412–(664)–898 x 4.2–(9.4)–12.9, n=21 QM G338728 1060–(1140)–1260 x 18.2–(21.5)–26.8, n=5 507–(556)–629 x 6.0–(9.7)–11.8, n=5 Cf. QM G339391 700–(991)–1250 x 12.1–(17.0)–21.0, n=28 518–(661)–700 x 5.6–(9.6)–16.0, n=14 Continued. Specimens Strongyles (root) Isochelae (length x spine width) Sigmancistras QM G339390 315–(478)–678 x 7.5–(11.0)– 26.3, n=21 44.6–(52.6)–61.4 x 3.5–(4.1)– 5.0, n=34 8.1–(11.2)–12.9 x 0.6–(1.4)– 2.0, n=28 QM G335997 42.6–(53.0)–61.2 x 3.5–(4.1)– 6.0, n=24 10.9–( 15.9 )–25.3 x 1.2–(1.3)– 1.4, n=3 QM G339387 24.5–(50.4)–57.9 x 3.9–(4.4)– 5.0, n=32 10.2–(12.3)–26.9 x 1.0–(1.4)– 2.0, n=27 QM G338728 46.3–(53.1)–59.0 x 3.5–(4.3)– 5.0, n=11 9.7–(10.1)–11.0 x 1.4–(1.4)– 1.5, n=12 Cf. QM G339391 57.9–(106.2)–138.0 x 7.8– (12.9)–19.8, n=63 9.1–(11.7)–14.8 x 1.0–(1.4)– 1.8, n=28 Colour: white in situ, on deck and in ethanol. Ectosomal skeleton: Thin membranous layer of the filaments is encrusted with small sigmancistras (Fig. 6 F). The sponge body and stem are encrusted isochelae overlaying the sigmancistras (Fig. 6 E). The lower stem and roots appear to be devoid of any specific ectosomal layer. Endosomal skeleton: The axis of the stem consists of longitudinally arranged styles. The main body and filaments consist of the radiating bundles of styles that form the filaments. The root axis consists of longitudinally arranged strongyles (Figs. 6 G, h). 444 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr FIGURE 6. Abyssocladia jeanvaceleti sp. nov.. A. underwater photo the specimens before collection. Arrows indicating individuals of the same species. The black arrow is the holotype QM G339390. B. A close up of the species in side profile. C. holotype QM G339390 on the right and paratype QM G335997 on the left. D. Scanning Electron Microscope image of the body face and filaments emerging. E. Close up of the ectosome of the sponge body showing the isochelae on the surface and the sigmancistras covering the styles of the filaments. F. Close up of the filaments showing the ectosomal membrane composed of sigmancistras over the style axis. G. The stem showing the axial bundles of styles. h. The root like appendage. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 445 FIGURE 7. Abyssocladia jeanvaceleti sp. nov.. holotype: QM G339390. A. Isochelae. B. Sigmancistras. C. Mycalostyle from the filaments with faint subtylostyle swelling. D. Magnified ends of the mycalostyle illustrated in C with faint subtylostyle swelling. E. Smaller mycalostyle from the filaments. F. Magnified ends of the mycalostyle illustrated in E. G. oxeote style from the stem. h. Magnified ends of the oxeote style illustrated in G. I. Thin sharp oxeote style from the stem. J. Magnified ends of the oxeote style illustrated in I. k. Strongyle from the root like appendages. L. Magnified ends of the strongyle illustrated in k. 446 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 3. Comparative morphological and distributional data for all stalked disc–shaped species of Abyssocladia. Zootaxa 5293 (3) © 2023 Magnolia Press · Species reference Morphology Total height x width (mm) Skeleton Abyssocladia falkor nov. sp This work Pedunculate, short erect stem and a vertically orientated disc shaped body with filaments radiating out in a single plane from the disc margin 50 x 10 Dense longitudinal bundles of mycalostyles and styles in the axis of peduncle, body with radial skeleton, bundles of mycalostyles extending into filaments, roots strongyle bundles Abyssocladia jeanvaceleti sp. nov. This work Pedunculate, short erect stem and a vertically orientated disc shaped body with filaments radiating out in a single plane from the disc margin 40 x 30 Dense longitudinal bundles of mycalostyles in the axis of peduncle, body with radial skeleton, bundles of mycalostyles extending into filaments, roots strongyle bundles Abyssocladia annae Ekins, Erpenbeck & hooper, 2020 Ekins, Erpenbeck & hooper, 2020: 23–25, Fig. 4 Pedunculate, erect stem supporting a slightly cupped-shaped obovate (leaf-like, flabellate, fan-shaped) apical body, lacking lateral filaments 3x2 Peduncle axis longitudinally arranged subtylostyles, body with radiating subtylostyles projecting only in one quadrant Abyssocladia carcharias kelly & Vacelet, 2011 kelly & Vacelet, 2011: 58–60, Figs. 2–3 Pedunculate, long thin, flattened circular body, with short blunt radiating filaments 7 x 3.4 Dense longitudinal bundles of mycalostyles in the axis of peduncle, body with radial skeleton, bundles of mycalostyles extending into filaments Abyssocladia dominalba Vacelet, 2006 Vacelet, 2006: 575–577, Fig. 14 Long thin pedunculate with an ovoid or subspherical body, body with dense spicular bundles laterally radiating 28–31 Peduncle axis with longitudinally arranged long fusiform styles, supported by radiating bundles of fusiform styles and smaller styles with tips directed outwards Abyssocladia fryerae hestetun, rapp & Pomponi, 2019 hestetun, rapp & Pomponi, 2019:4, Fig. 2 Pedunculate, short stem, disc-shaped dropletlike body, radiating long filaments in a single plane from disc margin 25 x 12 Stem with tightly packed subtylostyles entering center of disc-shaped body with radial skeleton, filaments and disc also composed of subtylostyles Abyssocladia huitzilopochtli Vacelet, 2006 Vacelet, 2006: 569–573, Figs. 9–11 Erect pedunculated, enlarged base, long thin peduncle, flattened semicircular disc-like body, with numerous free radiating spicule fascicles protruding 41 x 0.3– 0.6 Axis of peduncle tightly packed with longitudinal bundles of long substrongyles, body with radiating bundles and irregularly dispersed large substrongyles, filaments with smaller bundles of substrongyles Abyssocladia inflata Vacelet, 2006 Vacelet, 2006: 573–575, Figs. 12–13 Thin pedunculate, flattened discform body, hispid surface, with short filaments 8.5 x 3.5 Axis with longitudinally arranged styles extremely reduced living tissue, body with skeleton of radiating styles protruding from surface ......continued on the next page 447 448 · Zootaxa 5293 (3) © 2023 Magnolia Press TABLE 3. (Continued) Species reference Morphology Total height x width (mm) Skeleton Abyssocladia kellyae hestetun, rapp & Pomponi, 2019 hestetun, rapp & Pomponi, 2019: 5, Fig. 3 Long stalk with disc-shaped body and radial filaments in a single plane on disc margin, enlarged basal plate 43 x 7 Main skeleton with tightly arranged longitudinal mycalostyles, radiating filaments composed of mycalostyles, subtylostyles and tylostyles within sponge body Abyssocladia lakwollii Vacelet & kelly, 2014 Vacelet & kelly, 2014: 387–392, Figs. 1–3 Pedunculate, flattened disc attached to a thin peduncle with radiating filaments forming a flat to concave umbrella up to 57 