Ctenophora of Australia Lisa-Ann GERSHWIN South Australian Museum, North Terrace, Adelaide, South Australia 5000 (Honorary)

VOLUME 54 Part 3

Memoirs OF THE Queensland Museum

Brisbane 30 December 2010

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A Queensland Government Project Typeset at the Queensland Museum Ctenophora of Australia Lisa-ann GERSHWIN South Australian Museum, North Terrace, Adelaide, South Australia 5000 (Honorary). Email: [email protected] Wolfgang ZEIDLER South Australian Museum, North Terrace, Adelaide, South Australia 5000. Email: [email protected] Peter J.F. DAVIE Queensland Museum, PO Box 3300, South Brisbane, Queensland 4101

Citation: Gershwin, L., Zeidler, W. & Davie, P.J.F. 2010 12 30. Ctenophora of Australia. In, Davie, P.J.F. & Phillips, J.A. (Eds), Proceedings of the Thirteenth International Marine Biological Workshop, the Marine Fauna and Flora of Moreton Bay, Queensland. Memoirs of the Queensland Museum 54(3): 1–45. Brisbane. ISSN 0079–8835.

ABSTRACT An overview of the Ctenophora of Australia is presented based on limited collecting efforts, together with the description of seven species new to science. Euplokamis evansae sp. nov. (Cydippida: Euplokamididae), has only been found in Tasmanian waters; it is distinguished from its congeners in the comb rows being about 4/5 of the body length, the tentacle bulbs being long and close to the stomach, and having extremely long polar plates. Pukia falcata gen. nov., sp. nov. (Cydippida: Pukiidae fam. nov.), is found across the tropical and subtropical north of Australia from Port Hedland, Western Australia, to the Moreton Bay region of southeastern Queensland; it has beehive-coiled tentilla, similar to those of the Euplokamididae, but differs from all other cydippids in having crescentic-shaped tentacle bulbs. Bolinopsis ashleyi sp. nov. (Lobata: Bolinopsidae), from Moreton Bay, differs from its large-lobed congeners in its striking pattern of broad red streaks along the ctene rows. Leucothea filmersankeyi sp. nov. (Lobata: Leucotheidae), from northwestern Tasmania, differs from other species in its genus in having narrow, blind tentacular pits; long slender papillae; and a pale orange tint to the body core. Ocyropsis vance sp. nov. (Lobata: Ocyrop - sidae), from southeastern Tasmania, is distinguished from other ocyropsids with a deeply - indented hourglass-shaped stomach in lacking tentacle bulbs and spots. Finally, three new forms of benthic ctenophores are reported from the Great Barrier Reef: Coeloplana mellosa sp. nov. (Platyctenida: Coeloplanidae) is found on the soft coral Sarcophyton sp., and differs from its congeners in a unique combination of pigmentation pattern, papillae number and distribution, and statocyst palp morphology; Coeloplana reichelti sp. nov., a larger form found on algae and seagrasses, differs from others in the number and form of papillae, colour pattern, and distribution; the last, a minute form found on the Crown-of-thorns starfish, Acanthaster planci, must await additional material for proper description. The distribution of Neis cordigera is broadened to include South Australia and Western Australia, and a redes - cription of the species is provided. The potential fisheries impacts by a form of Bolinopsis in Spencer Gulf, South Australia, are discussed. An annotated classification and a key for the ctenophores of Australia are given. q Ctenophora, Ctenophores, comb jellyfish, Cydippida, Platyctenida, Lobata, Cestida, Beroida, new family, new genus, new species, new records.

Gelatinous zooplankton species are not typi - sen ted in nearly all marine pelagic zones, they cally well known, and this is especially true for are often difficult to capture and impossible (or members of the Ctenophora. Though well repre- nearly so) to preserve. Furthermore, their taxon-

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ÿ www.qm.qld.gov.au 1 Gershwin, Zeidler & Davie omy is poorly resolved, making identification cases comparative type material was non- often difficult and inconclusive. exis tent. Field-collected specimens were either Australian ctenophores are no exception and captured individually in plastic bags or glass are, in fact, even more poorly known than those jars while snorkeling, dipped individually from of many other regions. Ironically, von Lenden - jetties with a 500 µm mesh, 0.5 m-wide plankton feld (1885a) stated that the Australian cteno- net, or else ‘trawled’ with the net from a fixed phores ‘really appear to be rare.’ And while few loca tion relying on existing current for flow. species are reported, most are certainly not rare! Difficulties in preservation and inherent prob - In our experience, a small cydippid that is lems with shrinkage are a fundamental problem described as new herein and placed into a new in ctenophore taxonomy. Thus, whenever possible, family, is one of the commonest members of the live specimens were observed, photographed, Australian coastal gelatinous zooplankton com - and morphological and behavioural characters mu nity, and may be found throughout the year noted prior to preservation attempts. Tissue in just about any tropical and subtropical region. samples were also collected for future study Furthermore, an unidentified species of Bolinopsis when tissues from congeners become available. blooms in such plague proportions in the South Extensive efforts were made to relax and fix Australian gulfs in the summertime that one live material, often without success. For cydippids must wonder what effect this might have on the and beroids, room temperature asphyxiation larvae and food sources of other species in the gulfs. was the only effective method of relaxation, but The first mention of a ctenophore in Austra- was extremely labour intensive; preservation in lian waters was by Quoy & Gaimard (1824: 575), about 2% formalin produced moderately good who noted a Beroe from Port Jackson. Since that specimens. For lobates, no method of relaxation time, at least 24 additional species of ctenophores could be obtained, and efforts to preserve speci- have been reported from Australian coastal mens universally resulted in total tissue disinte- waters, with another ten newly reported herein, gration, leaving only scattered ctenes; the only including seven new to science. exception was Ocyropsis vance sp. nov., which During the course of the Thirteenth Inter- preserved quite perfectly in 1-2% formalin. national Marine Biological Workshop, in More- Measurements were made on preserved mate - ton Bay, Queensland, numerous collections were ri al with Max-Cal digital calipers, to the nearest made of ctenophores. As we were identifying 0.01 mm. Every effort was made to obtain true this material, we realised that it would be very dimensions across the widest points; however, useful to attempt to compile the first synopsis some specimens were too brittle to be spread of Australian ctenophores, and thus provide a out, in which case absolute measurements were foundation for productive future work. To do taken across the two farthest available points. this it was necessary to document not only the Body length (BL) was measured from the aboral historical literature, but also specimens in pole to the tip of the mouth for all specimens. In museum collections, and our own samples lobate ctenophores, lobe length (LL) was measured gathered over the last ten years. This has resul - from the inner corner where the lobe attaches to ted in numerous new distribution records, and the body to the distal tip; total length (TL) the descriptions of the seven new species includes the lobes. presented here. It is highly likely that many Morphological examinations were made more species will prove to be present than so far under a variety of dissecting scopes, depending recognised, and our knowledge of the on what was available at the institution where distibutional ranges of many will no doubt the specimens were studied. Microscopic and increase significantly with further targeted macroscopic digital images were made of all collecting. observable structures with Fujifilm MX-700 and MX-2700 cameras, Nikon CoolPix 995, and MATERIALS AND METHODS Sony DVD-201e in JPG format. While it was not The collections of ctenophores held in all possible to publish all photographs made of each Australian museums were examined; in most taxon, we have compiled a library of images of

2 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Table 1. An annotated checklist of ctenophores recorded from Australian waters, including all previous records from Moreton Bay. Nomenclatural notes are given in parentheses, where appropriate. Abbreviations used: Queensland (QLD), New South Wales (NSW), Victoria (Vic), Tasmania (Tas), South Australia (SA), Western Australia (WA), Northern Territory (NT), Great Barrier Reef (GBR), Australia ‘Unspecified’ (AU), Southern Australia (SO), Northern Australia (NO); Species names highlighted in bold are dealt with in more detail in the present work.

Family Species Moreton Bay Records Endemic/Australian Distribution Phylum CTENOPHORA Eschscholtz, 1829 Class TENTACULATA Eschscholtz, 1825 Order CYDIPPIDA Lesson, 1843 Euplokamididae Euplokamis evansae sp. Endemic: TAS. Mills, 1987 nov. Pleurobrachiidae Chun, Pleurobrachia pileus (O. F. Moreton Bay 1880 Müller, 1776) (Greenwood 1980; Gorman 1988); incorrect ID; referable to Pukia falcata, gen. nov., sp. nov. Pleurobrachia spp. NSW (Dakin & Colefax 1940); QLD (Hamond 1971; Greenwood 1980). [All previous records probably referable to Pukia falcata gen. nov., sp. nov.]. New records for SA, WA, and TAS. Pleurobrachiidae indet. Southern Australia (Gowlet-Holmes 2008). Pukiidae ‘Sea Gooseberry’ Moreton Bay (Davie fam. nov. 1998). Pukia falcata gen. nov., Moreton Bay [herein] Endemic tropical Australia: sp. nov. QLD, NT, WA. Order PLATYCTENIDA Bourne 1900 Coeloplanidae Coeloplana meteoris Thiel, QLD (Arnold 1993). Willey, 1896 1968 Coeloplana scaberiae Endemic: SA. Matsumoto & Gowlett-Holmes, 1996 Coeloplana thomsoni Endemic: WA. Matsumoto, 1999 Coeloplana willeyi Abbott, VIC (Smith & Plant 1976). 1907 Coeloplana spp. QLD (Stephenson et al. 1931). Southern Australia (Edgar 1997, 2000). Coeloplana mellosa sp. Endemic: Great Barrier Reef nov. on Sarcophyton sp. Coeloplana reichelti sp. Endemic: Great Barrier Reef nov. on algae and seagrass. continued ...

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Table 1 (continued) ... Family Species Moreton Bay Endemic/Australian Records Distribution Coeloplana sp. A Great Barrier Reef on Acanthaster planci. Ctenoplanidae Ctenoplana sp. Off Townsville, Willey, 1896 planktonic (Hamner, in Matsumoto 1999). Order LOBATA Eschscholtz, 1825 Bolinopsidae Bolinopsis chuni (von Endemic: NSW (Whitelegge Bigelow, 1912 Lendenfeld, 1885a) 1889; Dakin & Colefax 1933). Bolinopsis spp. NSW (Dakin & Colefax 1940; Dakin & Bennett 1987; Edgar 1997, 2000); SA (Gershwin & Zeidler 2003; Gowlett-Holmes 2008). New records for QLD, TAS, NT, and WA. Bolinopsis ashleyi sp. Moreton Bay [herein] So far only known from nov. Moreton Bay. Bolinopsis sp. SA (clear body with magenta canals). Mnemiopsis sp. Moreton Bay Probably erroneous ID. (Greenwood, 1980) Leucotheidae Leucothea Endemic: TAS, SA. Lesson, 1843 filmersankeyi sp. nov. Leucothea spp. Moreton Bay QLD (Harbison & Miller 1986); VIC-TAS-NSW (Edgar 1997, 2000); QLD (Gershwin & Zeidler 2003) Ocyropsidae Ocyropsis crystallina QLD Krumbach, 1925 crystallina Harbison & Miller, 1986 Ocyropsis maculata QLD immaculata Harbison & Miller, 1986 Ocyropsis maculata QLD maculata Harbison & Miller, 1986 Ocyropsis vance sp. Endemic: TAS. nov. Ocyropsis spp. New records for QLD, NT, and WA. Order CESTIDA Gegenbaur, 1856 Cestidae Velamen parallelum Australia-wide (Gowlett- Gegenbaur, 1856 (Fol, 1869) Holmes 2008). New records for TAS and QLD. continued ...

4 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Table 1 (continued) ... Family Species Moreton Bay Endemic/Australian Records Distribution Class NUDA Chun 1879 Order BEROIDA Eschscholtz, 1825 Beroidae Beroe cucumis NSW (Stiasny 1931); Eschscholtz, 1825 Fabricius, 1780 southern Australia (Edgar 1997, 2000). Beroe macrostomus QLD (Lesson 1829). Péron, 1807 Beroe ovale Bosc, 1802 NSW (Quoy & Gaimard 1824). Beroe ovata Bruguière, Moreton Bay WA (Goy 1990); QLD 1792 (Greenwood 1980) (Steene 1990). Beroe spp. Moreton Bay WA (Péron 1807); NSW (Hamond 1971) (Dakin & Colefax 1933, 1940); VIC (Zeidler & Gowlett-Holmes 1998); SA (Gowlett-Holmes 2008). New records for NT and TAS. Neis cordigera Lesson, Endemic: Australia. NSW 1829 (von Lendenfeld 1885b, 1885c; Whitelegge 1889; Stiasny 1931). New records for QLD, WA and SA.

Australian ctenophore specimens; images from in the SAM collection. The prefix ‘GZ’, refers to this library are available upon request. field collection numbers that correspond to Abbreviations used: Australian states are field notes and digital photos (archived in SAM); abbreviated as follows: South Australia (SA), in some instances where material was unpre - Western Australia (WA), Northern Territory servable (e.g. species of Pleurobrachia, Leucothea (NT), Tasmania (Tas), Queensland (Qld), Victoria and Bolinopsis) these notes and photos are the (VIC), and New South Wales (NSW). The Great only remaining evidence of the material exam - Barrier Reef is abbreviated ‘GBR’. Other insti - ined. Lots consist of single specimens, unless tutional abbreviations used: the Australian otherwise noted. Museum, Sydney, NSW (AM); the Museum Latin and Greek names were derived using and Art Gallery of the Northern Territory, Brown (1956). German and French texts were Darwin, NT (NTM); the Museum of Tropical translated with Globalink Power Translator v. Queensland, Townsville, Qld (MTQ); the 6.02 for Windows. Taxonomic classification is Museum of Victoria, Melbourne, VIC (MV); the modified from Mills (2007). Queensland Museum, Brisbane, Qld (QM); the Queen Victoria Museum and Art Gallery, Laun- SYSTEMATIC ACCOUNT ceston, Tas. (QVMAG); the Tasmanian Museum and Art Gallery, Hobart, Tas. (TMAG); and the PHYLUM CTENOPHORA Eschscholtz, 1829 Western Australian Museum, Perth, WA (WAM). CLASS TENTACULATA Eschscholtz, 1825 Copies of colour slides of specimens photo- ORDER CYDIPPIDA Gegenbaur, 1856 graphed by Karen Gowlett-Holmes are kept in the SAM photo-index collection (prefix ‘PH’), Family Euplokamididae Mills, 1987 with copyright retained by KGH; digital images of specimens photographed by LG are archived Euplokamis Chun, 1880

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Euplokamis evansae sp. nov. free. Comb plates about 25–30 per comb row, (Fig. 1A, B) spaced approximately 2–3 comb row-widths Material examined. HOLOTYPE: SAM-H1557 (PH apart in the live animal, about 2/3 to 1 comb 0248), Shag Rock Bay, Tasman Peninsula, Tas., 0–2 row-widths apart in the preserved animal. m, K. Gowlett-Holmes, 24.05.1995; 13.32 mm BL, 5.07 Tentacles 2, with distantly-spaced, coiled, mm tentacular width, 5.8 mm stomodael width. robust tentilla in life (Fig. 1B); could not be Diagnosis. Euplokamis with a nearly spherical completely studied in preserved retracted state body; long ctene rows, nearly 4/5 the body without damaging the delicate specimen. length; long tentacle bulbs, parallel to one Tentacle bulbs elongate, arranged closer to another and closer to the stomach than to the stomodaeum than to outer body wall; body wall; with extremely long polar plates; sub-parallel to each other; about 1/4 total body lacking pigment. length; located just forward of midline on Description. Preserved body elongated, cylin- oral-aboral axis. Tentacle sheaths long, slender, drical with rounded ends (Fig. 1A; somewhat opening very close to aboral end; with very more spherical in life: Fig. 1B), of a very soft small ostia. gelatinous consistency; with mouth protruding Infundibulum very narrow, straight, trans- both in life and preserved. parent, difficult to discern; short, less than 1/3 Comb rows 8, equally spaced around body, body length. extending from aboral end to about 4/5 body Stomodaeum broad, flat, about 2/3 body length toward oral end, leaving oral region length; narrowed somewhat behind mouth at

Table 2. Comparison of diagnostic characters of the species of Euplokamis as considered valid by Mills (1987). Data derived from original descriptions, plus Chun (1880), and Mills (1987). Abbreviations: Body length (BL), Substomodeal (SS), Subtentacular (ST), Comb row (CR). Body Comb rows Tentacle bulbs Other features E. crinita 4 mm BL; elongate, Whole BL Very small; between Extraordinarily long (Moser, slightly compressed the stomach and body ctenes covering most of 1909) wall the body; Seychelles E. dunlapae 20 mm BL; elongate, 2/3–3/4 BL Parallel to stomodaeum; Red patches along Mills, 1987 slightly compressed midway between comb rows, tentilla, stomodaeum and and tentacle bases; outer body surface Puget Sound E. helicoides 11 mm BL; Nearly Parallel to Very broad paragastric (Ralph & cylindrical whole BL stomodaeum; midway canals; faint pink colour Kaberry, between stomodaeum under comb rows; New 1950) and outer body Zealand surface E. octoptera ‘Pea-sized body’; 2/3–3/4 BL, Leaf-shaped, parallel Canals faintly pink; (Mertens, pear-shaped along crest of to stomodaeum; tentilla red; central 1833) raised midway between coast of Chile and Bay wing-like stomodaeum and of St Lawrence in the structures outer body surface Bering Strait E. stationis 25 mm BL; Full length of Small, oblique; Transparent and Chun, 1879 cylindrical the body midway between unpigmented; Gulf of stomach and body Naples wall E. evansae 13 mm BL; nearly 4/5 BL Long, parallel; closer Colourless; combs sp. nov. spherical live, to stomach than body distantly spaced; polar cylindrical wall plates extremely long; preserved; not Tasmania compressed

