Carnivorous sponges from the Australian Bathyal and Abyssal zones collected during the RV Investigator 2017 Expedition Author Ekins, Merrick Author Erpenbeck, Dirk Author Hooper, John N. A. text Zootaxa 2020 2020-05-12 4774 1 1 159 journal article 22182 10.11646/zootaxa.4774.1.1 269cf599-428d-464d-8f21-994d69110c5c 1175-5326 3825140 B0C4A2F8-F2AB-4147-BB12-63720EEF2516 Abyssocladia escheri sp. nov. Figure 2 , Tables 1 & 2 urn:lsid:zoobank.org:act: 49779C5D-8130-4573-808D-1CAB9F5469CE FIGURE 2. Abyssocladia escheri sp. nov. A. Abyssochelae. B. Mycalostyle. C. Magnified ends of the mycalostyle depicted in B. D. Substrongyles styles from the basal rhizoid rootlets. E. Magnified ends of the substrongyle depicted in D. F. Paratype QM G337506. G. Paratype QM G337507. H. Holotype QM G337545. I. Paratype QM G337546. J. SEM of the filaments showing the abyssochelae in the ectosome and the mycalostyles. K. SEM of the root like processes showing the substrongyles longitudinally arranged. TABLE 1. Comparative morphological and distributional data for all known species of Abyssocladia .

Species	Reference 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)	Sigmancistr as (L μm)	Sigmas Locality, (L μm) depth range and other microscleres
Abyssocladia escheri sp. nov.	this work ‘Pedunculate’, erect, with long very thin stem, multi-furcate root-like basal holdfast processes, and club-shaped to conical body with irregular lateral filaments	305 x 20	‘Pedunculate’, body and filaments supported by longitudinal bundles of longer styles, basal root-like processes with longitudinally aligned shorter style, stem contains both types of styles but mainly the shorter ones	shorter styles 92–555 x 3.4–16.7	longer styles 615–1280 x 7.0–15.2	shorter styles 92–555 x 3.4–16.7	Abyssochelae 18.4–53.1 x 10.2–24.0	absent	absent	off Fraser Island, Queensland, Australia, bathyal
Abyssocladia annae sp. nov.	this work ‘Pedunculate’, erect stem supporting a slightly cupped- shaped obovate (leaf-like, flabellate, fan-shaped) apical body, lacking lagteral filaments	3 x 2	Peduncle axis longitudinally arranged subtylostyles, body with radiating subtylostyles projecting only in one quadrant	subtylostyles 288-(504)-1000 x 3.1-(7.4)-15.2	undifferentiated	absent	Abyssochelae 48.2-(60.6)-72.2 x 5.1-(9.7)-14.9	11.6-(15.6)- 18.5	absent	off the continental shelf of central New South Wales, Australia, abyssal
Abyssocladia gliscofila sp. nov.	this work ‘Pedunculate’, erect, long thin stem, inflated filaments increasingly longer towards the apex, furcate root-like basal holdfast with fine basal rootlets anastomosing from roots	120–130 x 0.15–1.0	Peduncle axis, filaments and upper basal holdfast roots tightly bound longitudinal tracts of mycalostyles, axis of basal rootlets tightly bound longitudinal tracts of styles.	Mycalostyles 319–1560 x 2.0–22.7	undifferentiated	Styles 179–470 x 4–10.2	Arcuate isochelae 22.4–42 x 1.5–7	absent	absent	continental slope adjacent to the central and northern New South Wales coast, Australia, mesophotic -bathyal