Peduncle tightly packed longitudinal bundles of large mycalostyles, basal attachment tightly packed shorter mycalostyles 1 and substrongyles, mycalostyles 1 of the peduncle fan out in the disc, diverging into radiating bundles towards the rim, disc with many small diverging bundles of mycalostyles 2 Abyssocladia oxyasters Ekins, Erpenbeck, Goudie & hooper, 2020 Ekins, Erpenbeck, Goudie & hooper, 2020: 247–251, Figs. 4–6 Pedunculate, erect, long thin stem with plano-convex disc shaped body on apex encircled by radiating filaments, and conical basal disc holdfast 140–160 x 0.6 Axis of stem and filaments with longitudinal bundles of mycalostyles 1, basal holdfasts contain smaller thicker curved oxeote anisostyles 2 Abyssocladia natushimae Ise & Vacelet, 2010 Ise & Vacelet, 2010:889–892, Figs. 2–5 Erect pedunculate long stem on circular base, mop-like inflated apical body with long filaments in one plane, ending with inflated bulbous tips 88 x 1–2.2 Base cored by substrongyles, short microstrongyles and microscleres, axis of peduncle tightly packed with long mycalostyles longitudinally and spirally arranged, upper part of peduncle covered by soft tissue packed with microstrongyles and few microscleres, axis of filaments supported by bundles of mycalostyles and microstrongyles Abyssocladia polycephalus hestetun, Pomponi & rapp, 2016 hestetun, Pomponi & rapp, 2016: 523–525, Figs. 2–3 Erect pedunculate central stem with side branches each ending in a disclike body bearing filamentous projections 35 Densely packed bundles of mycalostyles in the central stem and branches, radiating bundles of mycalostyles projecting from the body and constituting the skeleton of the filaments, disc-shaped body also with a network of less well organized subtylostyles Abyssocladia stegosaurensis hestetun, rapp & Pomponi, 2019 hestetun, rapp & Pomponi, 2019: 11, Fig. 5 Stalked, vertical disc-shaped radial body, filaments radiating in a single plane from disc margin, 48 x 1–3 core axis of longitudinal, tight tracts of mycalostyles, fleshy parts of radial body with subtylostyles and tylostyles bases, base with styles or substrongyles ......continued on the next page EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 3. (Continued) Zootaxa 5293 (3) © 2023 Magnolia Press · Species Spicules of main axis (Lxw um) Spicules of lateral filaments or body (Lxw um) Spicules of basal attachment (Lxw um) Chelae (L um) Abyssocladia falkor nov. sp Mycalostyles 715–1020 x 10.0–30.7, oxetote styles 282–621 x 6.4–14.7, sinuous styles 222–540 x 4.9–13.5 Subtylostyles 192–830 x 3.2–12.3 Strongyles 220–490 x 5.5–14.7 Abyssochelae 27.5–52.8 x 1.9–8.6 Abyssocladia jeanvaceleti sp. nov. mycalostyles I 620–1620 x 12.6–29.8 mycalostyles II 387–962 x 19.1–35.1 mycalostyles I 738–1710 x 11.9–26.8 mycalotyles II 364–898 x 5.0–12.9 strongyles 315–678 x 7.5–26.3 Isochelae 24.5–61.4 x 3.5–6.0 Abyssocladia annae Ekins, Erpenbeck & hooper, 2020 subtylostyles 288–1000 x 3.1–15.2 undifferentiated absent Abyssochelae 48.2–72.2 x 5.1––14.9 Abyssocladia carcharias kelly & Vacelet, 2011 mycalostyles 510–1070 x 8–19 microstyles 140–240 x 2.5–5 undifferentiated base missing unguiferate abyssochelae 1, 116–197 abyssochelae 2, 60–86 abyssochelae 3, 35–48 Abyssocladia dominalba Vacelet, 2006 styles 620–2500 x 7–35 undifferentiated base missing arcuate isochela 80–170 abyssochelae—cleistochelae 40–45 anisochelae 9.5–11 Abyssocladia fryerae hestetun, rapp & Pomponi, 2019 subtylostyles 582–1130 x 3.8–23.5 undifferentiated undifferentiated arcuate isochelae 77.9– 110.3 Abyssocladia huitzilopochtli Vacelet, 2006 styles—substrongyles 1 (peduncle, filaments), 1050–2500 x 15–30 substrongyles 2 (body, filaments), 260–660 x 5–10 undifferentiated substrongyles 3 (base), 560–750 x 21–30 abyssochelae (body, upper stem) 60–80 arcuate isochelae 1 (body, filaments), 67–90 arcuate isochelae 2 (base of stem), 40–55 Abyssocladia inflata Vacelet, 2006 styles 1075–1800 x 21–33 undifferentiated base missing abyssochelae—cleistochelae 80–100 arcuate isochelae 14–150 Abyssocladia kellyae hestetun, rapp & Pomponi, 2019 mycalostyles 1262–2321 x 19–33 subtylostyles 1018–1994 x 20–35 tylostyles 275–592 x 6–15 undifferentiated arcuate isochelae78– 132 449 ......continued on the next page 450 · Zootaxa 5293 (3) © 2023 Magnolia Press TABLE 3. (Continued) Species Spicules of main axis (Lxw um) Spicules of lateral filaments or body (Lxw um) Spicules of basal attachment (Lxw um) Chelae (L um) Abyssocladia lakwollii Vacelet & kelly, 2014 mycalostyles 1, 750–1800 x 15–31 mycalostyles 2, 330–1150 x 6–20 undifferentiated mycalostyles 1, 380–980 x 18–30 substrongyles 250–1150 x 12–30 anchorate isochelae 1, 110–150 anchorate isochelae 2, 58–92 arcuate isochelae 3, 27–36 cleistochelae 48–70 Abyssocladia oxyasters Ekins, Erpenbeck, Goudie & hooper, 2020 mycalostyles 1, 1380–3810 x 15–54 undifferentiated oxeote anisostyles 2, 392–1560 x 20–50 palmate cleistochelae 62–123 x 17–37 Abyssocladia natushimae Ise & Vacelet, 2010 mycalostyles 1, 1350–(1657)–1940 x 19 – (34)–26.5 microstrongyles 14–(64)–250 x 4–(6)–10 mycalostyles 2, 395 –(1016)–1790 x 10–(16)–24 Sub–strongyles 395–(642)–980 x 22–(36)–45 cleistochelae–abyssochelae 38–(54)–75 Abyssocladia polycephalus hestetun, Pomponi & rapp, 2016 mycalostyles 720–(933)–1070 x 14–(17)–22 subtylostyles-mycalostyles 430–(686)–960 x 5–(10)–13 strongyles 380–(568)–780 x 15–(18)–22 base missing arcuate isochelae 28–(43)–50 Abyssocladia stegosaurensis hestetun, rapp & Pomponi, 2019 mycalostyles 846–1741 x 15–53 mycalostyles same as stem subtylostyles 491–1134 x 7–17 tylostyles 181–573 x 4–9 undifferent–iated palmate isochelae 20–40 ......continued on the next page EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 3. (Continued) Zootaxa 5293 (3) © 2023 Magnolia Press · Species Sigmancistras (L um) Sigmas (L um) and other microscleres Locality, depth range Abyssocladia falkor nov. sp 6.9–10.9 x 0.5–1.8 Sigmas I 50.6–67.2 x 0.8–1.5, sigmas II 16.8–33.1 x 1.1–3.1, Spherical microstrongyles 8.4–26.0 x 5.8– 12.9, Tylostyle microtrongyles 9.0–73.3 x 2.8–8.5 Great Barrier reef, Queensland, Australia, bathyal Abyssocladia jeanvaceleti sp. nov. 8.1–26.9 x 0.6–2.0 absent Great Barrier reef, Queensland, Australia, bathyal Abyssocladia annae Ekins, Erpenbeck & hooper, 2020 11.6 – (15.6)–18.5 absent off the continental shelf of central New South wales, Australia, abyssal Abyssocladia carcharias kelly & Vacelet, 2011 sigmancistras 1, 15–16.2 x 1.9–2 sigmancistras 2, 8–128–12 absent kermadec Seamounts, New Zealand, bathyal Abyssocladia dominalba Vacelet, 2006 1, 30–40 2, 9.5–12.5 absent North-Fijian back-arc Basin, bathyal Abyssocladia fryerae hestetun, rapp & Pomponi, 2019 17.9–22.9 absent Marianas, Nw