6 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia about the level of ctene termination; mouth E. octoptera (Mertens, 1833) having red tentilla with straight sides. Stomodaeum bears a sharp, and the ctene rows set upon raised gelatinous aborally-orientated out-pocketing along one wing-like structures. edge, believed to be an artifact. Meridional canals round in cross section; Family Pleurobrachiidae Chun, 1880 smooth-edged; smaller diameter than ctene rows they underlie, and thus obscured by them. Pleurobrachia Fleming, 1822 Internal canals could not be discerned in The following Pleurobrachia species have been preserved specimen without dissection, or in previously reported from Australia: photograph of live specimen. Pleurobrachia pileus (O. F. Müller, 1776) — Green - Statocyst in shallow, broad, nipple-shaped wood, 1980: 91 (Moreton Bay, Qld); Gorman, indentation, less than body length deep. Polar 1988: 17 and throughout, pl. 12; (Moreton plates extremely long. Bay, Qld) [incorrect identification = Pukia Polar plates extremely long, extending orally falcata sp. nov.]. up onto sides of body wall for a distance of Pleurobrachia spp. —Dakin & Colefax, 1940: 211 about 7 comb rows. (NSW); Hamond, 1971: 27 (Moreton Bay); Etymology. The specific name, evansae, is Greenwood, 1980: 88, 91 (Moreton Bay, Qld) named after Jill Evans, librarian at the South [All probably erroneous identifications.] Pleurobrachiidae indet. — Gowlett-Holmes, Australian Museum, in recognition of the heroic 2008: 54 (southern Australia). service, often unsung, that librarians perform in the pursuit of scientific knowledge. Material examined (mostly unpreservable). SA: Kangaroo I., L. Gershwin, 5.05.1999, with short Remarks. The morphological characters of this tentacle bulbs orientated parallel to stomodaeum in species are inconsistent with known cydippid aboral half of body. Robe, W. Zeidler & L. Gershwin, genera, but most similar to the genus 9.02.2002 (GZ0232); very spherical body with short Euplokamis. However, the specimen was unfor - parallel tentacle bulbs; found in large numbers; tunately too delicate to be dissected in order to many specimens had either ?parasitic worms, or prove the existence of striated muscle in the bright yellow eggs (believed to be from the worms), or both, in the infundibulum and tentacle bulbs tentilla, considered a defining character for the (GZ0234). Robe, W. Zeidler & L. Gershwin, 21– genus by Mills (1987). Nonetheless, we conser- 22.01.2002 (GZ0103, GZ0108); with short parallel ten- vatively place this species in the genus Euplo - tacle bulbs but with distinctively teardrop-shaped kamis, rather than destroying the type specimen body; found in very large numbers; some infested or erecting a new genus, pending collection and with the hyperiid amphipod Hyperoche mediterranea. examination of more material. WA: Derby Jetty, W. Zeidler & L. Gershwin, 22.11.2000; several small specimens. Port of Broome, Euplokamis was reviewed by Mills (1987) who W. Zeidler & L. Gershwin, 25.11.2000; 5 mm juvenile. regarded five species as valid (Table 2). Of TAS: Triabunna, W. Zeidler & L. Gershwin, those, E. evansae is most similar to E. dunlapae 26.01.2002 (GZ0146). Constitution Dock, Hobart, W. Mills (1987) and E. helicoides (Ralph & Kaberry, Zeidler & L. Gershwin, 29.01.2002 (GZ0170). CSIRO 1950) in general size of the body and shape of wharf, Hobart, W. Zeidler & L. Gershwin, 1–2.02.2002. the tentacle bulbs, but differs from both in Field identification. Body spherical in life, colouration and position of the tentacle bulbs, cylindrical preserved; colourless and transparent, and the body is more spherical in the living about 1 cm. Tentacle bulbs short, parallel, and specimens, and not compressed. It is also closer to stomach than to body wall. similar to E. stationis Chun (1879) in colouration, Remarks. No morphological comments or i.e., both are transparent and unpigmented; figures were given by most of the authors that however, the two would be unlikely to be have reported on Australian material, so it is confused, with E. stationis having obliquely- impossible to determine which species they orientated tentacular bulbs. Euplokamis evansae found. While it is clear to us from our own bears less resemblance to the other species in collections that Pleurobrachia does exist in the genus, with E. crinita (Moser, 1909) having Australia, it is uncommon relative to Pukia very long ctenes covering most of the body, and falcata gen. nov., sp. nov. Thus, it seems likely

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FIG. 1. A–B, Euplokamis evansae sp. nov., holotype (Tasmania). A. Preserved. B. Live, in situ; scale bar = 2 mm; photograph by Karen Gowlett-Holmes, used with permission. C–E, Pukia falcata, gen. et sp. nov. C. Holotype (Moreton Bay), in life. D. Sagittal view. E. Tentacular view. Note in C and D crescentic tentacle bulbs that wrap around base of stomodaeum rather than running parallel to it. Note in D and E coiled tentilla near proximal ends of tentacles. Note also permanently protruding mouth. Yellowish colouration under comb rows in C & D are due to recent consumption of brine shrimp nauplii in captivity. F–G, Coeloplana mellosa sp. nov., living specimens on Sarcophyton soft coral (Great Barrier Reef). Note tentacle branching pattern in F, and tentacle streaming off to left near mid-height of G. Photographs F & G by Bette Willis (James Cook University); used with permission.

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Table 3. Comparison of diagnostic characters of ctenophore genera within the order Cydippida. Data derived from original literature, plus Bigelow (1912), Mayer (1912), Mills (1987), Harbison (1996), Wrobel & Mills (1998). Tentacle bulbs Tentilla Other features Pleurobrachiidae Ceroctena Carré & Wide, subcylindrical, Filiform Body pear-shaped with two Carré, 1991 red, opening aborally large aboral digitiform papillae Hormiphora L. Agassiz, Long, running parallel to Two or more Body oval or egg-shaped 1860 stomodaeum kinds of side branches Minictena Carré and Large, globular sheaths Filiform with 5 Body small with two Carré, 1993b types of unequal pairs of aboral colloblasts papillae; comb rows 2/3 body length Moseria Ghigi, 1909 Very long, reaching the Not described Body cylindrical; comb rows oral collar, parallel to broad, full body length, stomodaeum ending before the mouth Pleurobrachia Fleming, Short, parallel to Numerous, Body more or less spherical; 1822 stomodaeum filiform comb rows about 2/3 body length; mouth non-extensile Sabaudia Ghigi, 1909 Long, voluminous, Numerous, Cylindrical, boxy body running parallel to simple, yellow stomodaeum Tinerfe Chun, 1898 Long tentacle bases Filamentous Body elongate; two kidney-shaped gelatinous apical protuberances Euplokamididae Euplokamis Chun, 1880 Orientated midway Sparsely Body elongate; comb rows (sensu Mills, 1987) between stomodaeum arranged, coiled, extend at least 2/3 of body and outer body wall, with smooth length opening aborally muscle Haeckeliidae Aulacoctena Mortensen, With lateral processes Lacking; Body ovate, distinctly 1932 tentacles with compressed in sagittal axis, large terminal with deep lateral furrow and knob apical prolongation Haeckelia Carus, 1863 Long, narrow, opening Lacking Body ellipsoidal; tentacles toward oral pole with kleptocnidae; comb rows about 1/2 body length Ctenellidae Ctenella Carré & Carré, Teardrop-shaped, in oral Tentacles lacking Comb rows arranged in 1993a third of body, opening tentilla and pairs; kleptocnidae in medially colloblasts gastrovascular wall; suckers on lips Bathyctenidae Bathyctena Mortensen, Crescentic at base of Present Body spherical, firm, darkly 1912 stomodaeum, opening pigmented; canals with close to mouth numerous blind side branches

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Table 3 (continued) ... Tentacle bulbs Tentilla Other features Lampeidae Lampea Stechow, 1921 Short, at midpoint of Numerous, long, Long, cylindrical body, with body, parallel to filamentous highly eversible pharynx stomodaeum Cryptocodidae Cryptocoda Leloup, About half the body Present Fusiform body, with large 1938 length, parallel to combs running nearly full pharynx length Mertensiidae Callianira Péron & Short, oblique, pointing Numerous, long, Body strongly compressed, Lesueur, 1810 oral-abaxially filamentous with two very long aboral processes Charistephane Chun, Small and angled Short, widely Strongly compressed body, 1880 obliquely, at the midline spaced lacking aboral ‘keels’ of the body very near the outer body wall Mertensia Lesson, 1836 Large, crescentic against Filiform, long, Strongly compressed body, pharynx; opening red with two short aboral ‘keels’ aborally Dryodoridae Dryodora L. Agassiz, Small, globular, near the Tentacles fine, Oral opening is actually 1860 midline of the body very unbranched margin of a large near the outer body wall medusa-like chamber where food is captured; mouth is deep inside. Pukiidae fam. nov. Pukia gen. nov. Crescentic, curled Numerous, short, Apple-shaped body; around base of alternately extensile mouth; no aboral stomodaeum, opening filiform and papillae aborally coiled that most or all earlier reports are probably P. ‘beehive’ form), as Pleurobrachia. As suggested falcata gen. nov., sp. nov., which is present in by Mortensen (1912) and Bigelow (1912), and abundance throughout most of the year in the echoed by Ralph (1950), ‘great variability in areas they studied. Gorman (1988) is the only shape, size, and the position of various body author who provided figures, and the species structures is found … observations on a large she studied was unquestionably Pukia falcata number of living specimens are necessary in gen. nov., sp. nov. order to decide whether these differences occur However, while the new form is common in the living animal’ (Ralph 1950: 79). We agree throughout the tropics and sub-tropics of with these authors that Pleurobrachia is a Australia, we have found Pleurobrachia spp. problematic genus, and because we have not occasionally in temperate waters of Tasmania had sufficient comparative material available, and South Australia, as well as twice in tropical we hesitate to identify any of these forms to Western Australia. We identify those cydippid species. The genus is in need of revision. ctenophores with spherical or globular bodies; This is the first report of Pleurobrachia in the short, parallel tentacle bulbs; and with ‘normal’ waters of Tasmania, South Australia, and cydippid tentilla (i.e., without the Euplokamis Western Australia.

10 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Pukiidae new family open aborally. It may then be said that, at least Diagnosis. Cydippida with tentacle bases cres- when it comes to the tentacle bulbs, Pukia is most centic, curling around aboral end of stomo- similar to Mertensia; however Mertensia has a daeum; tentacles with numerous fine coiling strongly compressed body with two short tentilla; with very long polar plate; with apple- aboral ‘keels’. shaped body, with protruding mouth. Pukia falcata sp. nov. Type genus. Pukia gen. nov. (Fig. 1C-E) Remarks. Pukia shares some characters with various members of different families through- Pleurobrachia pileus — Gorman, 1988: 17, pl. 12, and throughout (Moreton Bay, Qld); ? Greenwood, out the Cydippida (a comparison of generic 1980: 91 (Moreton Bay, Qld). characters of cydippids is given in Table 3). ? Pleurobrachia sp. — Dakin & Colefax, 1940: 211 While we hesitate to erect another monotypic (NSW); Hamond, 1971: 27 (Moreton Bay, Qld); family, the conspicuous crescentic tentacle Greenwood, 1980: 88, 91 (Moreton Bay, Qld). bulbs and the combination of other characters Sea Gooseberry — Davie, 1998: 240 (Moreton Bay, Qld). preclude it from existing families. Material examined. HOLOTYPE: QM-G315861, 200 m offshore of Bribie I., in Moreton Bay, Queensland, surface to 2 m, water temp 24°C, 0700 hours, Puk Pukia gen. nov. Petersen for display at Underwater World (Sunshine Diagnosis. As for family. Coast), 25.11.1999. 16.2 mm total length, 14.37 mm widest diameter, measured live. PARATYPES: Qld: Type species. P. falcata sp. nov. QM-G315862, same data as holotype, 34 spec.; 2 spec. Etymology. The generic name Pukia (pronounced also distributed from this series each to AM, SAM, ‘pook-ee-uh’) is to honor Puk Scivyer (nee NTM, MTQ, and WAM. QM-G322302, Dunwich Petersen), who while at Underwater World on fishing jetty, North Stradbroke I., L. Gershwin, 10.02.2005; 4 spec. QM-G322303, same loc. as the Sunshine Coast, Queensland, gathered these G322302, 23.02.2005; 1 juv. spec. QM-G329015, same and other interesting gelatinous animals for loc. as G322302, 18.02.2005; 2 spec. QM-G329019, public display and research, and generously same loc. as G322302, 10.02.2005; 1 spec. QM-G329029, allowed us to study and keep many specimens. same loc. as G322302, 12.02.2005; 2 spec. NT: NTM Her name has Danish origins and is also the unreg., Stokes Hill Wharf, Darwin Harbour, Darwin, name of Shakespeare’s merry trickster in A L. Gershwin, 17.08.1998; 2 spec., A) 16.92 mm BL, 10.92 mm BW; B) 11.5 mm BL, 7.86 mm BW. SAM Midsummer Night’s Dream. unreg. (= GZ0005), Mandorah Jetty, 12.11.2000; Remarks. The crescentic tentacle bulbs are the numerous spec. in dilute formalin plus 3 in liquid most obvious and diagnostic character for Pukia nitrogen. WA: SAM-XH00427 (= GZ 0034 and 0041), falcata gen. nov., sp. nov. They are readily visible Port Hedland main jetty, 20°18.679’S, 118°34.438’E, in all specimens with even casual examination; 27–28.11.2000, W. Zeidler and L. Gershwin; many spec. of various sizes; preserved in dilute formalin, they are prominently curled around the aboral EtOH, and frozen. end of the pharynx. In most cydippids, the Non-type material. Qld: SAM-H1586, Palm Cove, tentacle bulbs are more or less cylindrical and Cairns, L. Gershwin, 16.12.1999; 4 juv. spec., c. 1 mm. run parallel to the pharynx; generic distinctions Palm Cove, Dec 1999 to Feb 2000; hundreds of spec. are often based on relative length of the bulbs captured during routine sampling and examined (e.g., Pleurobrachia, Hormiphora, see Wrobel & and kept to various stages. Breakwater Marina, Mills (1998)). In a few other forms the bulbs are Townsville, 24.09.2003; numerous, examined in field globular, as in Minictena (Carré & Carré, 1993b) and released. Weipa, Evans Landing, 12°39’52.6"S, 141°50’51.4"E, 10.01.2004; numerous, examined in and Dryodora (Agassiz, 1860), or teardrop- shaped, field and released. NT: SAM-H970 (= XH0174), Stokes as in Ctenella (Carré & Carré, 1993a), while those Hill Wharf, Darwin Harbour, L. Gershwin, 17.08.1998; of Aulacoctena have lateral processes (Mortensen preserved in EtOH. WA: 19°13.021'S, 121°15.399'E, 1932). Two other forms have crescentic tentacle off Eighty Mile Beach, L. Gershwin, 16.04.2004; bulbs like Pukia, but are wholly unlike Pukia in numerous small, pointy, with rows half body length. all other respects, namely Bathyctena and Description. Body apple-shaped (Fig. 1C), with Mertensia. In Bathyctena, the tentacle bulbs open the oral end broader and flatter than the indented orally, whereas in Mertensia and Pukia they aboral end; octagonal to circular in cross-section;

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 11 Gershwin, Zeidler & Davie of fairly rigid gelatinous consistency. Mouth The mouth is generally held in the extended typically protruding (Fig. 1C-E). state. When disturbed, the animal might Comb rows 8, of equal length, nearly reaching partially retract the mouth briefly, but will soon mouth, extending aborally to halfway point of resume the extended state. each wing of polar plate. Combs 30–35 per row. Captive specimens were maintained for Tentacles reaching over 175 mm when relaxed, months in pseudo-kreisels and modified box- with numerous tentilla, spaced approximately kreisels, on a twice-daily feeding of newly- 1mm apart at equal intervals. Tentilla filamen- hatched Artemia nauplii enriched with Super tous, all alike, held coiled much of the time in Selco. As in the other forms with aborally- life (Fig. 1D–E). When tentacles are fully directed tentacle sheaths, P. falcata whirls to retracted into bulbs, portions of tentacle extend ingest its captured prey. part way into sheath. Tentacle sheaths running Bioluminescence. Numerous attempts to stimu - obliquely from mid-line to body wall, opening late light responses under different conditions closer to the midline than to the aboral pole; have been unsuccessful; it seems fairly con - with short projection toward oral end, giving vincing that this species is not bioluminescent. the overall appearance of a triangle with the Haddock & Case (1995) noted that while most shortest side braced along the stomodaeum, the ctenophores are capable of producing light, longest being the oral side of the sheath to the ‘several species from the family Pleurobrachi - body wall, and the middle length being the idae produced no evidence of bioluminescence aboral side of the sheath. Tentacle bulbs crescen - capability’; thus, it is possible that as more tic, curled around aboral end of stomodaeum species in the Pukiidae become known, this (Fig. 1C–D). may also prove to contain other non-luminous Stomodaeum broad, extending halfway to forms. aboral pole. Infundibulum long, narrow, slightly Distribution. In Moreton Bay, Queensland, conical, tapering toward statocyst. Pukia falcata was found in large numbers near Statocyst deeply embedded within the aboral the surface throughout September-December, indentation of the body. 1999. Individuals ranged in size from about 5 to Meridional canals broad, with bilateral diver- 15 mm, though smaller specimens were lost ticula beneath each comb plate. Interradial canals through the outflow mesh. This species is the short, with adradial canals branched close to most abundant ctenophore from at least the infundibulum, as in Fig. 6.2b of Harbison Moreton Bay, Queensland, up around the coast (1985); perradial canals lacking. to Darwin, Northern Territory. It was also found in large numbers through much of the summer Polar plate very elongated, with two opposing (1999–2008) in the Cairns area, especially narrow, straight-sided wings, each extending during northerly winds, in Darwin Harbour in approximately 5 mm in length beyond stato - August 1998, and along much of the coastline of cyst, nearly reaching the curvature of the body. Western Australia in spring 2000 and autumn Body colourless and extremely transparent in 2004. Gorman (1988: 21) reported that this life, though radial canals may retain pigment species (as Pleurobrachia pileus) was ‘found in from food for many hours; in preservative, trans- large numbers’, and that it was among the three parent with whitish or yellow-orange tentacles. most common species in four of six sampling Etymology. The specific name is derived from trips. It may also be the species reported as the Latin adjective falcatus (= sickle-shaped, Pleurobrachia by Dakin & Colefax (1940: 211), feminine falcata), in reference to the shape of the and said to be ‘so numerous as to block up not tentacular bulbs (Brown 1956: 314). only plankton nets but even the dredge and Behaviour in life. The tentilla are uncoiled in fishing nets such as the seine used from the the relaxed state. The tentacles continuously shore…’, but this is not certain. pull in and out of the sheaths, with the tentilla Field identification. Body apple-shaped in life, along the proximal half coiling into beehive- cylindrical preserved; colourless and trans- like beads as they retract. parent, about 1.5 cm. Ctene rows almost entire