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Species	Reference	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)	Sigman cistras (L μm)	Sigmas Locality, (L μm) depth range and other microscleres
Abyssocladia sp. 1	‘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	mycalostyles 1, 1380–3810 x 15–54	undifferentiated	oxeote anisostyles 2, 392–1560 x 20–50	palmate cleistochelae 62–123 x 17–37	21.5–40 x 2-7	oxyasters 74–136	Great Australian Bight, bathyal
Abyssocladia atlantica Lopes & Hajdu, 2014	Lopes & Hajdu, 2014: 331–333 , Fig. 2	Erect, short peduncle, ‘centipedal’ lamellar body, lateral filaments	4–5 x 3.4	Axis of peduncle longitudinal styles to substrongyles, axis of filaments plumose mycalostyles	styles to substrongyles 226-(521.1)-903 x 10-(18.3)-33	mycalostyles 431-(505.6)-575 x 8-(8.9)-10 styles 226-(320.5)-369 x 13-(14.4)-15	Styles to substrongyles 58-(112.2)-193 x 8-(10.4)-15	Cleistochelate isochelae 23-(45.9)-70	absent	absent	Campos Basin, Brazil, bathyal
Abyssocladia boletiphora Hestetun, Rapp & Xavier, 2017	Hestetun et al. , 2017: 169–170 , Fig. 2	‘Bottle-brush’, erect, single- axis, main stem connected to substrate by small basal plate, axis with rows of numerous filaments, side branches terminally inflated	15–30 x 2	Main axis tight bundles of mycalostyles, basal part also with strongyles, spicules in filaments perpendicular between stem bundles, terminal branches scattered styles	mycalostyles 465–(831)–2023 x 17.7–(24.4)–41.3	styles 283–(338)–400 x 4.2–(6.3)–9.8	strongyles 164-(289)-431 x 11.2-(16.8)- 23.9	Arcuate isochelae I, 4.8–(74.4)–91.5 Arcuate isochelae 2, 21.4-(32.4)-51.2 Abyssochelae, 25.7-(42.6)-54.8	8.3-(9.9)- 11.4	absent	Southwestern Indian Ocean Ridge, off South Africa, bathyal

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Species	Reference	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
Abyssocladia bruuni Lévi, 1964	Lévi, 1964: 78 , Fig. 30; Hestetun, Fourt, Vacelet, Boury- Esnault & Rapp, 2015	‘Pedunculate’, stem with a short apical club-shaped umbrella and irregular terminal lateral spicule bundles	13 x 1	Not described	subtylostyles 1300–1650 x 25–28	undifferentiated	base missing	Arcuate isochelae (‘thaumatochetes’) 70–75 x 30	absent	29–30 x 2–3	Kermadec Trench, New Zealand, abyssal
Abyssocladia Kelly & carcharias Vacelet, Kelly & Vacelet, 2011: 2011 58–60, Figs 2–3		‘Pedunculate’, long thin peduncle, 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 skelton, bundles of mycalostyles extending into filaments	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	Sigmancistras 1, 15–16.2 x 1.9–2 sigmancistr as 2, 8–128–12	absent	Kermadec Seamounts, New Zealand, bathyal
Abyssocladia claviformis Koltun, 1970	Koltun, 1970: 191–193 , Fig. 17	‘Pedunculate’	100 x 1.5	Main skeleton of axial type and composed of fusiform styles	Styles 600–1600 x 22–50 tylotes	subtylostyles 1, 175–850 x 5–11 subtylostyles- styles 2, 175–650 x 5	not recorded	arcuate isochelae 38–44	absent	8–11	Northwest Pacific, Russia, abyssal