Pacific, abyssal Abyssocladia huitzilopochtl i Vacelet, 2006 1, 20–24 2, 11–12 orthancistras (body, filaments) 150–195 microxeas (possibly foreign) 30–95 x 0.3–1 Middle America Trench, off Mexico, bathyal Abyssocladia inflata Vacelet, 2006 15–18 Acantho–microxeas 130–350 x 3–5 Easter microplate, East Pacific rise, bathyal Abyssocladia kellyae hestetun, rapp & Pomponi, 2019 22–32 absent Marianas, Nw Pacific, abyssal Abyssocladia lakwollii Vacelet & kelly, 2014 15–20 absent Far eastern Solomon Islands, bathyal Abyssocladia oxyasters Ekins, Erpenbeck, Goudie & hooper, 2020 21.5–40 x 2–7 oxyasters 74–136 Great Australian Bight, bathyal Abyssocladia natushimae Ise & Vacelet, 2010 1, 20–23 2, 9–12 absent Izu-ogasawara Arc, Japan, mesophotic Abyssocladia polycephalus hestetun, Pomponi & rapp, 2016 9.4–(9.8)–11.0 absent Muir Seamount, Bermuda, bathyal Abyssocladia stegosaurensis hestetun, rapp & Pomponi, 2019 6–9 absent Marianas, Nw Pacific, bathyal 451 Megascleres: The sponge filaments are composed of two different but overlapping size groups of styles. The measurements of spicules for all the specimens are listed in Table 2, the spicule measurements for the holotype only are listed in the following text. The large mycalostyles present in the filaments are long and straight, with a blunt tip, they are 880–(1120)–1480 x 12–(16)–23 µm in size, with the slightest tylostyle thickening (Figs. 7 C, D). The smaller styles in the filaments (Figs. 7 E, F) are blunt styles, they are 360–(520)–703 x 5–(8)–12 µm in size. The stem is also composed of two different sizes of oxeote styles, the larger are long and straight, with a slight central thickening, and sharp points (Figs. 7 I, J), which are 1060–(1240)–1470 x 18–(25)–30 µm in size. The shorter oxeote styles of the stem have an obvious central thickening and a blunt tip (Figs. 7 G, h) and are 390–(670)–960 x 19–(27)–35 µm in size. The roots are composed of just one class of strongyle (Figs. 7 k, L) and are 320–(480)–680 x 8–(11)–26 µm in size. Microscleres: The microscleres are composed of a single size class of palmate isochelae (Fig. 7 A) 45–(53)–61 µm in size and a single size class of sigmancistras (Fig. 7 B) 8–(11)–13 µm in size. Remarks: This species is morphologically very similar to Abyssocladia falkor sp. nov., with the same pedunculated disc shaped body. however, it differs markedly by the shape of the isochelae and the lack of the protective microspheres and microstrongyles. Most of the specimens contained filamentous curved raphide-like spicules, which were often on the outside of the ectosome (such as in Fig. 6 E), and sometimes presented as bundles in spicule digests. however, examination of the spicules (~100–200 x 1–2 µm) at extreme magnification (X8000) on the SEM, revealed them to have minute thorn-like scales. It is concluded these are non-native and perhaps were snagged from an hexactinellid glass sponge. one of the specimens (QM G339391) has a much larger range of sizes of chelae (almost two separate size classes of chelae), with a much more robust spine. This specimen was badly damaged during the collection. It may well be a different species, but until there is the collection of another complete specimen, it is retained here as Abyssocladia cf. jeanvaceleti sp. nov. Both this species and the uncertain specimen (cf. QM G339391) are very different to the umbrella-shaped sponge Abyssocladia bruuni Lévi, 1964 (p.78, Fig 30), with respect to the sponge size and the styles, and the isochelae are 70–75 µm long with the single tooth 30 µm wide, and are of a crab-like shape. The sigmancistras in Lévi’s species are twice as large, being 29–30 µm in length. Abyssocladia bruuni sensu koltun, 1970, illustrated in his Fig. 17 (1– 5) and Plate III, Fig. 5, displays a similar shape of chelae and a similar range in sizes of chelae, a sigmancistra and as well as a large style and a much smaller subtylostyle. however, it differs from this new species (including the cf. specimen QM G339391) mainly by the differences in shape of the sponge body. koltun’s specimen came from the Vityaz collections, collected in the South Pacific ocean (Station 3655, Bougainville Trench). Abyssocladia bruuni Lévi, 1964 and Abyssocladia bruuni sensu koltun, 1970 are clearly unrelated to each other. This is also supported by the redescription of Abyssocladia bruuni by Vacelet (2020). we propose koltun’s species Abyssocladia bruuni to be renamed Abyssocladia vladimirii nom. nov. The alternate name Abyssocladia koltuni (Ereskovsky & willenz, 2007) is preoccupied. Neocladia flabelliformis koltun, 1970 is accepted as Koltunicladia flabelliformis (koltun, 1970) (type by original designation). Genus Asbestopluma Topsent, 1901 [Cometella] Schmidt, 1870:49 (nomen oblitum) [Asbestopluma] Lankester, 1882: 478 (nomen nudum) Asbestopluma Topsent, 1901b: 23 Helophloeina Topsent, 1929: 8 Not Lycopodina Lundbeck, 1905: 58 Cotylina Lundbeck, 1905: 68. Diagnosis. Cladorhizidae with at least one small category of palmate isochelae, in a few cases modified to anchorate unguiferate, anisochelae. usually with a second larger type of palmate to arcuate anisochelae that may in some cases be modified to isochelae, anisoplacochela or tridentate anchorate chela. Sigmancistras and basal acantho(sub)(tylo)styles are also present with a few exceptions. Never forceps spicules (from hestetun et al. 2016b). Type species. Cladorhiza pennatula (Schmidt, 1875) (by subsequent designation, Topsent, 1901b). 452 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr Subgenus Asbestopluma Topsent, 1901b Diagnosis. Asbestopluma without spear-shaped microtylostyles (from Lopes et al. 2011). Type species: Cladorhiza pennatula (Schmidt, 1875) (by subsequent designation, Topsent 1901b). Remarks. The genus Asbestopluma has been recently summarised, including remeasuring the spicule dimensions of some old types, by Lopez et al. (2011) and Goodwin et al. (2017), with these data updated to include additional information, and more recently described species, in Table 4 of Ekins et al. (2020a). There have been three species recorded from the Australian EEZ, viz. Asbestopluma (Asbestopluma) desmophora kelly & Vacelet, 2011 from Seamounts on the Macquarie ridge, SE Pacific, As. (As.) obae koltun, 1964 from wilkes Land, Antarctica and As. (As.) maxisigma Ekins et al., 2020a previously known from off Jervis Bay, New South wales, and redescribed here from new material off the Great Barrier reef. Asbestopluma (Asbestopluma) maxisigma Ekins, Erpenbeck & Hooper, 2020 Figures 8–9, Table 4 Material Examined: holotype of Asbestopluma (Asbestopluma) maxisigma: QM G337488 off Jervis Bay, Station 56, New South wales, Australia, -35.333003, -35.332000, 151.258000 – 151.214000, 