12 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia body length. Tentacle bulbs crescentic, curving (in Matsumoto (1999)) reported tropical forms, around aboral end of stomach. as we do below. Remarks. The spherical apple shape of the body bears some discussion. Most cydippids have an Family Coeloplanidae Willey, 1896 oval or cylindrical body, while some are pear- Coeloplana Kowalevsky, 1880 shaped. However, the body of Pukia is distinctly apple-shaped, i.e., largest near the oral end, The following Coeloplana species have been slightly smaller toward the aboral end. This is previously reported from Australia: most similar to that of Pleurobrachia, which is Coeloplana meteoris Thiel, 1968 — Arnold, 1993: more spherical. 16 (Pioneer Bay, Orpheus I., Qld, free-living in The mouth is somewhat different, too. In muddy sand). Pukia, the mouth is typically held rigidly in the Coeloplana scaberiae Matsumoto & Gowlett- extended position, whereas in some cydippids Holmes, 1996: 33–40, figs 1–4 (on both sides of it is highly extensile, while in others it is just a the Yorke Peninsula, SA, on the brown alga slit. Scaberia agardhii). Pukia often coils its tentilla; the coiled tentilla Coeloplana thomsoni Matsumoto, 1999: are a key character of the Euplokamididae 385–393, figs. 2, 3 (Thomson Bay, Rottnest I., (Mills, 1987). While we were unable to deter - WA, on the coralline alga Jania sp.). mine the type of muscle present in the tentilla of Coeloplana willeyi Abbott, 1902 — Smith & P. falcata, the overall description does not match Plant, 1976: 43–46 (near Portsea, Port Philip that given by Mills (1987) for Euplokamis. Bay, VIC, on the green alga Caulerpa sp. and on Specifically, she indicates that the side branches red algae). of euplokamids are widely spaced, and nor- Coeloplana spp. — Stephenson et al., 1931: 72 mally held coiled when relaxed and not in use, (Low Is., Qld, on an alcyonarian); Edgar, 1997: but stretching out at high velocity when trig - 149 (southern Australia); Edgar, 2000: 149 gered by contact with prey. She further refers to (southern Australia). ‘the natural tendency for all ctenophore tentilla to coil to some extent’. In P. falcata, the tentilla Remarks. We describe below two new tropical are sometimes coiled and sometimes filamen - species of Coeloplana. One additional tropical tous, frequently switching between the two form has been identified, but is presently states, but are most often filamentous when at awaiting additional material, and is thus rest; furthermore, the tentilla are numerous and beyond the scope of this paper. It is a minute closely spaced. form with a more or less transparent body, observed in abundance on the Crown-of-thorns ORDER PLATYCTENIDA Bourne 1900 starfish Acanthaster plancii, collected off Townsville 2003–2007. Remarks. Benthic ctenophores were first identi- fied in Australian waters by Stephenson et al. Coeloplana mellosa sp. nov. (1931), but he gave no morphological details to (Figs 1F–G, 2 A, B) allow a species determination. Subsequent records have indicated fairly high rates of Material examined. HOLOTYPE: QM-G329570, off endemism, suggesting that the biodiversity of Townsville, Qld, Bette Willis, Jun 2005, commensal Sarcophyton this peculiar group is potentially much greater on sp. (Octocorallia, Alcyoniidae); c. 2.5 cm long (live). PARATYPES: SAM-H1596, same coll. in Australia than currently known. data as holotype; 1 spec., c. 2 cm long (live). SAM- The Australian benthic ctenophores have been XH00437, same data as holotype; 2 spec. in EtOH, c. well studied in comparison to their pelagic 1-2 cm long (live). cousins. They appear to be relatively common Diagnosis. Coeloplana on Sarcophyton host; with in the temperate waters of southern Australia, brown translucent body with whitish ecto- with two endemic species described by Matsu- dermal fine meshwork pattern, appearing to moto & Gowlett-Holmes (1996) and Matsumoto the unaided eye as a whitish body with (1999). However, Arnold (1993) and Hamner hundreds of tiny brown dots; in live animals,

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 13 Gershwin, Zeidler & Davie papillae about 20, cylindrical, in longitudinal Colouration: To unaided eye, uniformly whitish X-pattern through the statocyst; or in preserved crowded with numerous tiny brown pin-point specimens over 100, in 16 rows, in three sizes, of dots (Fig. 1F); under microscopic examination, two types; with four-lobed statocyst palps. body translucent brown, with cream-coloured Description of living specimens [primarily ectodermal reticulations, giving an overall based on holotype]. Body extremely flattened, impression of honeycomb. Papillae resemble filmy, modified for creeping, resembling a platy- colourless bubbles, and are very difficult to helminth flatworm; approx. 1.5 times as long as observe at length due to their absolute wide, more or less oval in shape with undu - transparency; when viewed dorsally, they act lating margin, with outline constantly changing as windows through to the underside of the in amoeboid manner. Dorsal surface with deep ctenophore’s body. Tentacles are coloured with groove down centre of body longitudin ally, alternating whitish and clear bands. Crescentic defined along both sides by ridges bearing statocyst-palp structures are opaque off-white. ephemeral papillae; longitudinal ridges divided Notes on preserved holotype. The specimen at statocyst by latitudinal groove. preserved extremely well, and a thorough About 20 papillae, extensile, with different study of its morphology could be made. ones appearing and disappearing with move- Curiously, however, several features evident in ment, and in response to stimuli; circular in the live specimen differed markedly in the outline, with straight sides and an evenly preserved specimen. Most noticeably different rounded top; mostly of same size, but occa- were the number and arrangement of papillae, sionally one or two appearing about half again and the transparency of the animal. as broad. Arranged loosely in an X-pattern, con - The live colour pattern and opacity was such verging at statocyst. When papillae are not that the internal morphology, particularly the extended, their position cannot be distinguished. tentacle bulbs, gonads, and radial canals, could Tentacles 2, highly extensile and retractable not be interpreted; however, while preservation into sheaths inside the body; emitting through typically makes most coelenterates more opaque, two ‘chimneys’ (Fig. 1G), at opposite ends of C. mellosa actually became considerably more the body. Main shaft cylindrical, bearing nume - transparent (Fig. 2A–B). The tentacle bulbs are rous filiform, pointed tentilla, more or less large but without particular shape (e.g., anchor, evenly and sparsely spaced along length of cross-bar). The radial canals form an anasto - tentacle; the tentilla branch from main shaft mosing network along the periphery of the animal. dichotomously rather than laterally (Fig. The gonads are arranged in eight distinct rows: 1F–G). Tentacle bulb shape was uninterpre - four forming a double-turret-shaped figure-8 table due to opacity of the body; placing the through the statocyst, orientated along the ctenophores over or near a light resulted in short axis of the animal; the other four, form their balling up rapidly and tightly for over 30 loosely S-shaped structures along both sides of minutes. each of the two tentacle bulbs and sheaths. Sex Statocyst placed dorsally near midpoint of or maturity of the gonads was not investigated; body; appearing as tiny white granule, deeply however, they appear from external examin - embedded in niche between pair of perma- ation to be well developed. nently raised, opposing, crescentic, four-lobed, The papillae were difficult to study in the live palmate palps, resembling fleshy, scalloped specimens, due to their total transparency and lips (compare with Dawydoff 1938: fig. II). ephemeral nature. Over about six hours of study Gonads were not observed, but at least one of four specimens, it was concluded that they specimen was brooding live cydippid embryos typically had about 20 papillae arranged in an on ventral side of body, which began streaming X-pattern centred on the statocyst, running out in mucus strands when disturbed. along the tentacular axis, i.e., opening toward Radial canals were not observed due to the tentacles. However, a considerably more avoidance of light by ctenophores and partial complex arrangement was revealed in the opacity of their pigmentation. preserved holotype (Fig. 2A–B), where these

14 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia papillae have taken on an opaque pointed form, sides of the study bowl; in dim light it would protruding neatly from the body wall, and are quickly seek branch-axil crevices or the under- readily visible. A total of well over 100 papillae side of coral branches. Examined on a black- are present, of two different types and three back petri dish, it readily spread out, but balled different sizes, in 16 distinct rows. The four up again with direct light (e.g., for photo - primary papillae rows, i.e., crossing the stato- graphy). cyst, each contain about 20 small, cylindrical Preservation technique. Good formalin papillae, arranged in two opposing double- preserved specimens were obtained using the turret-shaped configurations along the short following technique. A specimen was placed axis, arising along the abaxial edge of the gonad into a small petri dish (55 mm diameter) nearly rows. Four rows of secondary papillae each full of the seawater that the specimen had been contain about five papillae of the same size as in living in. It was put in the dark and allowed to the primary rows; these papillae are arranged relax and adhere. If it hid in a corner, it would in a crescentic row, overlying the abaxial edge be gently prodded to get it moving, until it of the proximal portions of the lateral gonads, finally settled and relaxed on the flat bottom. It which run alongside the tentacle bases. Distal was then put into the refrigerator to immobilise to this row of papillae and along the same it. After about one hour, the petri dish gonad rows, lies the tertiary set of papillae, containing the relaxed ctenophore was very considerably smaller than the primaries and gently lowered with forceps into a 250 ml secondaries but of the same form; each of the container of chilled seawater, so as to create as four rows has about 15–20, crowded papillae, little water current as possible. Once settled on arranged along the adaxial side of the gonads. the bottom, a single drop of concentrated In addition, eight broad papillae, about ten formalin was added along one side of the jar, times the size of the tertiary papillae, are and allowed to diffuse over 30 minutes, then arranged two on each side of each tentacle another to a different side, and so on, until sheath near the opening, abaxial to, but not about 8–10 drops of formalin had been added overlying, the corresponding gonad rows, and and the specimen began to look dead (i.e., are of a different form, i.e., more like diverticula slightly crinkled and with muddled colour), at in the body wall rather than the more which time a little more formalin was added to numerous opaque, pointy, cylindrical papillae. bring the concentration to about 1–2%, and the Etymology. The specific name, mellosa, is from jar was left at room temperature. the Latin mellosus (= honey-coloured; feminine), Field identification. Benthic ctenophores, with in triple reference: A) to the honeycomb-colour an extremely flattened body modified for pattern of this species, B) to the overall honey creeping, often mistaken for flatworms, until colour, and C) to the sticky viscosity of honey in the tentacles are observed. Tentacles of the a slow trickle, for this benthic, creeping form. typical ctenophore form (i.e., with tentilla), Type locality. Reefs off Townsville, North emitting from two ephemeral ‘chimneys’ near Queensland, on Sarcophyton sp. The exact the farthest opposing ends upon the dorsal locality is unknown, as the ctenophores were surface. Comb rows lacking in adults. Colour first observed in the laboratory on corals that usually camouflaged with host, which may be had been collected from a variety of places for algae, soft coral, echinoderm, or other. the zoology classes at James Cook University. Characters specific to Coeloplana mellosa include Behaviour. In addition to the morphological its host, Sarcophyton sp. from the Great Barrier behaviour noted above in the description, Reef, the brownish body with whitish reticu - Coeloplana mellosa displays a marked sensitivity lations, the 20 or so papillae in an X-arrange- to light, actively and rapidly moving away ment in the live animal or over 100 in 16 rows in from lighted areas, or balling up tightly or the preserved specimens, and the four-lobed sinking into crevices if light is shone directly on form of the statocyst-palps. the animal. If left to its own will in darkness, it Remarks. Coeloplana mellosa differs from all would be found anywhere on the coral or the other species mainly in the number and

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 15 Gershwin, Zeidler & Davie

Table 4. Characters used to distinguish species of Coeloplana. Species are those considered valid by Matsumoto (1999). Data derived from original descriptions plus Dawydoff (1938) and Matsumoto (1999).

Host Colour Papillae Locality C. agniae Cnidarian: Clear, milky white, 4 distinct rows of 8–12 Vietnam Dawydoff, 1930b Alcyonacean: or violet brown simple papillae each in Sinularia an X-pattern through the statocyst, plus 3–4 each in 4 more rows, 2 flanking each tentacle bulb C. astericola Echinoderm: Deep red or claret 4 crescentic rows of 4 Amboina; Kei Mortensen, 1927 Asteroid: with large, irregular simple papillae, Is. Echinaster spots of emanating from the luzonicus creamy-yellow, or statocyst in a figure-8 the inverse pattern C. bannwarthi Echinoderm: Dark purple [Not described] Red Sea Krumbach, 1933 Echinoid: Diadema C. bocki Komai, Cnidarian: Dark vermillion, [Not described] Misaki, Japan 1920 Alcyonacean: dark red, brick red, Stereonephthya pink, orange, or grey stripes branching and anastomosing C. duboscqui Cnidarian: Intense orange or Papillae not observed Vietnam, Dawydoff, 1930c Pennatulid: vermillion to orange; Hawaii Pteroeides tentacles colourless C. echinicola Echinoderm: Yellow brown with 32 simple papillae: 8 Japan Tanaka, 1932 Echinoid: wide pale green larger, 24 smaller Toxopneustes margin pileolus C. gonoctena Cnidarian: Milky white or grey 5 simple papillae in Vietnam Krempf, 1920 Alcyonacean: with brown spots each of 4 rows forming Alcyonium an X-shape or figure-8, with the distal-most in each row larger C. komai Utinomi, Cnidarian: Uniformly milky 4–6 pairs of simple Sagami Bay, 1963 Alcyonacean: white or seashell papillae Japan Alcyonium pink, with yellow tentacle bases C. krusadiensis Echinoderm: Red 6–20 simple papillae India Devanesan & Asteroid: Varadarajan, 1942 ‘Pentaceros hedemanii‘ C. lineolata Fricke, Cnidarian: Milky white or 60–70 simple papillae Madagascar 1970 Alcyonacean: greenish with pale Sarcophyton yellow or yellow-green spots C. mesnili Planktonic Transparent pale 32 simple orange Vietnam Dawydoff, 1938 green, with bright papillae arranged in 8 orange statocyst and short rows papillae Continued ...

16 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Table 4 (continued) ... Host Colour Papillae Locality C. meteoris Thiel, Free-living on Clear with 4 rows of simple Somalia; 1968 soft sediments yellow-white papillae, 3–4 per row North reticulations Queensland, covering body, and Australia red pigmentation around canals, tentacle sheaths and papillae C. metschnikowii Seagrass Dorsal grey, ventral [Not described] Red Sea Kowalevsky, 1880 Zostera white C. mitsukurii Algae: red Pigmentless to 4 curved rows of 6–8 Japan Abbott, 1902 Melobesia sp., chocolate brown, papillae per row with or brown with yellow white 2–5 digitate processes, Sargassum sp. cells around margin radiating around the and two bands of sense organ in a yellow around figure-8 statocyst C. perrieri Seagrass Deep olive green Lacking papillae (?, Vietnam Dawydoff, 1930a Posidonia with sepia spots, but see Dawydoff, with narrow yellow 1938) orange margin; tentacles yellow-brown C. punctata Fricke, Cnidarian: Transparent with 70–100 simple papillae Madagascar 1970 Alcyonacean: brown or grey lines Sarcophyton parallel to the tentacle axis C. scaberiae Algae: Scaberia Solid dark orange or Four rows of simple Yorke Matsumoto & agardhii vivid red, without papillae in a figure-8 Peninsula, Gowlett-Holmes, spots of any kind pattern, and also along South 1996 the margin Australia C. sophiae Cnidarian: Brick red with milky 4 rows of 3–5 simple Vietnam Dawydoff, 1938 Gorgonian: white spots papillae Solenocaulon C. tattersalli Planktonic Transparent green 8 simple papillae; India Devanesan & sometimes with Varadarajan, 1942 secondary papillae, which may appear branched C. thomsoni Algae: Jania Pale green or white Green morph has 16 Rottnest I., Matsumoto, 1999 with yellow white papillae (8 rows of 2), Western cells scattered over four branched, others Australia dorsal surface simple; white morph with 4 large permanent papillae and 12 small ephemeral papillae C. weilli Echinoderm: Uniform brownish Lacking papillae Gulf of Dawydoff, 1938 Echinoid: red Thailand Heterocentrotus mammillatus Continued ...