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Species	Reference	Morphology	Total height x width (mm)	Skeleton	Spicules of main axis (LxW μm)	Spicules of lateral filaments or body (LxW μm)	Spicules of basal attachment (LxW μm)	Chelae (L μm)	Sigmancistras Sigmas (L μm) (L μm) and other microscleres		Locality, depth range
Abyssocladia corniculiphora Hestetun, Rapp & Xavier, 2017	Hestetun et al. , 2017: 170 , Fig. 3	‘Centipedal’, erect, single axis of a flattened stem with 2 rows of filaments in opposite directions, and a few special stalked cup-like projections	25 x 0.5	Main stem with longitudinal mycalostyles, filaments arranged in 2 rows with mycalostyles perpendicular to stem, cuplike stalked bodies made exclusively of smaller styles	mycalostyles 375-(577)-722 x 11.6-(18.1)-21.7	mycalostyles undifferentiated from main axis	styles in cup- like stalked bodies 189-(367)-573 x 5.3-(9.0)-13.2	cleistochelae 23.0-(36.1)-59	7.5-(9.2)-11.9 absent		Southwestern Indian Ocean Ridge, off South Africa, bathyal
Abyssocladia desmophora ( Hooper & Lévi, 1989 )	Hooper & Lévi, 1989: 437–440 , Figs 1–4	Erect, bifurcate ‘pedunculate’, cylindricdal stem, fusiform body strongly hispid	24 x 2	Peduncle and stem with axial column of longer subtylostyles loosely surrounded by desmas, body with diverging radial bundles of shorter subtylostyles protruding through ectosome	subtylostyles (peduncle axis) 450–510 x 8–10 subtylostyles (body) 320–420 x 6–8 monocrepedial desmas 320–420 x 10–20	undifferentiated from axis	undifferentiated from axis	arcuate isochelae 60–85	absent	30–40 x 2	continental slope, off Townsville region Great Barrier Reef, bathyal

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Species	Reference	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)	Sigman cistras (L μm)	Sigmas Locality, depth (L μm) range and other microscleres
Abyssocladia diegoramirezensis Lopes, Bravo & Hajdu, 2011	Lopes, Bravo & Hajdu, 2011: 410–413 , Figs 3–4	Erect, short thin peduncle and a dichotomously branched ‘centipedal’ body in a single plane, with numerous lateral filaments regularly arranged in two rows	5–7 x 2–4	Main axis with large axial styles/ styles/ mycalostyles mycalostyles and substrongyles 257–1125 arranged longitudinally, x 7–33 also with few short branch styles/ strongyles, axis mycalostyles of filaments with 287–975 mycalostyles anchored x 3–20 in the main axis by strongyles their bases 175–1025 x 15–48		Substrongyles 287–1098 x 13–38	not recorded	Arcuate isochelae- cleistochelae 50–83 Abyssochelae 15–35	13–20	10–13	Diego Ramírez Archipelago, South America, bathyal
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	styles 620–2500 x 7–35	undifferentiated	base missing	arcuate isochela 80–170 abyssochelaecleistochelae 40–45 anisochelae 9.5–11	1, 30–40 2, 9.5–12.5	absent	North-Fijian back-arc Basin, bathyal
Abyssocladia faranauti Hestetun, Fourt, Vacelet, Boury- Esnault & Rapp, 2015	Hestetun, Fourt,Vacelet, Boury-Esnault & Rapp, 2015: 3–4 , Fig. 2	Elongate, pinnate ‘centipedal’ body, bearing thin regularly spaced lateral filaments	50 x 1	Central axis a dense core of large longitudinal mycalostyles, filaments composed of smaller mycalostyles anchored symmetrically and crossing main axis	mycalostyles 1, 1020-(1364)-1640 x 31.4-(43.8)-55.0 mycalostyles 2, 280-(395)-560 x 7.9-(13.4)-17.3 mycalostyles 3, 130-(193)-270 x 3.1-(5.7)-7.9	mycalostyles 530-(689)-940 x 14.1-(20.7)- 29.8	base missing	arcuate isochelae 48.7–(61.2)- 67.5 arcuate iso-or cleistochelae 22.0–(25.6)- 33	6.3-(6.4)- 7.9	absent	Mid-Atlantic Ridge- Fracture Zone intersection, bathyal