2636-2342 m, Beam Trawl, Coll. Merrick Ekins on RV Investigator, Cruise IN2017_V03, Sample 56-236, 29/v/2017. QM G339376, Noddy reef, Great Barrier reef, Queensland, Australia, -13.51773023, 144.1015761, 815.495 m, Site: So398, Sample: 130, roV SuBastian, Coll. Martie McNeil on RV FALKOR, cruise Fk200930, 15/X/2020. Comparative Material: Asbestopluma (Abestopluma) biserialis (ridley & Dendy, 1886): NhMuk 1887.5.2.187, South Pacific -22.35, -150.283, 4361 m, Challenger St. 281, 6/X/1875 Lectotype; NhMuk 1887.5.2.190, South Pacific -39.2167, -118.81.67, 4115 m, Challenger St. 291, Paralectotype. Distribution: This species is known from continental slope Queensland and New South wales, Australia, at bathyal depth. Description: Growth form: The new specimen (QM G339376) consists of an erect columnar pedunculate sponge with pinnate filaments projecting at right angles to the stem (Figs. 8 E, F). This specimen is 135 mm long, up to 2 mm wide. Ninety mm of this length comprises the below ground stem and basal root, whilst only 45 mm projects above the substrate, the top part of the specimen was lost during processing. The filaments of the preserved specimen are up to 7 mm in length, but possibly longer in vivo (Figs. 8 A–D). They are 0.5 mm in width and project into four columns, with a right angle between them. Colour: The above ground parts are white in situ and on the deck, whilst the below ground components are tan coloured. Ectosomal skeleton: The ectosome of both the stem and the filaments consist of soft tissue encrusted with anisochelae and sigmas. The ectosome of the lower stem and roots is encrusted with the acanthostyles. Endosomal skeleton: The axis of the stem and the filaments consists of tightly bound longitudinal tracts of mycalostyles. The mycalostyles are also arranged as buttresses providing support for the filaments that are also composed of the same styles and arise tangential to the stem, so that at their ends the mycalostyles converge onto the filament mycalostyles. In addition, there are supplementary smaller very fine and short filament columns composed of the subtylostyles projecting at right angles to the stem and similarly converging with buttressing mycalostyles. The subtylostyles are not present in the stem nor the roots, which is composed of the mycalostyles only. Megascleres: Styles of three types exist in different parts of the sponge. Large mycalostyles, thickest at the middle of the spicule and tapering at both ends (531–(730)–897 µm x 12.5–(20.3)–29.9 µm, n=59) occur in the filaments, the stem and the basal root like appendage (Figs. 9 C, D). Smaller subtylostyles with slightly swollen bases and tapering to fine points (393–(509)–604 x 4.2–(9.8)–12.6 µm, n=22) occur in the filaments (Figs. 9 E, F). Very thin long, often subtly bent fragile acanthostyles occur in the lower stem and root like appendage (66.8– (119.1)–246.0) x 0.8–(2.8)–6.6 µm, n=44) (Figs. 9 G, h). Microscleres: Palmate anisochelae (Fig. 9 A)., head with the lateral alae fully fused to the shaft and a large frontal alae significantly detached from lateral alae, foot with two fully fused nearly atrophied lateral alae and a single larger frontal ala with a tooth-like termination (Length 10.3–(12.8)–14.1 µm, large frontal alae width 3.2–(4.4)–6.5 NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 453 µm, small lateral alae width 1.8–(2.2)–2.7 µm, n=24) Small sigmancistras (Fig. 9 B), with an almost 90o twist (28.2–(32.3)–36.0 µm, n=20) were found. The larger sigmas recorded in the holotype by Ekins et al. (2020), were not found in this specimen. It is possible these may have been contaminants from Chondrocladia (Chondrocladia) clavata ridley & Dendy, 1886, multiple samples that were collected from the same beam trawl station. FIGURE 8. Asbestopluma (Asbestopluma) maxisigma Ekins et al. 2020a. A. QM G339376 underwater. B–D. Close up of QM G339376 underwater as being harvested by the roV SuBastian. E. QM G339376 fixed specimen. F. Close up of the stem and filaments. G. Scanning electron microscope of the stem showing the radial filaments. h. Ectosomal layer of the stem showing the anisochelae and sigmas, with the underlying styles. I. Filament emerging from the stem heavily encrusted with anisochelae and sigmas. 454 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr FIGURE 9. Asbestopluma (Asbestopluma) maxisigma Ekins et al., 2020. A. Anisochelae. B. Sigmas. C. Large mycalostyle. D. Magnified ends of the mycalostyle depicted in C. E. Smaller subtylostyle. F. Magnified ends of the subtylostyle depicted in F. G. Acanthostyles. h. Magnified ends of the acanthostyles in G. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 455 456 · Zootaxa 5293 (3) © 2023 Magnolia Press TABLE 4. Comparison of branched pinnate Absestopluma (Asbestopluma) species. Spicule dimensions measurements smallest–(average)–largest (µm) Species reference As.(As.) maxisigma Ekins et al. 2020a: 28–46, Fig. 6, Table 4 As.(As.) maxisigma This study As. (As.) biserialis Size (mm) Filament rows Mycalostyles Subtylostyles 4 855–(1000)–1130 x 16–(21)–28 427–(586)–805 x 4–(9)–15 135 4 531–(730)–897 x 12–(20)–30 393–(509)–604 x 4–(10)–13 (ridley & Dendy, 1886: 40);(ridley & Dendy, 1887: 75–76, Pl. XIV, Figs. 2–3, Pl. XV Figs. 8, 8a–b.) 92 x 1 2 2000 x 38 440 x 10 As. (As.) biserialis Lévi, 1964: 74–75, Fig. 21, Pl. III, Fig. h 120 x 1 2 1500–1900 x 18–20 200–500 x 2–3 As. (As.) biserialis koltun, 1970: 188–190, Fig. 15, Pl. III, Fig. 1 140 x 2 2 2000 x 18–38 400–1300 x 13–25 As. (As.) biserialis Lévi, 1993: 39, Fig. 14 B 95 x 1 2 1600–1800 x 20–21 375–800 x 10–14 As. (As.) biserialis var. californiana de Laubenfels, 1935: 4, Fig. 5 200 x 14 2 1150 x 18 As. (As.) belgicae (Topsent, 1901:VIII) 200 6 1400 x 23 800 x 12–15 As. (As.) belgicae hestetun et al. 2015:1316–1318 Table 1, Fig. 4 A–J 350 x 2 8 1010–(1449)–2070 x 17–(24)–31 610–(734)–880 x 11–(15)–19 As. (As.) belgicae Goodwin et al. 2017: 48 As. (As.) quadriserialis Tendal, 1973: 34–35, Figs. 1–2 75 x 1 4 480–(876)–960 x 6–25, filaments 880–(968)–1040 x 12–25 400–(511)–640 x 5–10 ......continued on the next page EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 4. (Continued) Species Strongyles Acanthostyles Anisochelae Sigmas Location Depth (m) 11–(13)–16 I: 20–(28)–38 II: 43–(54)–69 NSw, Australia 2600 10–(13)–14 28–(32)–36 Queensland, Australia 816 As. (As.) biserialis 12.6 25 South Pacific 4364 As. (As.) biserialis 9–10 25–27 kermadec Trench, South Pacific 2640 9–13 25–27 Nw Pacific 2640-6282 As. (As.) biserialis 8–10 26–28 New Caledonia 1618-1740 As. (As.) biserialis var. californiana 6 22 California As.(As.) maxisigma 67–(119)–246 x 1–(3)–6 As.