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 17 Gershwin, Zeidler & Davie

Table 4 (continued) ... Host Colour Papillae Locality C. willeyi Abbott, Not specific: Deep purple, red or 20–30 cylindrical or Japan; Hawaii 1902 seagrasses, all orange fading to club-shaped papillae colours of pink, with white algae, spots along margin echinoderms and yellow blotches at base of papillae C. wuennenbergi Cnidarian: Whitish to grey, with 40 simple papillae Madagascar Fricke, 1970 Alcyonacean: dark red/violet Sarcophyton spots C. mellosa sp. nov. Cnidarian: Overall impression 20 simple papillae Great Barrier Alcyonacean: whitish with arranged in a X-shape Reef off Sarcophyton hundreds of tiny in life; preserved: over Townsville, brown dots; 100 papillae in 16 Queensland close-up, body rows, 3 sizes, and two brownish with types cream-coloured ectodermal reticulations C. reichelti sp. Botanical: Transparent pale 8–20 large, branched, Great Barrier nov. variety of red yellowish body with permanent papillae, Reef off and green hundreds of plus 8 smaller, Townsville, algae and ectodermal and cylindrical, ephemeral Queensland seagrass endodermal green papillae midway to specks, with ‘frosty’ margin white cells on papillae and margin arrangement of papillae (Table 4), although this The colour of live specimens of C. mellosa is character becomes clearer upon preservation. most similar to C. gonoctena with a milky white In the preserved condition, it is the only species body with brown spots; however, in C. gonoc - with 16 rows of papillae. Some species do have tena the brown spots are quite large, whereas in complex papillae patterns, e.g., C. agniae C. mellosa they are minute. Dawydoff, 1930(c), and C. mesnili Dawydoff, 1938, with eight rows, C. gonoctena Krempf, Coeloplana reichelti sp. nov. 1920, with the distal-most in each of four rows (Fig. 2C–F) larger than the others, C. scaberiae Matsumoto & Material examined. HOLOTYPE: QM-G329571, off Gowlett-Holmes, 1996, with papillae along the Townsville, Qld, 14.05.2008 from Reef HQ Aquarium, margin as well as in the primary figure-8, and living on broad-leafy green algae; c. 1.0 cm body C. thomsoni Matsumoto, 1999, with two differ- length (live). PARATYPES: QM-G329572, same coll. data as holotype; 1 spec., c. 0.4 cm BL (live); living on ent types of papillae in both colour morphs. seagrass overgrown with dark red algae. QM- Similarly, C. lineolata Fricke, 1970, and C. G329573, same coll. data as holotype; 1 spec., c. 0.5 punctata Fricke, 1970, are both characterised by cm BL when stretched out (live); living on green having a very large number of papillae, 60–70 algae. SAM-H1629, same coll. data as holotype; 2 in the former and 70–100 in the latter. However, specs, c. 0.5 cm BL (live); living on green algae. in all these cases, the hosts and colour patterns Diagnosis. Coeloplana on various green and red serve to readily differentiate C. mellosa from algal and seagrass hosts; with yellowish trans- each, and even though there are generalities lucent body with numerous tiny ectodermal such as ‘a large number of papillae’ or ‘multiple green specks, with whitish cells on branches of rows or types of papillae’, they are nonetheless papillae and around margin, giving frosted dissimilar in actual number and arrangement. appearance; in live animals, 20 permanent

18 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia irregularly branched, primary papillae, arranged Gonads were not observed. in figure-8 pattern, plus 8 ephemeral secondary Radial canals numerous, throughout body papillae, 4 each in two rows, cylin drical, unbran- region peripheral to tentacle bulbs and papillae; ched; with smoothly rounded statocyst palps. not anastomosing; repeatedly branching toward Description. (Based on notes from laboratory margin into progressively finer canals (Fig. examination of live specimens, primarily the 2E–F). Gastrovascular origin of some canals not holotype (Fig. 2C)). apparent. Body extremely flattened, filmy, modified for Colouration: uniform transparent pale green creeping, resembling a platyhelminth flatworm; to the unaided eye; under microscopic examin- approximately 2–3 times as long as wide, more ation, the body is transparent and colourless, or less oval-shaped to lemon- shaped with an with hundreds of bright green ectodermal and undulating margin, with outline constantly endodermal specks, particularly over the vital changing in an amoeboid manner. During region between tentacle bulbs and in a ring study period, specimens changed shape several around just proximal to the periphery, and with times from a primarily ‘short, broad form’ to a frost-like whitish granules haphazardly arranged primarily ‘long, narrow form’ and back again, around the margin. Branched papillae trans - with respect to the direction of movement parent, with conspicuously ‘frosted’ appearance (perpendicular to the tentacular axis). Dorsal on some surfaces of tips and branches (Fig. surface lacking a deep groove down centre; 2C–D). Cylindrical papillae completely trans- ventral surface with deep groove down centre parent and colourless, resembling bubbles, and of body in tentacular axis. remarkably difficult to see; microscopic exam- Papillae 28, of two types, arranged as follows ination through these windows into the inside in holoytype: permanent papillae 20, irregularly of the animal revealed presence of numerous digitate, arranged in a figure-8 pattern through particles whirling around in a random motion, statocyst (Fig. 2C); two other rows of smaller, contained within papillae. Tentacles slightly unbranched, ephemeral cylindrical secondary translucent whitish. Crescentic statocyst-palp papillae, 4 in each row, arranged about midway structures are opaque pale orange coloured, between primary rows and margin. In para - with granular appearance, resembling sand. types, which are smaller in body size, perma - Etymology. The specific name, reichelti, is given nent papillae are arranged in 2 rows of 4, close to honour Dr Russell Reichelt, Chairman of the to, but either side of, midline (Fig. 2E). Great Barrier Reef Marine Park Authority Tentacles 2, highly extensile and retractable which owns the Reef HQ Aquarium where this into sheaths inside body, at opposite ends of species was discovered; also because Dr. Reichelt body; ‘chimneys’ not apparent; instead, has been extremely supportive of scientific tentacles emitted from an opening just projects relating to jellyfish and marine-stinger proximal to ‘edge’ of body margin. Main shaft safety; and more personally because Russell of tentacles cylindrical, bearing numerous has been an inspiring mentor to the senior author. filiform, pointed tentilla, more or less evenly Type locality. Algal and seagrass beds off and sparsely spaced along length of tentacle; Townsville, north Queensland. The exact locality tentilla branch from main shaft laterally rather is unknown as they were first discovered at the than dichotomously. Tentacle bulbs in shape of Reef HQ Aquarium flourishing on algae and angular ‘C’ or half a hexagon, with tentacle seagrasses that had been collected from a emitting from ‘outside’ or ‘back’ of central bar. variety of locations nearby. Statocyst dorsal, conspicuous, located at centre Behaviour. Coeloplana reichelti showed a marked of body, midway between four central-most propensity toward the air/water interface, and branched papillae; appearing as a tiny pale had to be repeatedly drawn off it during orange granule, deeply embedded in a niche examination. Whether this is due to some between a pair of permanently raised, opposing, feature of the study conditions, or is perhaps a crescentic, simple, smoothly rounded, unscal - natural dispersal mechanism, is unknown. In loped palps, resembling fleshy lips. its undisturbed condition, C. reichelti spends

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 19 Gershwin, Zeidler & Davie

FIG. 2. A–B, Coeloplana mellosa sp. nov., holotype, preserved. A. Whole animal; note one tentacle extending beyond right edge of image, the other tentacle retracted into bulb left of centre; note also arrangement of papillae. B. Close up of left side of A. Note three different sized papillae: Primary papillae (PP) forming double-turret-shaped figure-8 in centre of A; secondary papillae (SP) along lateral gonads in A & B; tertiary papillae (TP) along adaxial side of lateral gonads in B; and large papillae (LP) along abaxial side of lateral gonads in A & B. C–F, Coeloplana reichelti sp. nov. C. Holotype, dorsal view, note figure-8 arrangement of permanent papillae. D. Paratype QM-G329572, in life, dorsal view, direction of movement is to lower right. E. Paratype QM-G329573, in life, ventral view. F. Dorsal view of E; note branching nature and arrangement of papillae in two rows. Abbreviations in C–F: radial canals (RC), tentacle bulb (TB), statocyst (ST), permanent papillae (PP).

20 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia most of its time near the tips of algae and and conspicuously branched, and greatly seagrass, with the tentacles streamed out, up to outnumber those in C. tattersalli. C. tattersalli is about 20 cm, both day and night. also planktonic, whereas C. reichelti is benthic. Field identification. C. reichelti is highly cryptic C. thomsoni Matsumoto, 1999, has four large, but appears to prefer tropical algal and plant branched, permanent papillae, plus 12 smaller, hosts. Diagnostic characters are: yellowish body simple ephemeral papillae, whereas C. reichelti with green flecks and whitish frost-like cells on has 20 and 8, respectively; furthermore, C. papillae and margin; conspicuous, irregularly thomsoni is found only on a single algal host, branched permanent papillae, plus simple, whereas C. reichelti is less discerning. As ephemeral papillae; and the smoothly rounded already described C. reichelti has a distinctive form of the statocyst-palps. colour pattern within the genus. A comparison Remarks. Coeloplana reichelti appears to be most of primary diagnostic characters for species of similar in general form to C. mitsukurii from Coeloplana is presented in Table 4. Japan. Both have branched papillae, and both Family Ctenoplanidae Willey, 1896 have white cells around the margin and are found on various species of algae. However, C. Ctenoplana Korotneff, 1886 mitsukurii has 24–32 branched papillae and no mention of secondary, simple papillae, whereas Ctenoplana sp. there are only 8–20 branched papillae in C. Only one unidentified Ctenoplana species has reichelti, plus eight simple, ephemeral papillae. been previously reported from Australia: Interestingly, Abbott (1902) described the papillae of C. mitsukurii as being ‘entire or Ctenoplana sp. — Hamner, in Matsumoto, digitate and fringed’; this raises the possibility 1999: 386 (off Townsville, Qld; planktonic). of further difference between the two in the Field identification. Essentially a planktonic actual form of the papillae, as one would not version of a benthic ctenophore. describe the papillae of C. reichelti as digitate Remarks. The above report by Hamner noted and fringed, but possibly more as ‘jaggedly only that the specimen was transparent and branched, or haphazardly dendritic’. Without planktonic, and found off Townsville, but gave seeing fresh material of C. mitsukurii, it is no further details. It will be interesting to see difficult to compare the papillae branching what species is/are present in Australian forms between the two species. The two species waters, when found again. further differ in colour, with C. mitsukurii being pigmentless to chocolate brown, but C. reichelti ORDER LOBATA Eschscholtz, 1825 being whitish to yellowish with many bright Family Bolinopsidae Bigelow, 1912 green flecks. Furthermore, whereas C. mitsukurii is found in temperate northern hemisphere, C. Bolinopsis L. Agassiz, 1860 reichelti is found in the tropical southern (sensu Mayer, 1912) hemisphere. C. reichelti is also found on sea - grass, but this does not appear to have been Bolinopsis species recorded for C. mitsukurii. Whilst the ephemeral (Fig. 3A–B) papillae and seagrass host could have been overlooked, the number of primary papillae, The following Bolinopsis species have been the colour differences, and the great geograph - previously reported from Australia: ical separation, would seem to differentiate the Bolina chuni von Lendenfeld, 1885a: 929–931, two quite readily. pl. 44–45 (Port Jackson, NSW); Whitelegge, Two other species of Coeloplana also have 1889: 197 (Port Jackson); Dakin & Colefax, 1933: branched papillae. In C. tattersalli Devanesan & 198 (NSW). Varadarajan, 1942, the secondary papillae are Bolina sp. — Dakin & Colefax, 1940: 211 (NSW). said to sometimes appear branched, but in C. Comb jellies — Dakin & Bennett, 1987: 179 reichelti the primary papillae are permanently (NSW).

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 21 Gershwin, Zeidler & Davie

22 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Bolinopsis spp. — Edgar, 1997: 150 (NSW); numerous. WA: 19°07’56.3S, 121°18’26.3E, off Eighty Edgar, 2000: 150 (NSW); Gershwin & Zeidler, Mile Beach, 10.05.2004; several with thick comb rows. 2003: 237 (Great Australian Bight); Gowlett- Field identification. Body similar in outline to Holmes, 2008: 54 (SA). a chicken egg in size and shape, i.e., with one end somewhat larger and more rounded than Material examined (mostly unpreservable). Wholly the other; more or less colourless and trans - transparent forms: Qld: Palm Cove, Dec 1999 to Feb 2000; hundreds of specimens captured during routine parent; to about 8 cm long. Two lobes at larger sampling and examined to various stages; to about 7 end, capable of expanding out somewhat like cm long (Fig. 3B). SAM-XH436, Palm Cove, 10.12.1999; scoops, or often held curled in, giving the 1 spec. in EtOH. Palm Cove, 19.02.2000; transparent animal the egg-like shape. The body is extremely with 2 red dots on each side. Palm Cove, 22.02.2000; 1 soft and breaks apart easily. Ctene rows extend spec. with 2 red dots on each side, 2 cm BL. Quarter- along on the body, four extending out along the deck Marina, Townsville, 5.11.2003; 1 small, examined lobes, and the other four ending in rabbit- in the field and released. NT: Stokes Hill Wharf, Darwin Harbour, 6.08.1998; numerous specs. Man - ear-like auricles between the lobes. dorah Jetty, 12.11.2000; 2 specs in liquid nitrogen and Remarks. Unidentified, unpreservable forms EtOH. Dundee Beach, Fog Bay, 15.11.2000; numerous. of the genus Bolinopsis were found in great WA: Cockburn Groin, Cockburn Sound, 7.03.1999; abundance along much of coastal Qld, SA numerous specs examined. Derby Jetty, 22.11.2000; (especially upper Spencer Gulf and Kangaroo several small to medium, purplish clear. Port of Broome, 25.11.2000; 4 specs, 1–2 cm; 5 frozen (GZ0024). I.), along the south-west coast of WA, and in Port Hedland Jetty, 27–28 Nov 2000; numerous. Darwin Harbour, NT. Bolinopsis has not been Hearson Cove, near Dampier, 30.11.2000; 1 spec. previously reported in the waters of these 19°13.021S, 121°15.399E, off Eighty Mile Beach, states. No species identification was made, 16.04.2004; numerous. 19°07’56.3S, 121°18’26.3E, off though medium to small lobes and a trans - Eighty Mile Beach, 10.05.2004; numerous. 19°02’09.0S, parent/colourless body were noted. Another 121°21’44.1E, off Eighty Mile Beach, 11.05.2004; numerous. SA: Whyalla Marina, 12.05.1999; in plague transparent and colourless form from Tasmania proportions up to about 4cm long, typically orientat- is characterised by short, triangular aboral ing in water column with aboral end up; highly bio - extensions (Fig. 3A); these have not been luminescent. Murat Bay Jetty, Ceduna, 19–20.02.2002; observed in other Australian forms. It is pos- large numbers; 6 frozen in liquid nitrogen (GZ0528). sible that one or more of these forms can be NSW: Bare Rock, Botany Bay, 20.12.1998; numerous referred to B. chuni (von Lendenfeld, 1885a), the examined. Tas.: Photograph by Karen Gowlett-Holmes, Waterfall Bay (Fig. 3A); specimen not preserved. only Bolinopsis previously described from Austra- Transparent body with magenta canals: SA: Whyalla lian waters, but von Lendenfeld’s description of Marina, 18.02.1999; thousands of specimens 3–8cm, this NSW species is insufficient for confident casually examined in the field; numerous growth determination. stages examined in detail. Kingscote Jetty, Kangaroo A common form throughout Queensland is I., 1.05.1999; numerous, to 12 cm long; very highly completely colourless and transparent (Fig. bioluminescent; preyed upon in large numbers by Cyanea (Scyphozoa). Streaky Bay and Smoky Bay 3B), whereas another small Queensland form jetties, 19.02.2002; numerous. St Francis and Dog Is., lacks lobes and has two small red granular dots Nuyts Arch., Great Australian Bight, 21–25.02.2002; along each side, and a common form in South

FIG. 3. A–B, Bolinopsis spp. A. Bolinopsis sp. A (Tasmania). B. Bolinopsis sp. B (North Queensland). Note aboral extensions in A, lacking in B; complexly winding radial canals in the lobes of A compared to B; and comparatively heavier, longer lobes in B than in A. Both specimens photographed live and orientated with oral end toward upper right. C–D, Bolinopsis ashleyi sp. nov., holotype (Moreton Bay) in life. C. Sagittal view. D. Tentacular view. E–F, Ocyropsis spp. E. Ocyropsis vance, gen. et sp. nov., holotype (Tasmania) in life, orientated oral end toward lower right. Note deeply indented hourglass-shaped stomach, indicating affinity with the O. maculata species group. F. Ocyropsis sp., in life (Cairns Harbour, Far North Queensland). Note ctene rows longer but with comb plates more sparsely arranged than O. vance. G-H, Leucothea spp. G. Leucothea sp. (southeastern Tasmania). H. Leucothea sp. (Moreton Bay). Both G and H are orientated with oral end toward the left. Photographs A, E, and G by Karen Gowlett-Holmes; used with permission. Photographs C, D by Ross Easton (UnderWater World, Sunshine Coast, Australia); used with permission. Photograph H by Puk Petersen (UnderWater World, Sunshine Coast, Australia); used with permission.

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 23 Gershwin, Zeidler & Davie

Australian waters has a fine but conspicuous the body, have a similar functional biology and bright magenta colouration to the canals. predation potential (Main 1928; Nagabhushanam Attempts to preserve this third form ended in 1959; Reeve et al. 1978; Kremer 1979; Schulze- the animals writhing to the point of exploding Röbbecke 1984; Kasuya et al. 1994; Costello & and disintegrating. Experiments to relax the Coverdale 1998). For example, Bolinopsis vitrea specimens prior to fixation included magne - (Agassiz, 1860), which is similar to the Spencer sium chloride, menthol, extremely dilute forma - Gulf Bolinopsis morphologically, was demonstrat- lin, dilute ethanol, cooling, freezing, warming, ed by Kremer et al. (1986) to be a more efficient tea tree oil, eucalyptus oil, oxygen deprivation, predator than Mnemiopsis. Similarly, according and seawater diluted 50% with tap water. All to Mills (2001: 65), ‘Uye & Kasuya (1999) relaxing efforts proved unsuccessful; however, suggest that numbers of indigenous cteno - the dilute ethanol and the tap water dilution phores, especially Bolinopsis mikado (Moser, both produced a temporary specimen. Specimens 1907), may be rising in some Japanese coastal killed in dilute ethanol lasted a few days but waters; this situation bears following in coming exploded with further preservation attempts; years.’ We assert, therefore, that Bolinopsis in specimens killed with tap water dilution died Spencer Gulf could pose a similar ecological quickly in a good form, but disintegrated threat to that of Mnemiopsis in the Black Sea. quickly if left untreated, and also exploded in response to other treatments. Bolinopsis ashleyi sp. nov. Another unidentified form was found once in abundance off Palm Cove, Cairns, Queensland, (Fig. 3C–D) in about 1–2 m water. It had peculiar double Material examined. HOLOTYPE: SAM-XH450, approx. round marks midway inside the body. It was 10 cm total live length, 1 km off Mooloolaba, Qld, coll. P. found along with Pukia falcata sp. nov. (this Petersen, M. Callaghan, and M. Rego, October, 1999. paper), during a time of low medusa diversity. Preserved in 100% ethanol. Non-type material. Video and photographs of The Bolinopsis species found in the gulfs of several additional specimens (kept in the South South Australia warrants further study. It appears Australian and Queensland Museums). that it may pose a nuisance in the semi-enclosed Diagnosis. Bolinopsis with large, hemispherical waters of Spencer Gulf where blooms in plague lobes; brilliant red pigmentation along most of proportions have been observed (unpublished the length of the ctene rows. data, and correspondence with fishermen and researchers). While the trophic dynamics of Description. Total length of specimens known Bolinopsis in Spencer Gulf are unknown, the to approximately 12 cm, slightly compressed in predatory impact of this species is of concern, the tentacular plane. Aboral end of the body and should receive priority in sustainability bluntly rounded, with mesogleal extensions studies of commercially important species that slightly protruding past the aboral cleft. Lobes depend on the Spencer Gulf, and Gulf St. Vincent, very broadly rounded, spanning approx. 4–5 as a nursery or feeding ground. times the body width in the sagittal plane. Coelenterates, and ctenophores in particular, Auricles narrow, ribbon-like, tapered distally, are very successful competitors, preying on approximately 1/2 body length, reaching well eggs and larvae of other animals, and also beyond the mouth. competing with them for food resources Ctene rows 8, wide, all with densely-packed (Purcell 1990; Purcell 1991; Purcell & Arai 2001). comb plates. The four subtentacular comb rows, A well documented example is Mnemiopsis, which with about 60 comb plates, extend the entire was accidentally introduced into the Black Sea distance from the apical pole to immediately in 1982, and by about 1990, was estimated to short of the auricles; the four subsagittal rows, have a biomassover of a billion tons, and had with well over 100 comb plates, almost touch led to the collapse of fisheries industries in the where they begin slightly inward of the apical region (Shiganova 1998; Kideys 2002). pole, and extend nearly to edges of the lobes, Bolinopsis and Mnemiopsis, although separa - flaring a short distance along the path of the ted based on the length of the lobes relative to meridional canals before termination.