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Species	Reference	Morphology	Total height x width (mm)	Skeleton	Spicules of main axis (LxW μm)	Spicules of lateral filaments or body (LxW μm)	Spicules of Chelae basal attachment (L μm) (LxW μm)		Sigman cistras (L μm)	Sigmas Locality, (L μm) depth range and other microscleres
Abyssocladia Lehnert , flagrum Stone & (Lehnert, Heimler, Stone & 2006: Heimler, 20–23, 2006) Figs 12–13		Whip-like ‘bottlebrush’, cylindrical, with upper portion (body) highly filamentous	540 x 2.5–4	Central axis consists of interwoven polyspicular tracts of mycalostyles, filaments supported by single polyspicular tracts of mycalostyles	mycalostyles 980–1320 x 20–28	undifferentiated	absent	arcuate isochelae 98–112 palmate- arcuate isochelae 28–43	absent	sigmas 1, 37–48 sigmas 2, 17–20	Aleutian Islands, Alaska, mesophotic
Abyssocladia Hestetun , fryerae Rapp & Hestetun, Pomponi, Rapp & 2019: 4, Pomponi, Fig. 2 2019		Pedunculate, short stem, disc-shaped droplet-like 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	subtylostyles 582–1130 x 3.8–23.5	undifferentiated	undifferentiated	arcuate isochelae 77.9–110.3	17.9–22.9	absent	Marianas, NW Pacific, abyssal
Abyssocladia Hestetun e t hemiradiata al. , 2017: Hestetun, 170–172, Rapp & Fig. 4 Xavier, 2017		‘Bottlebrush’ erect, slender single-axis stem with projections in all directions, apex enlarged hemispherical, slightly flattened	25 x 1.5	Main stem with several bundles of tightly packed mycalostyles surrounding less dense core, swollen apex hispid composed of longest tylostyles radiating outwards	tylostyles 1, 2000–5000 x 5.9-(9.6)-12.1 tylostyles 2, 775-(988)-1327 x 6.1-(8.4)-11.4 tylostyles 3,333-(536)-687 x 5.6-(7.3)-10.0 tylostyles 4,108-(185)-303 x 4.3-(5.8)-8.2 mycalostyles 444-(868)-1290 x 15.6-(19.9)-22.8 styles 1, 508-(665)- 1166 x 8.6-(12.7)-16.8 styles 2, 141-(202)-224 x 2.9-(4.2)-5.3	undifferentiated	base missing	arcuate isochelae 20.3-(30.8)- 56.7	5.3-(6.9)- 10.3	11.0-(16.1)- 22	Southwestern Indian Ocean Ridge, off South Africa, bathyal

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Species	Reference	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)	Sigman cistras (L μm)	Sigmas (L μm) and other microscleres	Locality, depth range
Abyssocladia huitzilopochtli Vacelet, 2006	Vacelet, 2006: 569–573 , Figs 9–11	Erect ‘pedunculate’, 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	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	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	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	styles 1075–1800 x 21–33	undifferentiated	base missing	abyssochelae- cleistochelae 80–100 arcuate isochelae 14–150	15–18	acantho- microxeas 130–350 x 3–5	Easter microplate, East Pacific Rise, bathyal
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	mycalostyles 1262–2321 x 19–33	subtylostyles 1018–1994 x 20–35 tylostyles 275–592 x 6–15	undifferentiated arcuate isochelae 78–132		22–32	absent	Marianas, NW Pacific, abyssal

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Species	Reference	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
Abyssocladia koltuni ( Ereskovsky & Willenz, 2007 )	Ereskovsky & Willenz, 2007: 1383–1385 , Figs 2–6	‘Bush-shaped’, with basal plate supporting erect radiating cylindrical anastomosing branches, with terminations flat, oval and ending in filamentous projections	70–120	Branches with densely packed bundles of megascleres running parallel to body axis, flat distal skeleton with radiating bunches of larger styles terminating in filaments, and a tangential network of smaller styles and chelae	principal mycalostyles 1, 575.3-(717.4)-906.2 x 17.1-(23.5)-27.5	dermal subtylostyles 1, 474-(546.3)- 579.6 x 10.6-(13.6)- 15.4 dermal subtylostyles 2, 286-(346.8)- 398.2 x 7.7-(9.2)-10.4	not recorded	anchorate spatuliferous isochelae 58.2-(75.8)- 82.6 small anchorate isochelae 30.5-(37.5)- 46.3	10.9-(13.6)- 17.8	34.8-(54.2)- 69.6	Sea of Okhotsk, Russia, mesophotic
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	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	15–20	absent	Far eastern Solomon Islands, bathyal
Abyssocladia leverhulmei Goodwin, Berman, Downey & Hendry, 2017	Goodwin, Berman, Downey & Hendry, 2017: 40–41 , Fig. 2	Dichotomously branched, rounded ‘centipedal’ peduncle, small rounded basal attachment, upper branches flattened with projecting filaments	15–60 x 2–3	Peduncle with interlocked strongyles (the larger style-like), stem has a core of ascending styles with tangential bundles projecting with points outwards	mycalostyles -styles 409–1034 x 10–35	undifferentiated	strongyles 56–640 x 9–43	arcuate chelae 45–79 abyssochelae 22–40	8–19	absent	Drake Passage seamounts, South America, mesophotic -bathyal