(As.) maxisigma 100–500 x 1–3 As. (As.) biserialis As. (As.) belgicae 1400 x 23 200 x 3 12 33 Antarctica 400-569 As. (As.) belgicae 460–(659)–830 x 3–(6)–9 127–(198)–250 x 1.5 11–(13)–14 27–31–35 Central Atlantic 3147-3898 As. (As.) belgicae 820–(1105)–1384 x 13–(24)–36 116–232 x 3–4 8–(10)–12 28–(30)–32 75–(106)–125 x 1–3 I: 35–45 II: 10–12–13 I: 28–(31)–38 II: 75–(106)–125 North Atlantic 4600 As. (As.) quadriserialis Zootaxa 5293 (3) © 2023 Magnolia Press · 457 Remarks: As previously noted in Ekins et al. (2020a), As. (As.) maxisigma is most closely related to As. (As.) biserialis (ridley & Dendy, 1886), known from the South Pacific (ridley & Dendy 1886), kermadec Trench, (Lévi 1964), Coral Sea off New Caledonia (Lévi 1993), and the North Pacific, south of the Aleutian Islands (koltun 1970), from bathyal and abyssal depths (qv. Lopez et al. 2011; Ekins et al. 2020a). Both species have vaguely similar pinnate pedunculate morphologies but As. (As.) maxisigma has twice as many columns of filaments (i.e., four as opposed to two) and is round in cross-section rather than flattened (Fig. 10). This re-description of As. (As.) maxisigma with new material collected by roV, clearly shows that only about one third of the sponge is above the ground i.e., the ‘snow’ layer (Figs. 8A, E, F). The inclusion of the below-ground stem and the root like structures of QM G339376 from the Great Barrier reef shows the presence of acanthostyles, which were not found on the holotype of As. (As.) maxisigma. Since this species is only known to live in soft sediment, it is likely the spined surface of the acanthostyles serve as anchors securing the sponge in the soft sediment. Another example of exaptation of spicules in carnivorous sponges. These acanthostyles are similar to those found in As. (As.) biserialis but were only reported in Lévi (1964) and koltun (1970), not in the original description by ridley & Dendy (1886) nor Lévi (1993). with the inclusion of new acanthostyles this species is even more closely aligned with As. (As.) biserialis (ridley & Dendy, 1886). however, the species are kept separate as this distinction between the clearly biserial species As. (As.) biserialis, is also this distinction between it and another closely related species As. (As.) belgicae (Topsent, 1901a), which has 6–8 rows of filaments. Asbestopluma (As.) biserialis var. californiana de Laubenfels, 1935 is currently only distinguished by having slightly smaller chelae from a described single value of 6 µm, as opposed to the current species which begin at 10 µm. re-examination of the type material of As. (As.) biserialis var. californiana, may reveal a greater size range of these microscleres. FIGURE 10. Asbestopluma (Asbestopluma) biserialis (ridley & Dendy, 1886). A. Large fragment of Lectotype NhMuk 1887.5.2.187. B. Two fragments of NhMuk 1887.5.2.187 magnified showing flatted stem and biserial arrangement of the filaments. C. Paralectotype NhMuk 1887.5.2.190. 458 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr In its gross morphology this species also resembles As. (As.) belgicae, (qv. Lopez et al. 2011, hestetun et al. 2015; Goodwin et al. 2017). Although, As. (As.) maxisigma has acanthostyles, it has the following differences: fewer radial filaments, the absence of grooves, smaller mycalostyles, an absence of the strongyles. Asbestopluma (Asbestopluma) quadriserialis Tendal, 1973, from the North Atlantic also has four rows of filaments, however, it can be clearly distinguished by the presence of two sizes of anisochelae. These closely related species are compared in Table 4. Asbestopluma (As.) sarsensis Goodwin et al., 2017 is similar in spiculation to As. (As.) belgicae, and also differs from the present species for the same reasons given above, in addition to also having a very different growth form. Asbestopluma (Asbestopluma) obae koltun, 1964 from wilkes Land, Antarctica differs from the present species in lacking horizontal filaments. As shown in Fig. 3 of Ekins et al. (2020a), As. (As.) maxisigma does not have the same molecular sequence as any other Asbestopluma (Asbestopluma) spp. unfortunately, none of the closely related species listed above have any molecular sequence data either. The only one of the closest morphologically is As. (As.) cf. belgicae, which has a different molecular profile. As new collections of species of this genus and their corresponding sequences become available, will result in better resolution within the genus. Genus Axoniderma Ridley & Dendy, 1886 Type species: Axoniderma mirabilis (ridley & Dendy, 1886) (type by monotypy) Definition: Cladorhizidae with anchorate anisochelae and a ‘crinorhizoid’ parasol morphology. Species also included: Axoniderma mirabilis (ridley & Dendy, 1886), A. australis (Ekins, Erpenbeck & hooper, 2020), A. corona (Lehnert, watling & Stone, 2005), A. hubbsi (Lundsten, reiswig & Austin, 2017), A. kensmithi (Lundsten, reiswig & Austin, 2017), A. longipinna (ridley & Dendy, 1886), A. mexicana (Lundsten, reiswig & Austin, 2017), A. poritea (Ekins, Erpenbeck & hooper, 2020) and A. similis (ridley & Dendy, 1886). Remarks. The type species Axoniderma mirabilis was originally described as Trochoderma mirabile ridley & Dendy, 1886, type species by monotypy of Trochoderma ridley & Dendy, 1886 (ridley & Dendy, 1886: 344). however, the genus was already occupied and another genus Axoniderma ridley & Dendy, 1886 was created later, in the same publication (ridley & Dendy, 1886: 493) to accommodate the one species. The full description of Axoniderma mirabile was completed later (ridley & Dendy 1887: 96–98). The genus was later synonymized with Cladorhiza (hajdu & Vacelet, 2002). All of the species previously encompassed within the genus Cladorhiza were compared (Ekins et al. 2020a), and then subsequently split (Ekins et al. 2020b) into five genera: Cladorhiza Sars, 1872 (with arbuscular morphology), Axoniderma ridley & Dendy, 1886 (with classic parasol/umbrella morphology), Abyssosdiskos Ekins, Erpenbeck, Goudie & hooper, 2020 (with an upwards facing disc), Bathytentacular Ekins, Erpenbeck, Goudie & hooper, 2020 (with teardrop with tentacular processes) and Nullarbora Ekins, Erpenbeck, Goudie & hooper, 2020 (with pinnate morphology). Axoniderma wanda sp. nov. Figures 11–12, Tables 5–6 urn:lsid:zoobank.org:act:819293E6-E6BB-4032-B672-493676A91040 Material examined: holotype: QM G339384, wreck Plunge Pool, Great Barrier reef, Queensland, Australia, -12.131851, 143.979498, 1980.05 m, hanging upside down in a small cave, Site: So402, Sample: 178, roV SuBastian, Coll. Merrick Ekins remotely in Brisbane via video live satellite feed directing the roV pilots onboard the RV FALKOR on the Great Barrier reef and Martie McNeil onboard the RV FALKOR, cruise Fk200930, 27/10/2020. Paratypes: QM G339392, QM G339746, QM G339747, QM G339748, QM G339749—all paratypes have the same collection details as the holotype QM G339384. Etymology.This species is named for its resemblance to the alien wanderer/wanda that parasitizes the host Melanie Stryder, played by actress Saoirse ronan from the movie and novel by the same name i.e. “the host”, authored by Stephenie Meyer. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 459 Distribution. This species is presently known only from the bathyal zone of the Great Barrier reef, East Coast of Australia. Description: Growth form: The holotype consists of a pedunculated sponge, with a long thin stem that radiates at attachment in the centre of the posterior face of the sponge body. The body is an almost vertically orientated disc, concave on the anterior face, with up to 42 long thin filaments radiating out in a single plane from the disc margin in a complete circle (Figs. 11 A–G). From the top of the disc arises a thin stalk, longer and slightly thicker than all other filaments, terminating in a ‘lure’ in the shape of a minute circular fan (Fig. 11 J). The sponge body diameter is 8 mm, and 0.4 mm thick. The filaments are 23 mm long and 0.1 mm in width. The lure filament is 24 mm in length and 0.3 mm in diameter, whilst the circular lure is 1 mm in diameter. Interestingly, several specimens such as paratype G339747 has a raised bump on the stem which is most likely an oocyte-embryo swelling. Colour: The sponge body, filaments and stem are all off white, with a tan coloured lure. Ectosomal skeleton: The ectosomal skeleton, on the body, the stem and basal attachment and the filaments are covered on the external surface with tridentate ‘unguiferate’ anisochelae (Figs. 11 h–L). The lure and the filament supporting the lure are, however, covered with tridentate ‘unguiferate’ anisochelae as well as sigmancistras (Fig. 11 J). Endosomal skeleton: The endosomal skeleton of the body, the filaments, the stem and the basal attachment consist mainly of the larger mycalostyles in concentrated longitudinal bundles (Figs. 11 G–J, L), but also include the less common medium sized mycalostyles. The large mycalostyles originate in the centre of the body and radiate out to become the horizontal filaments. The stem and basal attachment and rarely the filaments also contain strongyles. The lure is composed of the medium sized mycalostyles as well as the thin ‘Lure Styles’ (Fig. 11 k). The filament for the lure is composed of the large and medium sized mycalostyles. Megascleres: The megascleres mainly consist of two different sized but overlapping mycalostyles with blunt terminations, with the largest diameter off-centre of the spicule (Figs. 12 C–F). The lure styles are long very thin and sinuous with no central thickening (Figs. 12 G, h). The strongyles are tapering with an offset thickening and blunt tips (Figs. 12 I, J). Dimensions are given in Table 5. Microscleres: The microscleres consist of abundant small anchorate ‘unguiferate’anisochelae with three large alae and five smaller alae on each end. The two outer small lower alae are paired, whilst the central lower small alae is solitary (Figs. 12 A, B). Sigmancistras are slightly contort, consist of a single size class and are restricted to the lure (Fig. 12 C, Table 5). Remarks. This species has the ‘unguiferate’ anchorate anisochelae and sigmancistras common to many other species of Cladorhiza, Axoniderma, Bathytentacular and Nullarbora. Axoniderma wanda sp. nov. is unique amongst the known species of these genera in having an external lure originating from the convex face of the body. This species is compared to other described species with any form of structure emerging from the body disk in Table 6. All species previously and currently belonging to Cladorhiza were also compared in Table 6 of Ekins et al. (2020a), and grouped morphologically by a visual icon in (Table 6, Ekins et al. 2020b). The new species Axoniderma wanda sp. nov., has a similar sclerite composition and basic morphological shape to Axoniderma corona (Lehnert, watling & Stone, 2005), Axoniderma kensmithi (Lundsten, reiswig & Austin, 2017) and Axoniderma mexicana (Lundsten, reiswig & Austin, 2017). It differs from all three by orders of magnitude of size, and also has a very different lure shape and size from those three species. Axoniderma longipinna (ridley & Dendy, 1886) is the only other sponge with a disc and some disc-like structures, but it displays the classic ‘crinorhizoid’ parasol morphology and is not related to the new species. Axoniderma corona, from the Northern Pacific ocean is the closest related species with the presence of anisooxeas similar to the strongyles in Axoniderma wanda sp. nov. and the sub(tylostyles) from the crown similar to the styles in the lure of Axoniderma wanda sp. nov. and the mycalostyles are of an overlapping size range. however, the much larger (by orders of magnitude) in size, the very short stem of the crown, the difference in crown versus lure shape, and the different shape of the lower alae in the larger anisochelae preclude A. corona from being the same as the new species. Axoniderma kensmithi differs from Axoniderma wanda sp. nov. by the much larger (by orders of magnitude) size, the convex downward facing filaments, the appearance of four lures on the shorter stems, the larger anisochelae and sigmancistras, the shape of the styles, and the shape of the lower alae of the anisochelae. Axoniderma mexicana differs from Axoniderma wanda sp. nov. by the much larger (by orders of magnitude) size, convex body shape, the double layer of filaments, the lack of strongyles and small styles, the presence of pseudoamphiasters and the larger sigmancistras, and the different lower alae of the anisochelae. 