24 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

At approximately 4/5 of the distance to the diet of small fish, adult and newly hatched outer edge of the lobe, the meridional canal Artemia, and unidentified mysid shrimp. They continues from the distal end of each were also offered dead fish, but did not take subsagittal ctene row, departs laterally, makes them. They survived approximately three a broad wavy crescent following the contour of months in captivity. Puk Petersen observed, the lobe toward the central axis of the animal, ‘We saw a definite reaction to light; as soon as then very near to the base of the lobe the canal we took the covers off for feeding they would loops back upon itself to the inside, once again locomote towards the surface. When feeding, following the contour of the lobe in a smaller, the flaps would open wide like a feeding whale somewhat wavy crescent, back to the axis of the shark, we would gently touch them on the ctene row, then dips inward toward the mouth, inside of the oral lobe with the fish and they then abruptly changes direction to unite with would instantly close and grab it, the fish was its mirror image in a finger-like, distal-pointing immobilised very quickly. At 18° it took a 10 cm projection at the midline between the two ctene animal about 2 hours to digest a 1 cm fish rows. The circumferential canal is not wavy, (approximately) .... I don’t think they intently and closely mirrors the contour of the outer hunted for a particular item of food as much as edge of the lobe along nearly its entire distance. hunting in general with the oral lobes wider Statocyst situated within deep aboral cleft. open than when they weren’t feeding.’ Stomodaeum reverse hourglass-shaped, widest Field identification. Like other Bolinopsis (see immediately oral to the midpoint, tapered both earlier), but with massive lobes, and red aborally and orally, though wider in the oral pigmentation along the ctene rows. half than the aboral half. Remarks. Although we have designated a Body colourless and transparent, except for a holotype, as mandated by Article 72.3 of the brilliant red band of pigment beneath the distal I.C.Z.N. (1999), we must acknowledge that this 2/3 of the 4 subsagittal comb rows and out specimen will of limited use for morphologi- along most of the outer crescent of the lobe cally discriminating this new species. Unfortun - canals. This same colour-pattern and vibrancy ately it is impossible to adequately preserve has been observed in approximately 40 speci - many types of ctenophore, including Bolinopsis mens of all sizes from 3 to 12 cm. ashleyi. Others have also attempted to deal with Type locality. About 1 km offshore, Mooloo- this problem including: Harbison & Miller laba, Sunshine Coast, Queensland. (1986), who deposited specimens of which only gonads and comb row fragments remained; Etymology. Named in honour of Ashley Robillard & Dayton (1972), who embedded Scivyer, formerly of Underwater World on the their specimens in wax; and Matsumoto & Sunshine Coast. Ashley and his wife, Puk Robison (1992), who preserved their specimens Petersen, have been extremely dedicated in in 4% buffered gluteraldehyde. However, discovering the medusa and ctenophore scattered ctenes and wax-embedded specimens biodiversity of Queensland and other tropical are unsuitable for study, and gluteraldehyde is regions of the world. problematical and only a temporary solution; Distribution. Presently known from Mooloolaba, ultimately, these species are based on photo- south to Moreton Bay. The material mentioned graphic evidence. While no method has so far was all caught within 2 m of the surface, been found to preserve these delicate taxa, in between October 1999 and about March 2000. fact, Matsumoto & Robison (1992: 20), have Bioluminescence. Could not be elicited with stated that morphological inform ation obtained tapping on the tank, but direct stimulation was from in situ observations, photographs and not attempted. video, ‘Often ... is far superior to that which can Ecological notes. Not observed during times of be inferred from preserved material.’ heavy salp blooms. Several specimens were The necessity of having a deposited holotype captured with small (1–1.5 cm long) fish in the specimen for a new species has also been gut. In captivity they were fed a maintenance recently questioned by Donegan (2008); and

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neknus neknus neknus neknus ylpeeD ylpeeD e s sisponiloB n e mc S gnol sa eciwt tuoba VS tuoba eciwt sa gnol otno yawflah naht ssel naht yawflah otno suoremun setalp bmoc setalp suoremun ,gnol yrev lartnevbuS yrev ,gnol gnidnetxe swor bmoC swor gnidnetxe )sebol otno gnidnetxe otno )sebol otno gnidnetxe yleraB gnidnetxe otno no 81–51( setalp bmoc setalp 81–51( no ylerab ,.e.i( htuom eht htuom ,.e.i( ylerab gnidnetxe swor enetC swor gnidnetxe SS no setalp 22-02 ,TS 22-02 setalp no SS suneg eht ni seiceps fo sretcarahc citsongaid fo nosirapmoC nosirapmoC fo citsongaid sretcarahc fo seiceps ni eht suneg no setalp 81-61 ;sebol 81-61 setalp no sebol otno gnidnetxe otno sebol no 53–03 ;swor trohs ;swor 53–03 no ;sebol otno yawflah otno ;sebol fo xepa ot ylraen SS ylraen ot xepa fo fo level ot VS ;TS sa ;TS VS ot level fo fo nigram ot ylraen ot nigram fo degnarra ylesrapS degnarra fo esab ot ylno SS ylno ot esab fo ro debircsed toN debircsed ro fo level ot ,sebol ot level fo sebol no lanac no sebol ton lartnevbuS ton debircsed toN debircsed

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26 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia napaJ ,oteS napaJ ,tseW yeK ,tseW ailartsuA nrehtuoS nrehtuoS laciporT ytilacoL dnalaeZ adirolF snriaC gnireB noiger tiartS weN dlQ dlQ yltcefrep ylraeN yltcefrep dnuof yllareneg dnuof ,enif dna llams dna ,enif elbuod ;slanac elbuod ediw selciruA ediw sesab elcatnet sesab elbisiv ylerab elbisiv ;nommocnU ;depahs-gge ,depahs-gge slessev eboL slessev notknalpooz srahc rehtO srahc yrev sbmoC yrev bmoc kcihT bmoc lanoidirem cirtsagarap esned htiw esned suonitaleg nioj slanac nioj ylemertxe ylemertxe dehcnarB etagnolE nommoc nommoc thgiarts moolb swor nworb-der peed nworb-der yrehpirep gnola yrehpirep ycnerapsnart sti ycnerapsnart slanac atnegaM slanac hcae gnola stod gnola hcae sselruoloc dna sselruoloc owt htiw ydob htiw owt ni elbakrameR ni hsidder slanac hsidder deruoloc-eulb ;ebol hcae fo hcae ;ebol stops egnaro stops -hsidder 6–4 -hsidder tnerapsnarT der ralunarg der ydob fo edis fo ydob eht gnikram eht slanac niam slanac ;der slanaC ;der yletelpmoC detnemgip sselruoloc tnemgip nosmirC ruoloC hcamots thgiL tfelc peed nI peed tfelc nagro esneS nagro ylpeed yreV ylpeed yletaredoM wollahS neknus neknus neknus neknus neknus neknus ylpeeD ylpeeD ylpeeD ylpeeD sa x2 tuoba lartnevbuS tuoba x2 sa ;ralucatnetbus sa gnol sa ;ralucatnetbus ;sebol otno gnidnetxe otno ;sebol htuom dnoyeb dnetxe dnoyeb htuom gnidnetxe lartnevbuS gnidnetxe gnidnetxe lartnevbuS gnidnetxe gnidnetxe lartnevbuS gnidnetxe yltnatsid setalp bmoc setalp yltnatsid ;swor trohs no 06–05 no trohs ;swor gnol sa x5.1 .c ,sebol .c x5.1 sa gnol ylthgils lartnevbuS ylthgils yawflah naht erom naht yawflah ton od lartnevbus od ton swor gnol no 021 no gnol swor fo egde ot ylraen ot egde fo fo egde ot ylraen ot egde fo fo egde ot ylraen ot egde fo ralucatnetbus sa ralucatnetbus ton swor enetC swor ton ;ralucatnetbus ralucatnetbus ralucatnetbus

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mm 05 mm detats y toN d mm mm o B ydob flah ot flah ydob lauqe htdiW lauqe )htgnel latot )htgnel trohs ,llamS trohs gnol si ydob si gnol tub ,worran tub ssel ylthgils ssel htgnel ydob htgnel trohs yreV trohs etaredoM 5/3( egraL 5/3( naht erom naht e latot flah latot ro htgnel ro sa daorb sa z sa tuoba sa gnikcaL i ,dnuor s ,eguH

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Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 27 Gershwin, Zeidler & Davie while his argument against this process was 1850), has short comb rows that barely extend primarily based on ethical and conservation onto the lobes, and also a more complex form of issues, he also emphasised that superior inform- lobe canals (see Agassiz, 1860: figs 88, 89). ation to assist identification can frequently be Several other species are said to contain red secured from non-corpse material. Our descript - pigment. First, Mayer (1912) stated that Bolinopsis ion of Bolinopsis ashleyi sp. nov. is thus primarily vitrea sometimes displays intense pink colour based on the excellent photographs that clearly in the peripheral parts of the chymiferous illustrate the morphology of this species. If a tubes, probably due to products of digestion or suitable method of preservation is developed, excretion. This is unlike B. ashleyi, in which the designation of a neotype of B. ashleyi (as well as pigment granules are epithelial, not digestive. its congeners!) would probably be desirable. Second, like B. ashleyi, B. paragaster has red The most striking feature of Bolinopsis ashleyi canals, but it also has a deep red-brown pig - is its brilliant colouration. While colour is often mented stomach, bifurcated paragastric canals, regarded as of little taxonomic value because it and double tentacle bases. Third, B. mikado cannot be studied in preserved specimens, (Moser, 1907) is said to be ‘transparent and many previous ctenophore workers have also almost colourless; only the canals are rosy used colouration (Mayer 1912; Bigelow 1912; when living’ (Komai, 1918: 455); however, there Tokioka 1964; Harbison & Miller 1986; Mills is another form of Bolinopsis in southern 1987; Matsumoto 1988; Matsumoto & Gowlett- Australian waters which has rosy canals, but Holmes 1996; Matsumoto 1999). In the case of would not be easily mistaken for B. ashleyi, in ctenophores, which often cannot be preserved which the red pigment might be more at all, colour remains an invaluable tool to be accurately likened to ‘racing stripes’. used in combination with other characters. Family Leucotheidae Krumbach, 1925 Bolinopsis ashleyi differs from its congeners by the following (see also Table 5). Its broad, Leucothea Mertens, 1833 nearly hemispherical lobes, separate it from the Leucothea filmersankeyi sp. nov. narrow lobed species: Bolinopsis microptera (Agassiz, (Fig. 4A–D) 1865), is more elongated, has very short lateral Leucothea sp. — Gershwin & Zeidler, 2003: 238 (Tas, lobes, has complexly winding canals, and is SA). transparent and colourless; Bolinopsis indosinensis Material examined. HOLOTYPE: QVM-20:4196, Stanley Dawydoff, 1946, has small, short lobes and Jetty, Stanley, Tasmania [40 46’02.4"S, 145 18’19.1"E], large, broad auricles; and Bolinopsis ovalis 8.03.2009, at surface in 1 m of water, observed for 3 (Bigelow, 1904), has sparsely-arranged comb days in captivity then preserved in EtOH; 10 cm TL plates and is colourless. live. PARATYPE: QVM-20:4197, same coll. data as holotype; 13 cm TL live. Of the species with large lobes, Bolinopsis Non-type material. TAS: Same coll. data as holotype; ashleyi can be distinguished by the following. 2 specs, 8–10 cm TL, observed in captivity for 2 days Bolinopsis rubripunctata Tokioka, 1964, is about before spontaneous disintegration occurred. Same half the size, and has numerous conspicuous coll. data as holotype; 7 specs, 7–10 cm TL, observed reddish orange spots along the periphery of the in the field and released. Stanley Jetty, 3.09.2002 lobes. Bolinopsis infundibulum (Müller, 1776), (GZ0210); one spec. torn beyond specific recognition, has much shorter subsagittal comb rows, and appeared to be approximately 15 cm TL; some tissue retained in 100% ethanol [SAM-XH445], some in rows of dark spots on the lobes, but lacks any liquid nitrogen [SAM-ABTC101331]. SA (probably trace of the brilliant red colouration of B. ashleyi. identical): North Point, St Francis I. [32°29’33.9”S, Bolinopsis chuni (von Lendenfeld, 1885a), has 133°16’59.6”E], S. Murray-Jones, 23.02.2002 (GZ0549); much shorter and less-dense subsagittal comb fragmentary specimen in alcohol [SAM-XH449] and rows that are violet in the adult. Bolinopsis liquid nitrogen. Fenelon I., Nuyts Archipelago [32 elegans (Mertens, 1833), is covered with tubercles. 34.474’S, 133 17.550’E], L. Gershwin, 25.02.2002 (GZ0559); 1 spec., about 15 cm TL, examined in the Bolinopsis paragaster Ralph & Kaberry, 1950, has laboratory until disintegration. Several specs were bifurcated paragastric canals and double observed at Masillon I. [32°33.581’S, 133°17.041’E], L. tentacle bases. Finally, Bolinopsis alata (Agassiz, Gershwin, 25.02.2002.

28 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Diagnosis. Leucothea with body to about 15 cm Tubercles prominent, numerous; most densely in length; with long, narrow, blind tentacular arranged and longest on aboral end and along pits; with long, slender papillae; with ctene long body corners, sparsely arranged and rows about 3/4 length of lobes, broad; with shorter on outer surfaces of lobes; slender, bimorphic diverticula beneath ctene rows; with cylindrical, tapered, many reaching nearly 1 cm pale orange tint to core of body, lacking other in length, about 1 mm diameter. colouration. Canals: Infundibular canal short, elongated Description from living specimens. Body hourglass-shaped (Fig. 4D). Meridional canals highly compressed in the tentacular plane, flat straight, heavily diverticulated under comb along both broad sides; approximately 5 times rows, ovaries on opposite sides of adjacent as wide in the stomodael plane. Aboral end of canals. Diverticula bimorphic: those underlying body deeply cavernous, with statocyst further subtentacular comb rows often alternate wide deeply embedded (Fig. 4A). Mouth small, incon- and narrow; those underlying substomodeal spicuous, midway along narrow blade-like oral rows typically of equally broad width. Beyond edge of body, concealed by lobes. Oral tentacles oral termination of substomodael comb rows, not observed. Tentacles 2, each located midway canals diverge at approximately 45º angles, across each broad side, near mouth on oral edge extending in straight line nearly to distal edge of body, emitting from tentacular canals. of lobes (Fig. 4B). Lobe canals winding Tentacular canals very long and narrow, about circuitously, forming large loops and smaller 1.5 mm wide, about 24 mm long, aboral- complex patterns (Fig. 4C), with left and right pointing portion about 3 times as long as oral- not necessarily in mirror image. pointing portion, blind-ending in both direct - Colouration: pale orange in long spindle- ions; oral canal portion contains a linear internal shaped region of body core; lobes and auricles structure of unknown function. transparent ghostly whitish; tubercles mostly Lobes 2, relatively small compared to other transparent and colourless. species, comprising about half the total length Etymology. Named to honour Mr Patrick Filmer- (TL) of the ctenophore (Fig. 4A), attached to Sankey, then Director of the Queen Victoria body such that approximately 1/2 of body and Museum and Art Gallery in Launceston, 3/4 of lobes overlap; with a broad figure-8 Tasmania, where the types are lodged. Patrick’s outline in normal posture, i.e., when lobes are enthusiastic approach to science and gentle - curled in. When extended for feeding, lobes are manly approach to life have been of great broadly rounded in outline, about half as broad inspiration to the senior author. as the body is long, with a deeply incised Field identification. Body covered in narrow, midline notch along distal edge, appearing gelatinous, retractable, flexible tubercles. Body heart-shaped (Fig. 4B). shape very compressed in the tentacular plane, Auricles 4, attached to body somewhat oral to considerably longer than wide, with small point of lobe attachment; long, slender, lobes always held curled; forward margin of cylindrical, gradually tapered; held tightly lobes with deep indentation. Colouration: pale coiled in a helical fashion most of the time (Fig. orange body core, with transparent and colour - 4A), or, when relaxed and extended, movement less tubercles; auricles and lobes whitish. Size by the animal causes ends of auricles to wave range: 6–15 cm long, with most specimens constantly, as if acting as lures. observed about 8–9 cm long. Ctene rows 8, broad, arranged as follows: 4 Distribution. Leucothea filmersankeyi has only substomodaeal ctene rows (SSCR) emanate been confirmed hitherto from Stanley, in the far from aboral end of body, extending to about north-west of Tasmania, along Bass Strait. 3/4 length of lobes, at point of divergence of Samples from Nuyts Archipelago, SA, are also meridional canals toward lobe margin; 4 sub- probably of this species, but more material is tentacular ctene rows (STCR) emanate from needed for positive identification. indented aboral end of body, extending some- Behaviour. Very active in captivity, spending what beyond origin of auricles. most of its time just under the surface of the