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Species	Reference	Morphology	Total height x width (mm)	Skeleton	Spicules of main axis (LxW μm)	Spicules of lateral filaments or body (LxW μm)	Spicules of basal attachment (LxW μm)	Chelae (L μm)	Sigmancistras Sigmas Locality, (L μm) (L μm) depth range and other microscleres
Abyssocladia marianensis Hestetun, Rapp & Pomponi, 2019	Hestetun, Rapp & Pomponi, 2019: 8 , Fig. 4	Stalked, long stem with small basal plate attachment, umbrella-shaped body with single horizontal filament crown	130 x 20	Stem with tightly packed mycalostyles slightly twisted tracts, umbrella body with radiating bundles of mycalostyles, oxeas in fleshy top part of body	mycalostyles 1824–2458 x 20–39	mycalostyles same as stem oxeas 289–477 x 4–7	undifferentiated	palmate chelae/ cleistochelae 1, 68–92 palmate chelae/ cleistochelae 2, 26–42	1, 50–65 2, 31–46	absent	Marianas, NW Pacific, abyssal
Abyssocladia myojinensis Ise & Vacelet, 2010	Ise & Vacelet, 2010: 892–893 , Figs 6–7	Erect, ‘centipedal’ pinnate, slender, flattened bearing numerous long lateral filaments bilaterally symmetrical in one plane, thin axis attached with enlarged base	19 x 0.5–0.8	Main axis with bundles of large styles-mycalostyles, spirally twisted in bundles in peduncle, axis of filaments bundle of styles, base with bundles of small styles	styles- mycalostyles 1 (axis), 278-(768)-1053 x 7–(10)-34	styles- mycalostyles 2 (filaments) 605-(773)-890 x 11-(15)-18	styles- mycalostyles 3 (base), 273–(301)-350 x 4–(5)-6	cleistochelae 59-(68)-78 abyssochelae 28-(36)-44	5–6	absent	IzuOgasawara Arc, Japan, mesophotic
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	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	substrongyles 395-(642)-980 x 22-(36)-45	cleistochelae- abyssochelae 38-(54)-75	1, 20–23 2, 9–12	absent	IzuOgasawara Arc, Japan, mesophotic

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Species	Reference	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
Abyssocladia naudur Vacelet, 2006	Vacelet, 2006: 577–579 , Figs 15–16	Centipedal’, small, erect, on enlarged base, with slender flattened spicular axis bearing numerous lateral filaments regularly arranged in 2 rows	40	Axis with large fusiform styles longitudinally arranged, spirally twisted at the basal where axis is lined by substrongyles, axis of filaments lightly conical at the base, with styles anchored by their head and the point outwardly directed	styles- substrongyles (main axis) 700–1600 x 10–37	styles (lateral branches) 330–1000 x 5–15	styles- substrongyles, strongyles (base and exterior coating of axis) 30–825 x 8–30	cleistochelae 30–72	1, 15–19 (6.5–9.5 x 1) 2, 5–8	absent	East Pacific Rise, bathyal
Abyssocladia oxeata Koltun, 1970	Koltun, 1970: 191 , Fig. 17, Plate 3	Club-shaped, long thin ‘peduncle’ supported by flattened base plate, swollen subspherical apex with microfilament ous surface	20	Main skeleton of axial type with the stem formed of long fusiform needles, but at the capitate parts they are shorter	mycalostyles 1300–3500 x 16–44	microxeas 88–375 x 4–6 microstyles 33	undifferentiated	arcuate isochelae 77–105	absent	33	Northwest Pacific, Russia, hadal
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	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	9.4–(9.8)–11	absent	Muir Seamount, Bermuda, bathyal