460 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr FIGURE 11. Axoniderma wanda sp. nov. A. underwater photo of the types in situ. B. QM G339747. C. QM G339746. D. QM G339749. E. holotype QM G339384. F. QM G339748. G. QM G339746, showing an amphipod caught on a filament and a semidigested crustacean in the sponge body. h. Filaments emerging from the sponge body. I. A filament showing the encrusting chelae. J. The lure of the sponge. k. The stem of the sponge. L. The basal attachment. NEw CArNIVorouS SPoNGES FroM AuSTrALIA Zootaxa 5293 (3) © 2023 Magnolia Press · 461 FIGURE 12. Axoniderma wanda sp. nov. holotype QM G339384. A. Tridentate ‘unguiferate’ anisochelae. B. Sigmancistra. C. Large mycalostyles. D. Magnified ends of the mycalostyle illustrated in C. E. Medium sized mycalostyle. F. Magnified ends of the mycalostyle illustrated in E. G. Lure style. h. Magnified ends of the lure style illustrated in G. I. Strongyle. J. Magnified ends of the strongyle illustrated in I. 462 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 5. Spicule dimensions measurements smallest–(average)–largest (µm) of the holotype and paratypes of Axoniderma wanda sp. nov. NM=Not Measured Specimen Large Mycalostyles Medium Mycalotyles Lure Styles Strongyles Anisochelae Sigmancistras QM G339384 holotype 903–(1619)–2130 x 23.8–(35.5)–56.5, n=28 573–(727)–998 x 12.4– (14.7)–20.2, n=19 262–(362)–452 x 5.0–(6.5)–11.0, n=11 370–(547)–657 x 20.0–(23.0)–26.7, n=4 22.3–(26.4)–28.3 x 1.5–(2.6)–3.9, n=33 24.0–(28.4)–33.4 x 1.3–(2.1)–3.6, n=22 QM G339392 961–(1730)–2360 x 20.0–(38.6)–56.5, n=44 420–(762)–1180 x 10.5–(17.1)–24.1, n=23 377–(455)–622 x 2.8–(6.0)–9.1, n=28 326–(573)–955 x 15.3–(23.3)–36.8, n=34 25.0–(26.7)–29.0 x 1.3–(2.4)–3.2, n=31 20.8–(29.2)–36.1 x 0.9–(1.7)–3.1, n=37 QM G339746 1040–(1630)–1920 x 16.1–(29.0)–40.5, n=5 NM NM 486 x 22.7, n=1 23.8–(27.3)–29.5 x 1.6–(2.7)–3.6, n=33 NM QM G339747 1310–(1679)–2030 x 22.6–(33.4)–44.6, n=36 569–(877)–1160 x 6.1–(15.1)–23.8, n=28 NM 635 x 22, n=1 22.9–(25.9)–28.4 x 1.6–(2.4)–4.1, n=36 NM QM G339748 1050–(1401)–1800 x 19.8–(28.4)–38.8, n=8 673–(685)–697 x 16.6– (16.6)–16.6, n=2 NM 283–(717)–1150 x 9.8–(23.8)–37.8, n=2 23.4–(26.8)–29.2 x 1.6–(2.5)–3.1, n=19 26.4 x 0.6, n=1 QM G339749 1030–(1430)–2040 x 17.1–(26.3)–35.7, n=17 529–(772)–1090 x 7.0–(14.7)–19.0, n=19 NM 642–(651)–660 x 23.5–(25.6)–27.7, n=2 22.3–(26.5)–29.0 x 1.6–(2.6)–3.4, n=18 NM Zootaxa 5293 (3) © 2023 Magnolia Press · 463 464 · Zootaxa 5293 (3) © 2023 Magnolia Press TABLE 6. Axoniderma wanda sp. nov. compared to other species of Axoniderma containing structures emerging from the body disc. Spicule dimensions measurements smallest–largest (µm). Species Source Morphology Total height x stem width (mm) Skeleton Axoniderma wanda sp. nov. This study Pedunculate erect, unbranched long thin stem with a disc shaped body with radial filaments and on a long stalk a circular terminal lure 60 x 0.5 Axis of stem and body cored by longitudinal bundles of mycalostyles of 2 size classes and an extra thin small style located in the lure Axoniderma corona (Lehnert, watling & Stone, 2005) Lehnert, watling & Stone, 2005: 1359, Figs. 1–2 Pedunculate erect, parasol ‘crinorhizoid’ morphology, long stem, two planes of different appendages, basal disk holdfast 225–325 x 1–9 Axis of stem and basal appendages with thick bundles of long mycalostyles, crown with tracts of long mycalostyles fanning out in 1 plane, with single thin subtylostyles perpendicular to the axial skeleton, basal plate with mycalostyles and short anisoxeas densely packed in one plane parallel to the substrate Axoniderma kensmithi (Lundsten, reiswig & Austin, 2017) Lundsten, reiswig & Austin, 2017: 250–253, Figs. 2–3 Crinorhizoid’ form with numerous filaments parasol-shaped, on long stem with densely branching basal rhizoid holdfast, 2–4 spermatocyst-bearing discs on short, slender stalks on apex of body 208–325 x 1–2.1 Axis of stem cored by mycalostyles 1–3, filaments cored by mycalostyles 1&3, basal rhizoid cored by strongyles and mycalostyles 3 Axoniderma longipinna (ridley & Dendy, 1886) ridley & Dendy, 1887: 92, P1. XX. Fig. 2; P1. XXI. Figs. 4, 21; koltun 1970: 185–186, Fig. 13, PL. VI Figs. 3–5 Pedunculate erect, ‘crinorhizoid’ parasol morphology, long thin stem with apical suspherical body bearing long radial projections outwards and downwards, basal attachment not recorded 27 x 5 Axes of stem and lateral processes cored by bundles of long mycalostyles Axoniderma mexicana (Lundsten, reiswig & Austin, 2017) Lundsten, reiswig & Austin, 2017: 253, Figs. 4–5 Erect parasol-shaped, ‘crinorhizoid’ body with radiating long filaments at base of body, on long stalk, presumed basal holdfast not collected 300 x 1–3 3 size classes of mycalostyles, presumed undifferentiated distribution in sponge EkINS & hooPEr ......continued on the next page NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 6. (Continued) Species Source Morphology Total height x stem width (mm) Skeleton Axoniderma mirabilis ridley & Dendy, 1886 ridley & Dendy, 1887: 98, Pl. XX, Fig. 5, Pl. XXI, Figs. 8–10; (Ekins et al. 2020a); [koltun 1970: 187, Fig. 13] Pedunculate erect, ‘crinorhizoid’ parasol-shaped with conical cap-shaped body with terminal papillae, perched on the end of a long slender stem, with numerous long filaments surrounding the body, basal attachment not recorded 56 ? (13 x 2) Axis of stem, body and filaments cored by bundles of styles-mycalostyles of a single size class, with single styles projecting on the terminal papillae, cortex encrusted with pseudoamphiasters and other microscleres Axoniderma similis (ridley & Dendy, 1886) ridley & Dendy, 1886: 343; ridley & Dendy, 1887: 93, Pl. XX, Fig. 7, Pl. XXI, Fig. 5, 18; (Lévi, 1993: 40, Fig. 40. Pl. IV, Figs. 4–6) Pedunculate erect, ‘chrinorhizoid’ parasol morphology, with cap-shaped conical body and long filaments encircling the lower body, perched on a thin stem, basal attachment missing ? Axis of stem, body and filaments cored by bundles of styles, dense close-set tylostyles mostly near surface, fewer in the axial skeleton ......continued on the next page Zootaxa 5293 (3) © 2023 Magnolia Press · 465 466 · Zootaxa 5293 (3) © 2023 Magnolia Press TABLE 6. (Continued) Species Spicules of main axis (Lxw um) Spicules of lateral filaments or body (Lxw um) Spicules of basal attachment (Lxw um) Chelae (L um) Sigmancistras (L um) Sigmas (L um) and other microscleres Locality/ depth range Axoniderma wanda sp. nov. mycalostyles 1, 903–2360 x 24–56 mycalostyles 2, 420–1090 x 7–24 Lure styles 260–320 x 3–11 Strongyles 280– 1150 x 15–38 tridentate unguiferate anisochelae 22–29 x 1–3 sigmancistras 20–36 x 1–4 absent Great Barrier reef, Australia, bathyal Axoniderma corona (Lehnert, watling & Stone, 2005) mycalostyles 600–4260 x 10–65 (sub–)tylostyles 510–1650 x 8–20 (in the crown) anisoxeas 140–660 x 38–43 anchorate anisochelae 30-42 35-42 absent Aleutian Islands, mesophoticbathyal Axoniderma kensmithi (Lundsten, reiswig & Austin, 2017) mycalostyles 1, 3138–4850 x 49–75 mycalostyles 2, 1009–1501 x 15–29 mycalostyles 3, 242–452 x 13–17 mycalostyles 1 &3 strongyles 421– 1023 x 10–22 mycalostyles 3 tridentate unguiferate anisochelae 33–37 43–50 absent west Coast, uSA, bathyalabyssal Axoniderma longipinna (ridley & Dendy, 1886) mycalostyles 1, 3000 x 50–55 mycalostyles 2, 200–1000 x 10–16 undifferentiated not recorded unguiferate tridentate anisochelae 30–78 absent absent Nw Pacific, bathyal - hadal Axoniderma mexicana (Lundsten, reiswig & Austin, 2017) mycalostyles 1, 3577–4133 x 58–74 mycalostyles 2, 2650–3246 x 45–63 mycalostyles 3, 1293–1931 x 23–39 undifferentiated undifferentiated