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 29 Gershwin, Zeidler & Davie water, hanging in a lobes-up orientation, L. filmersankeyi seems most similar to L. japonica, frequently sinking to the bottom of the tank in that the comb rows extend about 3/4 the then bouncing back to the surface. On contact distance toward the margin of the lobes. with the air-water interface, the lobes spread However, L. filmersankey has no trace of red out somewhat, with no ripple whatsoever on colouration, and does not squirt ink when the surface tension. Sometimes the animal poked, so it would be unlikely to be confused begins to spin on its axis, which it may persist in for L. japonica, which does. doing for several minutes without ascending or In colouration, L. filmersankeyi is quite readily descending from its position. separated from all others, and its size separates The animal is very responsive to mechanical it from many. Unfortunately, most structural stimulation. When touched gently on any side, features are not well described for the existing it will recoil and immediately move away from species of Leucothea, such that it is very difficult the direction of stimulation. When a single to compare across a range of features. tubercle is touched with a plastic probe, all However, logically, conspicuous differences in neighbouring tubercles within about a 1.5 cm colouration may be indicative of other radius shoot out immediately in the direction of differences, which simply cannot be assessed at the probe, and stay extended for several this time. For example, the striking colour seconds. The tubercles in a given area do not pattern of L. pulchra has no counterpart in any respond to continued stimulation, but will other species; the spotted markings in L. respond again if stimulated after another region. ochracea and L. tiedemanni, while different from Although no light-sensory apparatus was each other, are nonetheless not echoed in detected, it does respond to sudden illumination others; and finally, while most Leucothea species by immediately recoiling the body. are yellowish, the presence of red pigment in L. Bioluminescence was tested at night with a japonica may be interpreted as a key difference. simple probe. Small flashes of blue light were We wonder about the relationship between emitted when stimulated along the canals; L. filmersankeyi and a curious form from Little when stimulated near the base, all canals Waterfall Bay in southeastern Tasmania, flashed simultaneously. further discussed in the next section. Both share Remarks. Leucothea filmersankeyi differs from its the interesting characters of an elongate, congeners as summarised in Table 6. Structur - narrow and largely colourless body, and tightly- ally, L. filmersankeyi appears to be unique in h eld lobes. However, upon closer examination, having narrow, blind tentacular pits; the only the two have many differences. For example, other species for which the tentacular pits are the lobes of L. filmersankeyi comprise about half well characterised is L. pulchra, in which they the total length of the ctenophore, whereas are broad. The gonadal regions of the merid - those of the southern form are nearly 2/3 the ional canals are bimorphic in L. filmersankeyi, total length (compare this with the lobate whereas this pattern is undescribed for other family Bolinopsidae, in which a similar species. Furthermore, the papillae are relatively difference is used to separate the genera narrower and longer than those figured for Bolinopsis and Mnemiopsis). In addition, the two other species. For most species, the comb rows Tasmanian forms of Leucothea have about the are described either as extending only halfway same density of tubercles, and they are quite onto the lobes, or as ‘very long’; in this respect, slender in both forms, but the tubercles are

FIG. 4. A–D, Leucothea filmersankeyi sp. nov., holotype. A. Habitus, tentacular view. B. Stomodeal view; note broad heart-shape lobes. C. Detail of lobe canal pattern. D. Detail of sense organ and infundibular canal. E, Velamen sp. (Tasmania), in life. Oral end orientated toward right. F–H, Beroida, all specimens orientated with oral end ‘up’. F, Beroe sp. A (North Queensland); completely transparent and colourless, in life. G, Beroe sp. B (North Queensland), with broad ctene rows and red pigmentation, in life. H, Beroe sp. C (Tasmania), with a pronounced neck lacking ctene rows, preserved. I, Neis cordigera (South Australia), in life. Photograph A by Karen Gowlett-Holmes; used with permission. Photograph E by Thierry Laperousaz (South Australian Museum); used with permission.

30 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 31 Gershwin, Zeidler & Davie .)0881( nuhC sulp ,snoitpircsed lanigiro morf devired ataD . ataD devired morf lanigiro ,snoitpircsed sulp nuhC .)0881( ,ruobraH avuS ,ruobraH naenarretideM napaJ ,ikasiM napaJ ASU ,adirolF ASU AS ,saT WN ,saT AS ,ainrofilaC fo tsae aeS tsae fo ,sagutroT ytilacoL saT ES saT saT ES saT dlQ ES dlQ napaJ ASU ijiF .L eallipap rednels eallipap naht xelpmoc naht ssel raf slanaC raf ssel eallipap egraL eallipap eallipap egraL eallipap rednels ,trohS rednels !dekop nehw !dekop gnol ;eallipap gnol dnilb worraN dnilb s no stnemalif no e sinrocitlum kni striuqS kni stip daorB stip ,gnol ;stip ,gnol t lapicnirp o selcatnet n laretaL

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32 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia quite short in the southern form, and much short, slender papillae; the lobes comprise longer in the northern L. filmersankeyi. The exact much more that half the total length (Fig. 3G). relationship between the two forms remains No specimens were captured. unclear at this point, but it seems plausible that A second form, comparable to L. multicornis they represent different species. It also high- (Quoy & Gaimard, 1824), was caught by Puk lights the need for existing species to be re-exam- Petersen in Moreton Bay in September 2000 ined across a range of structural characters, so (Fig. 3H). Unfortunately, specimens could not that they can be properly compared. be preserved intact. From the photographs, and from first-hand accounts, it can be described as: Leucothea species body length c. 9 cm, with prominent gelatinous (Fig. 3G–H) extensions at the aboral corners; extensively covered by large papillae (c. 5 mm tall); ctene Other species of Leucothea that have been rows wide, running the entire length of the previously reported from Australia: animal; colouration transparent yellowish with Leucothea spp. — Harbison & Miller, 1986: fig. pale orange gut. The auricles were thick, 4 (Coral Sea); Edgar, 1997: 149 (Vic, NSW, Tas.); cylindrical, tapered and long, most often held Edgar, 2000: 149 (Vic, NSW, Tas.). coiled in a conical form, but occasionally Material examined [from photographs]. Qld: SAM- unreeling into long, writhing, tentacle-like XH444, Pumicestone Passage, Moreton Bay, ethanol- appendages. These characters are similar to L. preserved tissues from specimen in Fig. 3H, P. multicornis, described from the Mediterranean Petersen, Sept. 2000; about 9 cm TL. TAS: Little by Quoy & Gaimard (1824) and Chun (1880). Waterfall Bay, Tasman Peninsula, K. Gowlett- Holmes, 2.03.1998; about 12 cm TL. Dog Leg Cave, However, given the different biogeographic Waterfall Bay, M. Baron, 18.11.1998; 30–45 cm TL. provinces it is unlikely that these two Field identification. Body covered in gelatinous, populations will be conspecific. In colouration, retractable, movable tubercles; body usually the Moreton Bay form is different from L. longer than wide, with large or small lobes. grandiformis (Agassiz & Mayer, 1899), which is Colourless and transparent, or yellowy, orangy, cinnamon-yellow on the ctene rows, canals, or greenish; medium-sized or massive. and gastric cavity; from L. ochracea Mayer (1912), which has lateral filaments on the Distribution. Leucothea was reported from tentacles in the adult stage, and also has Victoria to New South Wales, including Tasmania conspicuous opaque yellow areas on the sides by Edgar (1997, 2000). We have also found of the oral lobes; from L. japonica Komai (1918), Leucothea species of uncertain identity in South which has a faintly brick-red body, deeper red Australia (Gershwin & Zeidler 2003) and canals, and yellow-edged lobes; and from L. Queensland (unpublished data). pulchra Matsumoto (1988), which has a very Remarks. Leucothea urgently needs revision. A large body with large lobes, and the body is good (though brief) summary of species was opaque whitish with prominent orange papillae. given by Matsumoto (1988), who recognised six A third large Australian form was videod by species (Table 6). In Australian waters, it seems Mick Baron at Dog Leg Cave, Waterfall Bay, that several additional species may be present, Tasmania on 18.11.1998. While no specimens based on our limited observations of specimens were retained, they were 30–45 cm long, with collected or photographed. However, any large papillae, a yellow-green gut, and formal descriptions beyond those herein will prominent diverticula beneath the ctene rows. require additional specimens with better photographs and/or preservation techniques. Family Ocyropsidae Harbison & Miller, 1986 Notes on the Australian forms are as follows. Ocyropsis Mayer, 1912 A specimen photographed by Karen Gowlett- Ocyropsis species Holmes on 2.03.1998 at Little Waterfall Bay on the Tasman Peninsula is about 12 cm long, and (Fig. 3E–F) is characterised by having a very elongate, The following Ocyropsis species have been totally transparent and colourless body, with previously reported from Australia:

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 33 Gershwin, Zeidler & Davie

Ocyropsis maculata immaculata Harbison & spots and a deeply-indented stomach as reported Miller, 1986: 413–424 (Heron I., Qld). by Harbison & Miller (1986) from Heron I., and Ocyropsis maculata maculata Harbison & 3) a small form from tropical North Queensland Miller, 1986: 413–424 (Heron I., Qld). with a deeply-indented stomach and two small Ocyropsis crystallina crystallina Harbison & black spots near the margin of each lobe. Miller, 1986: 413–424 (Heron I., Qld). The last form is of interest, because it does not Material examined (mostly unpreservable). Qld: seem to match any known species. In colour - Palm Cove, 29.12.1999, during rich bloom of gelatin - ation, it is most similar to O. crystallina guttata, ous zooplankton; many specs, c. 5 mm across, described by Harbison & Miller (1986) with lacking spots, with shallowly indented stomach, small brown to black spots closely associated with ctene rows starting beyond polar plates. Palm with the substomodaeal meridional canals, Cove, 24.01.2000; 2 cm across. Palm Cove, 19.02.2000; in thick jelly bloom. Palm Cove, 22.02.2000; 3 small located at the ends of the gonads where the specs in rich zooplankton bloom. Cairns Harbour, substomodaeal meridional canals make a sharp 25.12.2005, during rich jelly bloom (Fig. 3F). NT: bend to begin their windings on the oral lobes. Stokes Hill Wharf, Darwin Harbour, 6.08.1998; 1 However, in this last form, there are exactly two stout spec. 19°13.021S, 121°15.399E, off Eighty Mile small black spots very near the margin. We Beach, 16.04.2004; numerous c. 1 cm, some with have seen this form from time to time, but it is spots, some without. WA: SAM-XH428 (= GZ0055), unpreservable and short-lived. Port of Dampier, NW side of King Bay, 30.11.2000; 2 specs with 2 black spots on top end of each lobe above gonad; 1 in EtOH, 1 in liquid nitrogen. Ocyropsis vance sp. nov. Field identification. Body resembling the general (Fig. 3E) shape of two hands held together in prayer, i.e., Material Examined. HOLOTYPE: SAM-H935 (PH0322), two large lobes connected at the bottom. The Waterfall Bay, Tasman Peninsula, Tasmania, drifting lobes are massive, and used for clapping in 2–4 m, K.L. Gowlett-Holmes, 28.03.1999. together for rapid escape. Generally transparent Diagnosis. Hermaphroditic Ocyropsis lacking and colourless, may also have two large diffuse tentacular bulbs or canals, pigment spots, and brown spots, or two small black spots on each warts; with 26–27 ctenes per subtentacular row, lobe. Most specimens are small, but may reach 53–54 ctenes per substomodaeal row; with up to about 8 cm across. deeply indented hourglass-shaped stomodaeum; Distribution. In Australia, the species O. maculata with long substomodaeal ctene rows, origin - immaculata, O. maculata maculata, and O. crystallina ating well before polar plates and extending to crystallina were reported at Heron I., Qld, by the level of the stomodael indentation; with Harbison & Miller (1986). gonads in diverticula on both sides of the meridional canals, ovaries facing one another, Ocyropsis of uncertain affinity were also found with testes projecting in opposite direction. at Palm Cove, Qld, in the summer of 1999–2000. Found in abundance, the largest specimen was Description. Body 29.8 mm (aboral pole to never more than about 1 cm. They were active mouth); 47.7 mm total length preserved (aboral swimmers, and readily luminesced at night pole to lobe tip), lacking warts or pigmentation. when gently stimulated. Despite great effort, Tentacular canals and bulbs absent. specimens were impossible to preserve. Lobes large, glove-like around mouth in Remarks. Like most other ctenophores, Ocyropsis preserved specimen; 41.3 mm long. Lobe are difficult to preserve. The genus needs formal musculature densely crowded, in distinct pattern revision, but we tentatively recognise eight of approx. every second longitudinal fibre heavier species (Table 7), including O. vance sp. nov. than those between; cross-fibres minute. In addition to the new species, there seems to Auricles 15.0 mm long, nearly reaching level be at least three other species/forms of of mouth distally; long narrow triangles with Ocyropsis in Australian waters: 1) a small, straight edges. transparent form with a shallowly indented Subtentacular ctene rows proximally joining stomach and no spots (? = O. crystallina; Fig. 3F); at centerpoint of aboral ‘edge’ of body; distal 2) Ocyropsis maculata with large diffuse brown terminus at level of lobe origination; with 26–27

34 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia ctenes, arched obliquely over raised row. the polar plates, and in lacking spots; however, Substomodaeal ctene rows parallel along O. vance lacks tentacular canals and bulbs, as in aboral side; proximally well beyond polar O. crystallina. Primary characters of the species plates but not adjoining; distal terminus of Ocyropsis are summarised in Table 7. approximately half again longer than subten - Mayer (1900) described ‘wart-like protuber - tacular rows; with 53–54 ctenes, mostly perpen- ances upon the surface of the ctenophore’ for O. dicular over raised rows. crystallina (legend, pl. 31), but in fig. 105 (pl. 31) Stomodaeum 26.3 mm long, deeply indented these appear to be the internal gonads. Fewkes hourglass-shaped, 13.3 mm wide at mouth, 13.5 (1882) described Ocyropsis from the Tortugas mm at widest part of base, 3.1 mm wide at two decades earlier, but did not note surface constriction; mouth approx. 2/3 distance to tips warts. Harbison & Miller (1986, fig. 1b, p. 415) of lobes. also show what might be mistaken for Canals underlying ctene plates diverticul - protruding gonads in their O. maculata ated, but appear to lack gonadal material. Lobe immaculata, but these appear to be encased canals complexly looped. Paragastric canals fine, within a nearly-invisible envelope. If Mayer’s narrow, long. Tortugas form does indeed have surface warts, Gonads in diverticula on both sides of the this would separate it readily from the substomodaeal meridional canals, apparently Australian O. vance. confined to lobe regions distal to ctene Despite Ocyropsis being notoriously difficult terminus; diverticulations alternate and opposite, to preserve, the holotype is in excellent with ovaries in diverticula facing adjacent condition (preserved without special effort in canals, testes facing away; pattern irregular, 10% formalin/propylene glycol in seawater); with those toward outer edge of lobe simple whether there is any structural or physiological and short, approximately 1/3 to 1/2 as long as feature behind this should be investigated those toward center axis of lobe, which are further. Ocyropsis vance is a cold-temperate somewhat branched. species whereas all other ocyropsids inhabit Statocyst in shallow aboral cavity about half warm, tropical waters. depth to level of subtentacular rows; polar ORDER CESTIDA Gegenbaur, 1856 plates each 9.6 mm long, very narrow. Colour in life: transparent and colourless. Family Cestidae Gegenbaur, 1856 Etymology. The specific name ‘vance’ is derived Velamen Krumbach, 1925 from a Category 5 cyclone that destroyed the town of Exmouth in Western Australia in the Velamen species autumn of 1999 (while the senior author (Fig. 4E) happened to be visiting). After wiping out Exmouth, the cyclone then went on to travel Only one Velamen species has been previ- southward across Western Australia, emerging ously reported from Australia: in the Southern Ocean, and never losing Velamen parallelum (Fol, 1869) — Gowlett- cyclone-rating the whole journey over nearly Holmes, 2008: 54 (Australia-wide). 1,000 km of land. Upon reaching the Southern Ocean, it regathered its strength and moved Material examined. Qld: Palm Cove, Cairns District, L. Gershwin, summer of 1999–2000; several specs, c. east towards South Australia and Victoria. Like 1 cm BL, caught during routine plankton tows, in Cyclone Vance, Ocyropsis vance seems nearly cooler water believed to be of an oceanic current, indestructible. Noun in apposition. with a rich diversity of other gelatinous Remarks. Ocyropsis vance is most similar in zooplankton. Tas: SAM-XH448 (=GZ 0128), Bicheno overall appearance to O. maculata immaculata [41 52’24.5”S, 148 18’39.2”E], W. Zeidler and L. Gershwin, 25.01.2002; 2 spec. approximately 3mm Harbison & Miller, 1986, in having a deeply BL, tissues preserved in EtOH. Photograph K. indented, hourglass-shaped stomodaeum, gonads Gowlett-Holmes, south of The Thumbs, Tasman on both sides of the meridional canals and the Peninsula, 4.03.1998, c. 14 C; c. 9 cm, no spec. comb rows originating well before the end of retained.