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Species	Reference	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
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	mycalostyles 846–1741 x 15–53	mycalostyles same as stem subtylostyles 491–1134 x 7–17 tylostyles 181–573 x 4–9	undifferentiated	palmate isochelae 20–40	6–9	absent	Marianas, NW Pacific, bathyal
Abyssocladia symmetrica ( Ridley & Dendy, 1886 )	Ridley & Dendy, 1887: 77–78 , Pl. XIX Figs 6, Pl. XXVI Fig. 4, Pl. XLVI Fig. 7; Hestetun et al. , 2017: 172–173 , Fig. 5	Erect, ‘bottlebrush’, slender with a central stem with rows of filaments in all directions	22–44 x 2–3	Main stem with 2 concentric rings of longitudinal mycalostyle bundles with less densely packed tissue in the center, filaments are supported by radial rays of mycalostyles anchored in the main stem skeleton	mycalostyles 363-(688)-1105 x 4.8-(15.6)-26.3	undifferentiated	base missing	palmate/ arcuate isochelae 23.6-(28.0)- 34.2	7.3-(9.4)-16.6	absent	Prince Edward Island, Canada, mesophotic
Abyssocladia tecta Hestetun, Fourt, Vacelet, Boury-Esnault & Rapp, 2015	Hestetun, Fourt, Vacelet, Boury- Esnault & Rapp, 2015: 4–6, Fig. 3	Elongate ‘centipdal’ pinnate attached by basal plate, bearing long thin lateral filaments regularly spaced	30 x 0.2	Basal plate skeleton consists of 2 types of substrongyles, main axis a dense core of large longitudinal tracts of mycalostyles, filaments with plumose tracts of large and small mycalostyles points directed outwards	principal mycalostyles 1040–(1241)- 1490 x 20.4– (27.8)-34.5	mycalostyles 370-(617)-860 x 6.3-(11.5)-17.3	substrongyles 1, 220-(294)-430 x 15.7-(25.2)- 31.4 substrongyles 2, 45-(84)-180 x 7.9-(12.6)- 20.4	Cleistochelae 42.4-(47.4)- 55 Abyssochelae 11.0-(12.1)- 14.1	6.3-(6.6)-7.9	absent	Mid-Atlantic Ridge-Fracture Zone intersection, bathyal

...Continued on the next page TABLE 1. (Continued)

Species	Reference	Morphology	Total height x width (mm)	Skeleton	Spicules of main axis (LxW μm)	Spicules of lateral filaments or body (LxW μm)	Spicules of basal attachment (LxW μm)	Chelae (L μm) Sigmancistras Sigmas (L μm) (L μm) and other microscleres			Locality, depth range
Abyssocladia umbellata Lopes, Bravo & Hajdu, 2011	Lopes, Bravo & Hajdu, 2011: 413–417 , Figs 5–6	Erect, short thin peduncle, flattened ‘centipedal’ body, bearing numerous lateral filaments	51–57 x 7–12	Main axis with longitudinal bundles of large mycalostyles and few short strongyles, basal axis lined by substrongyles, filaments with mycalostyles inserted by their bases, base with styles or substrongyles	mycalostyles 1 308–1400 x 18–45 strongyles 72–185 x 8–50	mycalostyles 3 205–534 x 5–15	styles or substrongyles 133–1221 x 20–50	Arcuate 15–23 isochelae 75–133 Abyssochelae 23–63		absent	Diego Ramírez Archipelago, South America, bathyal
Abyssocladia villosa Hestetun, Rapp & Pomponi, 2019	Hestetun, Rapp & Pomponi, 2019: 12 , Fig. 6	Stalked, hispid, cup-shaped body, horizontal radial slightly apically oriented filament crown close to top margin, small basal plate	50 x 0.5–2 Solid core of tightly packed large longitudinal subtylostyles, plumose brushes of smaller tylostyles project out from stem producing hispid surface, top of body with radiating larger tylostyles		subtylostyles 718–1443 x 14–25 tylostyles 2, 154–488 x 4–15	tylostyles 1, 549–1356 x 7–19	undifferentiated	Cleistochelae, 8–12 arcuate isochelae 31–90		absent	Marianas, NW Pacific, bathyal