tridentate unguiferate anisochelae 33–36 47–55 asymmetrical pseudoamphiasters 102–115 west coast uSA, Mexico, bathyal ......continued on the next page EkINS & hooPEr NEw CArNIVorouS SPoNGES FroM AuSTrALIA TABLE 6. (Continued) Zootaxa 5293 (3) © 2023 Magnolia Press · Species Spicules of main axis (Lxw um) Spicules of lateral filaments or body (Lxw um) Spicules of basal attachment (Lxw um) Chelae (L um) Sigmancistras (L um) Sigmas (L um) and other microscleres Locality/ depth range Axoniderma mirabilis ridley & Dendy, 1886 styles >3500 (styles– mycalostyles 1, ~6000 x 11 styles 2, ~300 x 5) [mycalostyles ~3500 x 15–50] undifferentiated absent tridentate unguiferate anisochelae 38 (31–48) [‘anchors’ 33–38] absent sigmas 75.6 (88–114) [sigmas 76] birotules (‘amphiasters’) with 5 terminal alae 230 (pseudoamphiasters 127–295) [‘amphiasters’ 88– 230] SE Pacific, off S Easter Island, & N Pacific, abyssal Axoniderma similis (ridley & Dendy, 1886) styles ‘very long and slender’ tylostyles 210–600 x 16 (styles 1, up to 3600 x 50 styles 2, 650–700 x 30 styles 3, 250–300 x 8–10) undifferentiated undifferentiated tridentate unguiferous anisochelae 30 (anisochelae 30) absent absent Central Pacific & New Caledonia, bathyalabyssal 467 All of the specimens of Axoniderma wanda sp. nov. were collected from the underside of an overhang (Fig. 11 A), and so live in an inverted position (i.e. hanging upside down), such that the concave body disc actually faces downwards, ensuring the disc does not capture the falling debris ‘snow’, as does the habitat of taking shelter under overhangs. Not all spicules on each paratype were measured, as it was considered that enough of these fragile tiny specimens were already destroyed in order to observe part of the structure under an SEM, and so ‘NM’ (i.e. Not Measured) in Table 5 is a reflection of this, rather than an indication that the spicule type was absent from the specimen. Discussion The discovery of five species of carnivorous sponge in the present study, of which three new species were discovered from only four roV dives highlights the potentially much higher diversity of cladorhizid sponges at sub-mesophotic depths off the Great Barrier reef. up until the present study roVs and their skilled operators have rarely been deployed in this region, but the present carnivorous sponge discoveries demonstrate their immense value to carefully observe and collect from cryptic habitats such as overhangs and caves at depths of one kilometre or more, compared to more destructive sampling methodologies such as bottom trawls from which most of the carnivorous sponges around the world have been discovered. Together with scanning electron microscopy allowing detailed spicular morphology as well as molecular techniques that are now teasing apart the relationships between species and genera our knowledge of cladorhizids has escalated over the past few decades. Although there are some old records, the recent redescriptions of old materials (e.g hestetun et al. 2017) have fixed most of these, but this group of sponges lacks the impediments that plagues most modern sponge taxonomy (i.e. having to revise every genus in order to describe new species). The recollection of species only previously described from trawled specimens by roV, enabled the collection of the entire specimen. This enabled a fuller description of two previously described species i.e. (Chondrocladia (Chondrocladia) zygainadentonis Ekins et al., 2020a and Asbestopluma (Asbestopluma) maxisigma Ekins et al., 2020a) previously known from the East coast of Australia based on incomplete specimens. The holotype of Asbestopluma (Asbestopluma) maxisigma i.e. QM G337488, is the only current specimen of this species that has a molecular sequence, so future collections of more species of this genus with molecular analysis would be useful for confirmation of the pinnate species throughout the world’s oceans. Despite the skill of the roV operators in collecting and photographing intact specimens in situ, it was not possible to do molecular analysis of the new species due to contamination problems that occurred either during the collection or most likely during the processing of samples on the vessel. The robust morphological and spicular characteristics of these carnivorous sponges easily enable the classification using traditional sponge taxonomy. The examination of the new species Axoniderma wanda sp. nov. and its placement within the genus Axoniderma, has revealed one species Bathytentacular moniqueae (Ekins et al., 2020a), to be closely related to Cladorhiza pentacrinus Dendy, 1887. They both have similar sized and shaped anisochelae, mycalostyles and sigmas, however C. pentacrinus lacks the sigmancistras present in B. moniqueae. Cladorhiza pentacrinus was also collected from the southwest Pacific i.e. northeast of New Zealand at a shallower depth of 1280 m as opposed to 4200 m off the East Coast of Australia of C. moniqueae. Cladorhiza pentacrinus and C. bathycrinoides koltun, 1955 should both be moved to the genus Bathytentacular. Acknowledgements The collection of material was supported by a grant of sea time on rV Falkor from the Schmidt ocean Institute. we would like to thank the Master, ship’s crew and scientific staff, especially the pilots of the roV SuBastian who all helped with the collection of specimens. we would also like to thank rob Beaman, Mardi McNeil, Jeremy horowitz and Geosciences Australia for collection, trip organisation, site data and the bathymetric map. These samples were collected under Australian Marine Park Activity Permit PA2020-00040-1, Great Barrier reef Marine Park Authority permit G20/43974.1 V. 2 and Parks Australia Access to Biological resources in a Commonwealth Area for Non-Commercial Purposes Au-CoM2020-483. ME would like to thank Geoff Thompson for his help with 468 · Zootaxa 5293 (3) © 2023 Magnolia Press EkINS & hooPEr the AI sharpening of the underwater image in Fig. 4A and Tom white from the NhM for his photographs of the type material of Asbestopluma (Asbestopluma) biserialis. References de Voogd, N.J., Alvarez, B., Boury-Esnault, N., Carballo, J.L., Cárdenas, P., Díaz, M.-C., Dohrmann, M., Downey, r., hajdu, E., hooper, J.N.A., kelly, M., klautau, M., Manconi, r., Morrow, C.C. Pisera, A.B., ríos, P., rützler, k., Schönberg, C., Vacelet, J. & van Soest, r.w.M. (2022) world Porifera Database. Available from: https://www.marinespecies.org/porifera (accessed 12 July 2022) https://doi.org/10.14284/359 Ekins, M., Erpenbeck, D. & hooper, J.N.A. (2020a) Carnivorous sponges from the Australian Bathyal and Abyssal zones collected during the rV Investigator 2017 Expedition. 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