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 35 Gershwin, Zeidler & Davie dna ,lairetam detcelloc ylwen ,snoitpircsed lanigiro morf detalloc noitamrofnI . noitamrofnI detalloc morf lanigiro ,snoitpircsed ylwen detcelloc ,lairetam dna ,aeS ossagraS ,aeS ,aeS ossagraS ,aeS edreV epaC edreV ,citnaltA N ,citnaltA ,citnaltA N ,citnaltA ;lairotauqE citnaltA N citnaltA etarepmet ainamsaT ,nozamA ,nozamA sevidlaM nrehtuoS nrehtuoS .I noreH .I noreH .I .I noreH .I ytilacoL sellitnA -dloC epols ton tub ylesrevsnart gib ylesrevsnart tub ton slanac lanoidirem htiw lanoidirem slanac nworb ylmrofinu ydoB ylmrofinu nworb sdanog fo sdne laro ta laro ;tnesba sdne fo slanac sdanog relpmis 2 htiw sebol kciht ,giB kciht sebol htiw 2 stops nworb krad gib krad nworb stops stops ;trohs selciruA ;trohs stops ot nworb llams htiW llams nworb ot renni no stops kcalb stops no renni yrev sebol ;ruoloc ni ;ruoloc sebol yrev sebol laro fo ecafrus fo laro sebol wolley esuffid egraL esuffid wolley laro no stops kcalb stops no laro noitaicossa ni sebol ni noitaicossa hsinworb ot nworb ot hsinworb gnikcal ,sselruoloc gnikcal anillatsyrc .O anillatsyrc s dna tnerapsnarT dna r e t stops gnikcaL stops stops gnikcaL stops c a r a h c rehtO naht stops kciht sa gnol sa eciwt SS eciwt sa gnol sa bmoc ;setalp ralop ;setalp bmoc fo sdne ot ylno SS ylno ot sdne fo fo sdne ot ylno SS ylno ot sdne fo ralop fo dne ot SS ot dne fo ralop ralop fo dne ot SS ot dne fo ralop a ylno htiw ,trohS htiw ylno a bmoc ;tsycotats ot ;tsycotats bmoc tsomla dnetxe SS dnetxe tsomla tsomla dnetxe SS dnetxe tsomla suoremun setalp suoremun esraps setalp esraps ot ylraen ;TS ylraen ot s setalp ralop setalp tsycotats ot tsycotats sbmoc wef sbmoc w tsycotats o r

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36 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

Diagnosis. Ribbon-like ctenophore, with the CLASS NUDA Chun 1879 sides of the body drawn out to the extreme. In ORDER BEROIDA Eschscholtz, 1825 Velamen, the canals arise straight off the axial canal system toward the lobes, whereas in Family Beroidae Eschscholtz, 1825 Cestum, the canals parallel the stomach for a short distance then curve sharply outward Beroe Browne, 1756 along the lobes. Field identification. Body ribbon-like or belt- Beroe species like, with the sides of the body drawn out to the (Fig. 4F–H) extreme. In Velamen, the canals arise straight off The following Beroe species have been the axial canal system toward the lobes, previously reported from Australia: whereas in its close relative Cestum, which Beroe ovale Bosc, 1802 — Quoy & Gaimard, reaches about 1 m in length, the canals parallel 1824: 575 (Sydney). the stomach for a short distance then curve sharply outward along the lobes. Beroe macrostomus Péron & Lesueur, 1808 — Lesson, 1829: 105–106 (south of New Guinea). Remarks. Velamen parallelum (Fol, 1869) was originally described from the Mediterranean Beroe cucumis Fabricius, 1780 — Stiasny, 1931: and tropical Atlantic; it has subsequently been 40 (Watson Bay, Port Jackson, NSW); Edgar, reported from the Indian Ocean (Harbison et al. 1997: 150 (southern Australia WA to NSW, 1978) and off California and the Sea of Cortez in including Tas.); Edgar, 2000: 150 (southern the North Pacific (Stretch 1982). Ralph & Australia WA to NSW, including Tas.). Kaberry (1950) documented the closely related Beroe ovata Bruguière, 1789 — Greenwood, Cestum in cold New Zealand waters. It was 1980: 93 (Moreton Bay); Goy, 1990: 110 (Shark recently reported in Australian waters for the Bay, WA); Steene, 1990: 86 (Lizard I., GBR). first time by Gowlett-Holmes (2008). Beroe spp. — Péron, 1807: 105; WA. – Dakin & The taxonomy of Velamen is in need of Colefax, 1933: 198 (NSW); Dakin & Colefax, reassessment. Currently, most workers consider 1940: 211 (NSW); Hamond, 1971: 27 (Moreton that there is only one species of Velamen, Bay, Qld); Zeidler & Gowlett-Holmes, 1998: namely V. parallelum; the same has often been 117–118 (amphipod association in Port Phillip said for the closely related genus Cestum, Bay, Vic.); Gowlett-Holmes, 2008: 53 (southern although Harbison et al. (1978) thought differ- Australia). ently. With the ctenophores of most of the Beroe “n. sp.” — Dakin & Colefax, 1933: 198, world’s regions poorly studied, not to mention pl. 7, fig. 1 (NSW). the difficulty in their capture and preservation (Harbison et al. 1978), it is plausible that there Material examined. Qld: SAM-H1574, Palm Cove, may be additional species in either or both L. Gershwin, 23.02.2000; 2 spec., 2–3 mm, with red dots on stomach and red lines. Gershwin coll., Palm genera awaiting discovery. Specifically, we Cove, L. Gershwin, 19.12.1999; 2 spec.: 6.88 mm BL believe it is possible that there are species with red granules under combs; 3.53 mm BL with differences yet to be elucidated between our yellowish comb rows. Gershwin coll., Palm Cove, L. cold-water Tasmanian form and the warm- Gershwin, 29.12.1999, during rich bloom of water Mediterranean V. parallelum; thus, we gelatinous zooplankton; several spec., incl. one pink have not identified our form to species, in hope c. 2 cm BL; not preserved. Gershwin coll., Palm Cove, of stimulating further study. Thus, it seems L. Gershwin, 22.01.2000; 6 spec. in rich zooplankton bloom. Gershwin coll., Buchen’s Cove, L. Gershwin, unwise to assign these Australian forms to any 10.02.2000; 3 spec. Gershwin coll., Palm Cove, L. particular species without closer examination Gershwin, 22.02.2000; 2 spec. c. 1 cm BL, plus several and comparison with overseas material, especi - 2–3 mm juvs., in rich jelly bloom. NT: Cullen Bay ally since the Tasmanian specimen was much Marina, Darwin Harbour, K. Gowlett-Holmes, larger than those that we typically see in 18.08.1998; 1 large purple spec., disintegrated prior Australia, although still within the size-range to preservation. SA: SAM-GZ0537, Ceduna, W. Zeidler & L. Gershwin, 20.02.2002; 3 tiny spec. WA: reported for the species (see Harbison & Madin SAM-H1284 (GZ0054), Port of Dampier, NW side of 1982).

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 37 Gershwin, Zeidler & Davie

King Bay, W. Zeidler & L. Gershwin, 30.11.2000; 1 Neis Lesson, 1829 spec. with wide flared mouth. Tas: TMAG K1726, Slopen main beach, Tasman Peninsula, L. Turner, Neis cordigera Lesson, 1829 May 1999, in very shallow water; 1 spec. TMAG K1727, same data as K1726; 6 spec. SAM-H1577, (Fig. 4I) ‘Sloping Main’, Tasman Peninsula, J. Cossum, 29 Neis cordigera Lesson, 1829: 103–104, pl. 16, fig. 2 Apr [no year stated]; 2 spec., 34.72 mm BL and 36.44 (Port Jackson); Lesson, 1843: 97-98 (summary); mm BL. TMAG 602/K32, Maria I. West. QVMAG, von Lendenfeld, 1885b: 968–976 (comprehensive photograph by C. Reid, washed up on beach near redescription); von Lendenfeld, 1885c: 673–682, Musselroe Bay, far NE, 8.05.2009; conspicuous red pl. 33 (detailed description); Whitelegge, 1889: pigment under ctene rows, no specimen retained. 197 (Port Jackson); Stiasny, 1931: 40 (Gunnamatta Vic: SAM-PH0082, Portsea, Port Phillip Bay, Bay, Port Hacking, NSW; reg. no. AM-P12946). 5.07.1993, with commensal amphipods. Material examined. WA: WAM-Z4694, west side of Field identification. Body very soft and mucous, groin, Cockburn Sound Power Boat Association bag-like, typically flattened, with a very large Beach, Cockburn Sound, J. Fromont, 6.03.1999; 33.75 open mouth at one end; usually colourless and mm BL, 37.98 TL, mouth 25.4 mm wide. transparent, but may be faintly coloured WAM-Z4695, South Mole, Fremantle, c. 1 m water yellowish, purplish, or reddish, especially depth, J. Fromont, 8.03.1999; 11 cm BL, 7 cm wide. Gershwin coll., 19°13.021S, 121°15.399E, off Eighty along the ctene rows. Body of some species may Mile Beach, WA, L. Gershwin, 16.04.2004; numerous get quite large, but most specimens are about at sunset only. SA: SAM-H1070, North Arm Marina, 1–6 cm long. Commercial Fishing Harbour, Fisherman’s Wharf, Remarks. Beroe is common in Australian waters. Port River, Adelaide, T. Laperousaz & K. Phillips, There are several easily distinguishable forms, 24.05.2000; 6 spec.: 25.86 mm BL, 19.33 mm BW; 23.74 mm BL, 17.7 mm BW; 33.23 mm BL, 39.4 mm BW; although matching these up to existing species 34.83 mm BL, 33.2 mm BW; 62 mm BL, 46.1 mm BW; has proven difficult. One form from Darwin is 77.22 mm BL, 58.45 mm BW. SAM-XH233, same large, about the size of a man’s fist, faintly collection data as H1070; 3 spec.: 19.4 mm BL, 13.76 purple throughout with bright purple ctene mm BW; 24.46 mm BL, 16.4 mm BW; 17.98 mm BL, rows, with comb rows almost to the mouth, 11.83 mm BW. SAM (GZ0523), Cowell, L. Gershwin with two oval rings of papillae near the apical & T. Laperousaz, 20.02.2002; 1 spec. Qld: Reefworld organ; unfortunately, both specimens disinte- pontoon, Hardy Reef, Whitsundays, Great Barrier Reef Marine Park, 3.11.2008, photographs by Emily grated prior to study. A second form, from Smart / Fantasea Adventure Cruising; 1 specimen, 8 tropical Queensland, always 1–2 cm or less in cm TL (note that this species is said to be sighted body length, is transparent throughout (Fig. every year by Fantasea staff, and reaches up to 15 cm 4F). A third, also from tropical Queensland, long). only about 2 cm long, has numerous bright red Other material. SA: Streaky Bay, 5–7.03.08; approx. pigment granules around quite heavy comb 30–40 individuals per 2 hours each day, varied from rows and at the aboral end of the body, along 10 cm to 20 cm; observed and videotaped by T. with large, branched aboral papillae (Fig. 4G). Laperousaz, SAM; no spec. retained. A fourth, from Tasmania, has a long, narrow Diagnosis. Beroida with pronounced aboral body with flared lips, and short, unbranched lobes; with canal network uninterrupted across aboral papillae along both sides of a prominent the midline. conical cap over the statocyst (Fig. 4H). A fifth, Redescription. Body dimensions of largest speci- also from Tasmania, has a fairly short, stout men (ex SAM-H1070): length along center axis body, with conspicuous red pigment beneath 77.2 mm, total length including wings 96.5 mm; the ctene rows. Two others have been identified width between oral ridges 58.5, between mouth as B. cucumis and B. ovata by Edgar (1997, 2000) corners 52.0 mm; von Lendenfeld (1885b) and Steene (1990), respectively; the former has reported specimens up to 250 mm, with general a more sloping aboral body contour, whereas measurements being 2–2.5 times as long as this is more squared off in the latter. Without broad, and 4–5 times as long as thick. The body doubt, there are additional forms as well, yet to is thickest in the centre region when viewed be elucidated, pending more field work and a from the narrow sides. General shape square to formal review of the genus. slightly rectangular (excluding the wings),

38 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia quite compressed, with distinct large, triangular Several times throughout the first night aboral wings protruding to the posterior on following capture, tapping of the jar was found either side of the statolith and center axis, to produce a single response, approximately 1 x edged by a prominent ridge, creating a 1 mm. The exact point of light production could hollowed-out appearance of the two narrow not be established prior to the death of the sides. Laterally, the oral edge flares into a ridge specimen. positioned along the midpoint of either narrow Biology. At present, nothing is known about side, which ends about 2/3 back between the the reproductive or ecological biology of this ridges of the wings. species. In Western Australia, it was found at Ctene rows along the entire length of the around midday, in about 0.5 m depth, where body; the four peripheral rows running from the water was about 3 m bottom-depth, over a approx. 2 mm either side of the statocyst, along sandy area beside a rocky groin, in windswept the aboral margin, around the edge of the seas; in South Australia, large numbers were wings, and up the side edge to the corner where found in the harbour during the day; von the mouth opens. The four axial rows run from Lendenfeld (1885b) repeatedly found single adjacent to the statocyst, flaring outward specimens accompanying swarms of Bolinopsis slightly, up toward the mouth, ending on the in Sydney Harbour. Its behaviour was not outer portion of the lip. Thus, the peripheral observed in situ in either case. In the laboratory, ctene rows are considerably longer than the specimen number WAM-Z4694 vigorously axial ctene rows. avoided light, attempting to dive into the Mouth large, encompassing the entire leading bottom of the observation dish and remaining edge of the animal, though not flared. Stomach curled until the water level was lowered. vase shaped, largest at the oral end, spanning During observation, the aboral 1/3 of the sides from corner to corner, tapering back to a point of the stomach could be seen repeatedly at statocyst; not extending into the aboral wings. undulating peristaltically within the body. Gonads not observed. Distribution. Subsequent to Lesson’s (1829) Gastrovascular system forms a continuous description from Port Jackson, Neis has been network, i.e., it is not divided side to side. reported several times in the Sydney area: von Canals of two types: some continuous between Lendenfeld (1885b, 1885c) found it at Sydney ctene rows, with anastomoses to adjacent canals, Harbour, and Stiasny (1931) found it at Port to the stomach, and possessing blind branches; Hacking. We here broaden the distribution to others, singly or doubly between continuous include Adelaide, the west coast of South canals, unbranched and ending blindly. Australia, Western Australia from the southern- most Indian Ocean sector to the tropics, and the Aboral statocyst small, embedded shallowly Whitsunday region of the Great Barrier Reef. in a cup-shaped pit in the ‘saddle’ between the Because of its broad distribution as now two large aboral wings. Polar fields could not known, it seems likely that Neis may be found be found. in Northern Territory and Tasmanian waters. Aboral papillae of two types: lateral ones This is the first tropical report of Neis. It has not simple and short; axial ones biforked and yet been found outside Australian waters. approx. double the length of the lateral ones. Field identification. Body similar to Beroe, i.e., Colouration: The SA specimens were slightly flattened, bag-like, with large mouth at one cloudy with carmine red side edges (along end, but with the aboral end extending past the nearly the entire length of the short sides). The statocyst on both sides into lobes. Characterised bodies of the WA specimens were, in contrast, by brilliant red pigmentation, particularly colourless with a slightly cloudy translucency, along the two narrow sides. except for the ctene rows, under each of which Remarks. Although Lesson’s (1829) description runs a line of fine red points. was relatively brief, the figure is a perfect Bioluminescence. Bioluminescence was observed rendition of the species, making it immediately in the smaller of the two WA specimens. recognisable. A good account of the morphology

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 39 Gershwin, Zeidler & Davie and behaviour of the species was given by von 7. Body (including lobes) to about 15cm in Lendenfeld (1885b). Neis cordigera remains today length, elongate...... 8 a relatively common ctenophore in temperate — Body about 30-45cm long, with large Australian waters, and may occasionally occur papillae; gut yellow-green...... in large swarms...... Leucothea sp. A There has been debate as to whether Neis is 8. Body overall yellowish, with large papillae merely a Beroe. Chun (1880: 307) regarded Neis (Qld)...... Leucothea sp. B and Beroe to be identical. Von Lendenfeld — Body overall transparent and colourless, (1885b) quite forcefully asserted that Neis is not with small papillae (Tas)...... 9 identical with Beroe, and further remarked that 9. Body densely covered with long, slender Neis seems to represent a transition from Beroe tubercles; lobes 1/2 total length (north- to the Lobata. As elucidated by von Lendenfeld western Tas, SA)...... (p. 969), the vascular system of Beroe is divided ...... Leucothea filmersankeyi sp. nov. in the two halves of the body, whereas no such — Body sparsely covered with short, slender separation is present in Neis. tubercles; lobes approximately 2/3 total length (south-eastern Tas)...... Leucothea sp. C KEY TO THE CTENOPHORES OF 10. Lobes one-half total animal length or less. AUSTRALIA [Bolinopsis spp.] ...... 11 1. Body globular or ribbon-like, gelatinous — Lobes massively long, much greater than and planktonic...... 2 one half total animal length; animal moves — Body flatworm-like, creeping on algae, with a hand-clapping motion when disturbed. echinoderms, or other substrate. Benthic [Ocyropsis spp.] ...... 13 ctenophores...... 18 11. Lobes small, giving the whole animal a 2. Body more or less spherical or egg-shaped, compact egg-shape appearance; pigment with or without lobes...... 3 lacking or inconspicuous (e.g., fine canals — Body highly compressed, and sac-like or coloured, but not body)...... 12 ribbon-like...... 16 — Lobes broad and wing-like; with crimson pigment marking the ctene rows...... 3. Body lacking conspicuous lobes; more or ...... Bolinopsis ashleyi sp. nov. less spherical; with two conspicuous retract - able tentacles issuing from cylindrical bulbs 12. Body colourless; canals violet...... between stomach and body wall...... 4 ...... Bolinopsis chuni — Body with conspicuous lobes; more or less — Body colourless; canals finely magenta- egg-shaped; lacking tentacles...... 6 coloured...... Bolinopsis sp. A 4. With two conspicuous tentacles issuing 13. Stomach deeply indented into a strongly from crescentic-shaped bulbs...... pronounced hourglass shape; with or ...... Pukia falcata gen. nov., sp. nov. without spots...... 14 — With two conspicuous tentacles issuing — Stomach not deeply indented, more from cylindrical-shaped bulbs...... 5 shallowly hourglass-shaped, lacking spots...... Ocyropsis crystallina 5. Tentacles with distantly-spaced tentilla, usually held tightly coiled with a ‘beehive’ 14. Body small (c. 1 cm), found in the tropics. 15 appearance, like droplets along the tentacle; — Body large (c. 5 cm long), transparent, mouth protruding...... lacking spots; found in temperate waters...... Euplokamis evansae sp. nov...... Ocyropsis vance sp. nov. — Tentacles with closely-spaced tentilla, 15. Transparent, with large diffuse brownish usually held relaxed out with a comb-like or blackish spots on inner surfaces of lobes. or feather-like appearance...... Ocyropsis maculata ...... Pleurobrachia species — Transparent, with two small black spots 6. Body surface smooth, lacking tubercles. 10 near edge of lobes...... Ocyropsis sp. — Body covered in gelatinous tubercles. 16. Body compressed, sac-like, with large [Leucothea spp.] ...... 7 mouth at one end...... 17