Material examined : Holotype : QM G337545 , off Fraser Island , Station 115, Queensland , Australia , 25 o 19’31.1”– 25° 21’ 4.7” S , 154 o 4’5.9”– 154° 4’ 33.6” E , 2350– 2342 m , Beam Trawl , Coll. Merrick Ekins on RV Investigator , Cruise IN2017_ V03 , Sample 115–126, 11/vi/2017 . Paratypes : QM G337546 off Fraser Island , Station 115, Queensland , Australia , 25 o 19’31.1”– 25° 21’ 4.7” S , 154 o 4’5.9”– 154° 4’ 33.6” E , 2350– 2342 m , Beam Trawl , Coll. Merrick Ekins on RV Investigator , Cruise IN2017_ V03 , sample 115–144, 11/vi/2017 ; QM G337506 same collection data as QM G337546, Sample 115–144.1 ; QM G337507 same collection data as QM G337546, Sample 115–144.2 . Etymology : Named after the Dutch artist Mauritus Cornelis Escher, 1898–1972 , for the shape of the abyssochelae resembling a 3D puzzle similar to something that Escher might have illustrated. Distribution : This species is presently known only from the type locality off Fraser Island, Queensland , Australia , at abyssal depth. Description: Growth form : Erect, pedunculate sponges up to 305 mm long and 20 mm thick at the widest part of the body, anchored in muddy substrate by a multi-furcate rhizoid basal root, with a club-shaped to conical, fleshy body and long, irregularly branching filaments that extend laterally at the apex of the sponge. The stem is up to 250 mm long and only 0.15 mm wide. The main body including filaments is 39 mm in length, and including the filaments 20 mm wide. The filaments themselves are up to 10 mm in length and 0.5 mm in width. The remainders of the basal rhizoid rootlets are 15 mm long and 0.3 mm wide. Colour : Pale cream to orange on deck and beige in ethanol. Ectosomal skeleton : The ectosome of both the stem and the filaments consists of soft tissue encrusted with abyssochelae. Endosomal skeleton : The body, filaments and axis of the stem are composed of tightly bound, long, thin mycalostyles longitudinally arranged. The basal rhizoid rootlet also consists of longitudinally aligned axes of substrongyles that are shorter than those styles in the remainder of the sponge. Megascleres: Mycalostyles in the filaments and stem are long, thin and straight with sharp points (615– 1280 x 7.0– 15.2 µm ). Substrongyles from the roots are shorter and thicker, sometimes curved, often with blunt ends (92–555 x 3.4–16.7 µm ) (see Table 2 ). Microscleres : Abyssochelae with curved shaft (18.4–53.1 x 10.2–24.0 µm). In the smaller abyssochelae, the frontal and lateral alae are usually in contact with the alae offset from the opposing ones. In larger abyssochelae these alae become separated from those opposing alae. Abyssochelae of intermediate sizes were also present (see Table 2 ). TABLE 2. Spicule dimensions (µm) of the holotype and paratypes of Abyssocladia escheri sp. nov.

Specimen	Abyssochelae	Mycalostyles	Substrongyles
QM G337545 Holotype	18.4-(31.1)-53.1 x 10.0-(15.9)-23.5, n=140	745-(979)-1180 x 7.0-(9.8)-13.4, n=53	92-(346)-531 x 3.4-(7.4)-12.9, n=59
QM G337506 Paratype	20.0-(27.9)-48.1 x 10.9-(15.2)-24.0, n=41	712-(997)-1180 x 6.0-(10.6)-15.2, n=44	271-(424)-555 x 4.7-(8.3)-11.9, n=32
QM G337507 Paratype	21.6-(30.8)-45.0 x 11.6-(16.0)-22.0, n=29	615-(998)-1280 x 6.0-(9.8)-15.2, n=41	261-(384)-551 x 4.6-(10.7)-14.3, n=40
QM G337546 Paratype	19.3-(27.1)-44.7 x 10.2-(15.2)-23.4, n=39	632-(902)-1050 x 5.9-(9.1)-12.3, n=38	118-(314)-484 x 4.7-(7.4)-16.7, n=36

Molecular data: The 28S sequence of QM G337545 and G337546 are provided in the Sponge Barcoding Database under accession number SBD#2302, SBD#2303 respectively and the molecular difference to other congenerics displayed in Figure 3 . 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. Remarks: The presence of abyssochelae indicates this species belongs to the genus Abyssocladia . On gross morphology alone most species within this genus fall into three groups: ‘centipedes’ (feather-shaped), ‘bottlebrushes’ (short peduncle, large body), or ‘lollypops’ (with long pedunculate stems). The pedunculate species include A. bruuni Lévi, 1964 , A. carcharias Kelly & Vacelet, 2011 , A. claviformis Koltun, 1970 , A. desmophora ( Hooper & Lévi, 1989 ) , A. dominalba Vacelet, 2006 , A. fryerae Hestetun et al. , 2019 , A. huitzilopochtli Vacelet, 2006 , A. inflata Vacelet, 2006 , A. kellyae Hestetun et al. , 2019 , A. lakwollii Vacelet & Kelly, 2014 , A. marianensis Hestetun et a l., 2019, A. natushimae Ise & Vacelet, 2010 , A. oxeata Koltun, 1970 , A. polycephalus Hestetun et al. , 2016 c , A. stegosaurensis Hestetun et al ., 2019 and A. villosa . Abyssocladia that resemble ‘centipedes’ include: A. atlantica Lopes & Hajdu, 2014 , A. corniculiphora Hestetun et al. , 2017a , A. diegoramirezensis Lopes et al. , 2011 , A. faranauti Hestetun et al. , 2015 , A. leverhulmei Goodwin et al. , 2017 , A. myojinensis Ise & Vacelet, 2010 , A. naudur Vacelet, 2006 , A. tecta Hestetun et al. , 2015 , and A. umbellata Lopes et al. , 2011 . The third group, resembling ‘bottlebrushes’, include A. boletiphora Hestetun et al. , 2017a , A. flagrum ( Lehnert et al. , 2006 ) , A. hemiradiata Hestetun et al., 2017a and A. symmetrica ( Ridley & Dendy, 1886 ) . There is a solitary species that has a ‘bush-shaped’ anastomosing branched morphology representing multiple peduncles: A. koltuni ( Ereskovsky & Willenz, 2007 ) . Abyssocladia escheri sp. nov. is unique amongst this genus in its relatively large size (up to 30 cm total length, exceeded only by A. flagrum from the Aleutian Islands, which measures 54 cm in length); having reduced spiculation consisting only of styles as megascleres and abyssochelae as microscleres; and comparative spicule dimensions (see Table 1 ). The abyssochelae of A. escheri sp. nov. resemble in shape those of A. natushimae from the North Pacific. However the abyssochelae in this new species are much smaller than those of A. natushimae (18.4-(29.3)- 53.1 µm vs. 38-(54)- 75 µm , respectively), and additionally A. escheri sp. nov. lacks the sigmancistras and microstrongyles that are present in A. natushimae .