40 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

— Body highly compressed, ribbon-like or information (in alphabetical order): Phil belt-like...... Velamen sp. Alderslade, Peter Arnold, Australian Museum, 17. Body with rounded lobe extensions on Mick Baron, Penny Berents, Steve Cook, CSIRO- ‘corners’ of aboral end; oral end broad and CRIMP, Barb Done, Merrick Ekins, Peter Fenner, square; conspicuous crimson pigment on Jane Fromont, Amber Hansen, Tom Hatley, lobes...... Neis cordigera Chad Hewitt, Karen Gowlett-Holmes, John — Body with smoothly rounded aboral end; Hooper, Thierry Laperousaz, Museum and Art oral end broad or pursed; colourless or Galleries of the Northern Territory, Museum of sometimes purplish or pinkish. . Beroe spp. Tropical Queensland, Puk Petersen, Queensland 18. Benthic ctenophores found on algal or Museum, Reef HQ, University of Queensland, seagrass host...... 19 Ashley Scivyer, South Australian Museum, — Benthic ctenophores found on animal host Tasmanian Museum and Art Gallery, Ian or free-living in tropical waters...... 22 Tibbetts, Liz Turner, Underwater World (Sunshine 19. Benthic ctenophores found on algal host in Coast), Western Australian Museum, and Bette temperate waters...... 20 Willis. We sincerely thank the photographers — Benthic ctenophores found on algal or who generously agreed to let us use some of seagrass host in tropical waters; body their images; photographs not otherwise transparent with numerous green specks, credited were taken by L. Gershwin. Field work, with branched papillae, about 20 mm long. travel and other costs were generously funded ...... Coeloplana reichelti sp. nov. by the Australian Marine Sciences Association, 20. Body colour red, pink, or orange. ....21 Australian Biological Resources Study (ABRS — Body colour green or white, found on Jania grant no. 20045 to LG & WZ; ABRS grant no. sp. (coralline alga) at Rottnest I.; about 207-63 to LG), the Australian-American Fulbright 10mm long...... Coeloplana thomsoni Foundation, Surf Life Saving Australia, Surf 21. Found on Scaberia agardhii (brown alga) in Life Saving Queensland, Australian Rotary, the South Australia; body bright red or orange, Thyne-Reid Foundation, CRC Reef Research, 25 mm long...... Coeloplana scaberiae James Cook University, the Robert W. King — Found on red and green algal hosts in Port Memorial Scholarship Foundation, and the Philip Bay; body dark pink to orange-red Lions Foundation. L.G. and W.Z. both hold with transparent or whitish areas, 10–50 current honorary positions at the South mm long...... Coeloplana willeyi Australian Museum, and the senior author is 22. Found on animal host, e.g., starfish or soft also an honorary at the Museum of Tropical coral...... 23 Queensland. — Free-living on muddy sand on the Great Barrier Reef; body with yellowish-white LITERATURE CITED reticulate pattern and red pigment, about Abbott, J.F. 1902. Preliminary notes on Coeloplana. 36 mm long...... Coeloplana meteoris Annotationes Zoologicae Japonenses 4: 103–108. 23. On Sarcophyton sp. (soft coral) on Great Barrier 1907. The morphology of Coeloplana. Zoologische Reef; body colourless, mottled with ectodermal Jahrbücher, Abteilung für Anatomie und Ontogenie pale reddish-brown blotches, about 25 mm der Tiere Tiere 24: 41–70, pls 8–10. long...... Coeloplana mellosa sp. nov. Agassiz, A. 1865. North American Acaleph . Memoirs — Found on Acanthaster plancii (Crown-of-thorns of the Museum of Comparative Zoölogy at Harvard starfish) on Great Barrier Reef; body College 1(2): 1–234. transparent and colourless, about 5 mm Agassiz, A. & Mayer, A.G. 1899. Acalephs from the long...... Coeloplana sp. A Fiji Islands. Bulletin of the Museum of Comparative Zoology at Harvard University 32(9): 157–189, 17 pls. ACKNOWLEDGEMENTS Agassiz, L. 1850. Contributions to the Natural History of the Acalephae of North America. Part It is with deepest pleasure we thank the II. On the beroid medusae of the shores of organising committee of the Moreton Bay Massachusetts, in their perfect state of develop - Workshop. We are grateful to the following ment. Memoirs of the American Academy of Arts and people and institutions for specimens and Sciences, New Series 4 (2): 313–374, pls 1–8.

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 41 Gershwin, Zeidler & Davie

1860. Contributions to the Natural History of the United 1880. Die ctenophoren des Golfes von Neapel und States of America. vol. III. pt. I. Acalephs in general. der angrenzenden Meeres-abschnitte. Fauna and pt. II. Ctenophoroe. (Little Brown: Boston). Flora des Golfes von Neapel 1: 313 pp., 16 pls. Arnold, P. 1993. First record of ctenophore Coeloplana 1898. Die Ctenophoren der Plankton-expedition. (Lipsius (Benthoplana) meteoris (Thiel, 1968) from Australia. & Tischer: Kiel). Memoirs of the Queensland Museum 33(1): 16. Costello, J.H. & Coverdale, R. 1998. Planktonic feed- Bigelow, H.B. 1904. Medusae from the Maldive ing and evolutionary significance of the lobate Islands. Bulletin of the Museum of Comparative body plan within the Ctenophora. Biological Zoology at Harvard University 39: 245–269. Bulletin (Woods Hole) 195(2): 247–248. 1912. Reports on the scientific results of the Dakin, W.J. & Bennett, I. 1987. Australian Seashores expedition to the eastern tropical Pacific, in (Angus & Robertson: North Ryde, NSW). charge of Alexander Agassiz, by the U.S. Fish Dakin, W.J. & Colefax, A.N. 1933. The marine Commission Steamer Albatross, from October plankton of the coastal waters of New South 1904, to March 1905, Lieutenant Commander L.M. Wales. Proceedings of the Linnean Society of New Garrett, U.S.N., commanding. XXVI. The ctenophores. South Wales 58: 186–222, pl. 7. Bulletin of the Museum of Comparative Zoology at 1940. The plankton of the Australian Waters off New Harvard College 54(12): 369–408. South Wales (The Australasian Medical Publishing Bosc, L.A.G. 1802. Histoire naturelle des vers, Co.: Sydney). contenant leur description et leurs moeurs, avec Davie, P. 1998. Wild Guide to Moreton Bay (Queens- figures dessinées d’après nature. Medusozoa. Pp. land Museum: Brisbane). 130–168. In, Buffon, G.L.L. (Ed.) Histoire Naturelle de Buffon, classee d’apres le system de Linne par Dawydoff, C. 1930a. Sur une nouvelle Coeloplanide R. R. Castel. Vol. 2. (Déterville: Paris). (Coeloplana perrieri n. sp.) provenant du Golfe de Siam. Archives de zoologie expérimentale et générale Bourne, G. 1900. Chapter 7, The Ctenophora. Pp. 70 (Notes et Revue, Numéro 2): 52–54. 1–25. In, Lankester, R.A. (Ed.), A Treatise on Zoology. Part 2 The Porifera and Coelenterata 1930b. Une nouvelle Coeloplanide de la côte sud (Adam and Charles Black: London). d’Annam (Coeloplana agniae nov. sp.). Archives de zoologie expérimentale et générale 70 (Notes et Brown, R.W. 1956. Composition of Scientific Words. Revue, Numéro 3): 83–86. (Smithsonian Institution Press: Washington, DC). 1930c. Coeloplana duboscqui nov. sp., Coeloplanide Browne, P. 1756. The Civil and Natural History of provenant du Golfe de Siam, commensale des Jamaica. (Osborne and Shipton: London). Pennatules. Archives de zoologie expérimentale et Bruguière, M. 1792. Histoire naturelle des vers. 757 pp. générale 70 (Notes et Revue, Numéro 3): 87–90. In, D’Alembert, M.D.E. (Ed.) Encyclopédie Méthodique. 1938. Les Coeloplanides Indochinoises. Archives de Vol. 1. (Chez Panckoucke: Paris). zoologie expérimentale et générale 80: 125–162, pl. 1. Carré, C. & Carré, D. 1991. Ceroctena bicornis, new 1946. Contribution la connaissance des cténophores genus new species of ctenophore (Cydippida, pélagiques des eaux de l’Indochine. Bulletin Biolog- Pleurobrachiidae) from the Mediterranean. Comptes ique de la France et de la Belgique 80(2): 113–170. Rendus de l’Académie des Sciences Serie III Sciences de la Vie 313(12): 559–564. Devanesen, D.W. & Varadarajan, S. 1942. On three new species of Coeloplana found at Krusadai 1993a. Ctenella aurantia, new genus and species of a Island marine biological station, Gulf of Manaar. Mediterranean tentacled ctenophore (Ctenellidae Journal of Madras University 14: 181–188. fam. nov.) without colloblasts and with labial suckers. Canadian Journal of Zoology 71(9): 1804–1810. Donegan, T.M. 2008. New species and subspecies 1993b. Minictena luteola, new genus and new species descriptions do not and should not always require of Mediterranean Cydippida ctenophore with a dead type specimen. Zootaxa 1761: 37–48. five types of colloblasts. Beaufortia 43(10): 168–175. Edgar, G.J. 1997. Australian Marine Life (Reed Books: Carus, J.V. 1863. Handbuch der Zoologie (W. Engel- Kew, Victoria). mann: Leipzig). 2000. Australian Marine Life, Revised Edition (Reed Chun, C. 1879. Die im Golfe von Neapel erschein- New Holland: Sydney). enden Rippenquallen. Mittheilungen aus der Eschscholtz, F. 1825. Bericht über die zoologische Ausbeute Zoologischen Station zu Neapel Tome I (2 Heft): während der Reise von Kronstadt bis St. Peter und 180–217, Pl. 6. Paul. Isis (Oken) XVI: 733–747, Pl. V.

42 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

1829. System der Acalephen. Eine ausführliche Hamond, R. 1971. Some medusae from near Brisbane. Beschreibung aller medusenartigen Strahltiere (F. Search 2(1): 27. Dummlert: Berlin). Harbison, G.R. 1996. Ctenophora. Pp. 101–147. In, Fabricius, O. 1780. Fauna Groenlandica. (Impensis Gasca, R. & Suárez, E. (Eds), Introducción al Estudio Ioannis Gottlob Rothe: Hafniae et Lipsiae). del Zooplancton Marino (ECOSUR: Chetumal, Mexico). Fewkes, J.W. 1882. Notes on acalephes from the Harbison, G.R. & Madin, L.P. 1982. Ctenophora. Pp. Tortugas, with a description of new genera and 707–715, pls 68, 69. In, Parker, S.P. (Ed.), Taxonomy species. Bulletin of the Museum of Comparative and classification of living organisms, Vol. 1. (McGraw- Zoology at Harvard University 9(7): 251–289, 7 pls. Hill: New York). Fleming, J. 1822. The philosophy of zoology, or a general Harbison, G.R., Madin, L.P. & Swanberg, N.R. 1978. view of the structure, functions, and classification of On the natural history and distribution of oceanic animals (Archibald Constable: Edinburgh). ctenophores. Deep-sea Research 25(3): 233–256. Fol, H. 1869. Ein Beitrag zur Anatomie und Entwickelungs- Harbison, G.R. & Miller, R.L. 1986. Not all cteno- geschichte einiger Rippenquallen. Berlin, Medicin - phores are hermaphrodites. Studies on the system- ischen Facultat der Friedrich-Wilhelms Universitat: atics, distribution, sexuality and development of 12 pp., 4 pls. two species of Ocyropsis. Marine Biology 90: Fricke, H.W. 1970. Neue kriechended Ctenophoren 413–424. der Gattung Coeloplana aus Madagascar. Marine International Commission on Zoological Nomenclature. Biology 5: 225–238, 16 pls. 1999. International Code of Zoological Nomenclature Gegenbaur, C. 1856. Studien uber organisation und (International Trust for Zoological Nomenclature: systematik der Ctenophoren. Archiv fur Natur - London). geschichte 22(1): 163–205, pls 7, 8. Kasuya, T., Ishimaru, T. & Murano, M. 1994. Feeding Gershwin, L.-A. & Zeidler, W. 2003. Encounter 2002 characteristics of the lobate ctenophore Bolinopsis expedition to the Isles of St Francis, South Australia: mikado Moser. Bulletin of Plankton Society of Japan Medusae, siphonophores and ctenophores of the 41(1): 57–68. Nuyts Archipelago. Transactions of the Royal Kideys, A.E. 2002. Fall and rise of the Black Sea eco- Society of South Australia 127(2): 205–241. system. Science (Washington DC) 297(5586): 1482–1484. Ghigi, A. 1909. Raccolte planctoniche fatte dalla R. Komai, T. 1918. On ctenophores of the neighborhood Nave “Liguria” nel viaggio di circonnavigazione of Misaki. Annotationes Zoologicae Japonenses 9: del 1903–1905. I. Ctenofori. Pubblicazioni del R. 451–474, 1 pl. Instituto di Studi Superiori Practici e di Perfeziona- 1920. Notes on Coeloplana bocki n. sp. and its mento in Firenze Sezione di Scienze Fisiche e Naturali development. Annotationes Zoologicae Japonenses 9: 2(1): 1–24, pl. 1. 577–584. Gorman, N. 1988. Planktonic Cnidaria and Cteno - Korotneff, A. 1886. Ctenoplana kowalevskii. Zeitschrift phora of Moreton Bay. MS Thesis, Department of für Wissenschaftliche Zoologie 43: 242–251, pl. viii. Zoology. Brisbane, University of Queensland: 82 pp. Kowalevsky, A. 1880. Über Coeloplana metschnikowii. Gowlett-Holmes, K. 2008. A field guide to the marine Zoologischer Anzeiger 3: 140–141. invertebrates of South Australia (Notomares: Tas- mania). Kremer, P. 1979. Predation by the ctenophore Mnemiopsis leidyi in Narragansett Bay, Rhode Goy, J. 1990. Preliminary observations on the zoo- Island. Estuaries 2(2): 97–105. plankton of Shark Bay, Western Australia, with emphasis on medusae and description of a new Kremer, P., Reeve, M.R. & Syms, M.A. 1986. The rhizostome jellyfish (Cnidaria, Scyphozoa). Pp. nutritional ecology of the ctenophore Bolinopsis 107–113. In, Berry, P.F. et al. (Eds), Research in Shark vitrea: comparisons with Mnemiopsis mccradyi from Bay. Report of the France-Australe Bicentenary the same region. Journal of Plankton Research 8(6): Expedition Committee. (WA Museum: Perth). 1197–1208. Greenwood, J.G. 1980. Composition and seasonal Krempf, A. 1920. Sur un Ctenophore planariforme variations of zooplankton populations in Moreton nouveau Coeloplana gonoctena (nov. sp.). Comptes Bay, Queensland. Proceedings of the Royal Society of Rendus Hebdomadaires des Séances de l’Académie des Queensland 91: 85–103. Sciences (Paris) 171: 438–440. Haddock, S.H.D. & Case, J.F. 1995. Not all ctenophores Krumbach, T. 1925. Ctenophora. Pp. 905–995. In, are bioluminescent: Pleurobrachia. Biological Bulletin Kukenthal, W. & Krumbach, T. (Eds), Handbuch (Woods Hole) 189(3): 356–362. der Zoologie (W. de Gruyter: Berlin). Vol. 1.

Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) 43 Gershwin, Zeidler & Davie

1933. Ueber eine kriechende Ctenophore aus dem Mertens, H. 1833. Beobachtungen und untersuchungen Golfe von Suez und ein paar Thesen über die über die beroeartigen akalephen. Mémoires de Architektonik der Rippenquallen. Mitteilungen l’Académie Impériale des Sciences de St. Petersbourg aus dem Zoologischen Museum in Berlin 19: 475–479. Series 6, 2: 479–544, Pls 1–13. Leloup, E. 1938. Siphonophores et ctenophores. Résultats Mills, C.E. 1987. Revised classification of the genus du voyage du S.Y. Belgica en 1897–1898–1899 sous le Euplokamis Chun, 1880 (Ctenophora: Cydippida: commandement de A. de Gerlache de Gomery ... Euplokamidae n. fam.) with a description of the Rapports scientifiques Expedition Antarctique Belge new species Euplokamis dunlapae. Canadian Journal (J.E. Buschmann: Anvers.). of Zoology 65(11): 2661–2668. Lesson, R.P. 1829. Voyage de la Coquille. Zoophytes. (A. 2001. Jellyfish blooms: Are populations increasing Bertrand: Paris). globally in response to changing ocean con- 1836. Mémoire sur la famille des Béroides. Annales ditions? Hydrobiologia 451: 55–68. des Sciences Naturelles, Paris (Zoology), Series 2 5: 2007. Phylum Ctenophora: list of all valid species 235–266 [pls 1–2 referenced, but could not be names. Electronic internet document available at located]. http://faculty.washington.edu/cemills/Ctenoli 1843. Histoire Naturelle des Zoophytes. Acalèphes st.html Published by the author, web page estab- (Librairie Encyclopédique de Roret: Paris). lished March 1998, last updated 13 May 2007. Main, R.J. 1928. Observations on the feeding Mortensen, T. 1912. Ctenophora. Danish Ingolf- mechanism of a ctenophore, Mnemiopsis leidyi. Expedition 5(2): 1–95, pls 1–10, 1 chart. Biological Bulletin (Woods Hole) 55: 69–78. 1927. Papers from Dr. Th. Mortensen’s Pacific Matsumoto, G.I. 1988. A new species of lobate Expedition 1914–1916. XXXIX. Two new cteno - ctenophore, Leucothea pulchra sp. nov., from the phores. Videnskabelige Meddelelser Dansk Natur - California Bight. Journal of Plankton Research 10(2): historisk Forening 83: 277–288. 301–311. 1932. Ctenophora. Report on the Scientific Results of 1999. Coeloplana thomsoni sp. nov., a new benthic the “Michael Sars” North Atlantic Deep Sea Expedition, ctenophore (Ctenophora: Platyctenida: Coeloplani- 1910 3 (Part 2, No. 1): 1–12, 1 pl. dae) from Western Australia. Pp. 385–393. In, Moser, F. 1907. Neues ueber Ctenophoren. II. Walker, D.I. & Wells, F.E. (Eds), The Seagrass Flora Zoologischer Anzeiger 32: 449–454. and Fauna of Rottnest Island, Western Australia. 1909. Die Ctenophoren der Deutsche Sudpolar- (Western Australian Museum: Perth). Expedition 1901–1903. Deutsche Sudpolar- Expedition, Matsumoto, G.I. & Gowlett-Holmes, K.L. 1996. XI. Zoologie III: 116–192, 3 pls. Coeloplana scaberiae sp. nov., a new benthic Müller, O.F. 1776. Zoologiæ Danicæ prodromus, seu ctenophore (Ctenophora: Platyctenida: Coeloplani- animalium Daniæ et Norvegiæ indigenarum characteres, dae) from South Australia. Records of the South nomina, et synonyma imprimis popularium (Impensia Australian Museum 29(1): 33–40. auctoris, typis Hallageriis: Havniae). Matsumoto, G.I. & Robison, B.H. 1992. Kiyohimea Nagabhushanam, A.K. 1959. Feeding of a cteno- usagi sp. nov., a new lobate ctenophore from the phore, Bolinopsis infundibulum (O.F. Muller). Nature, Monterey Submarine Canyon. Bulletin of Marine London 184 (4689): 829. Science 5(1): 19–29. Péron, F. 1807. Voyage de découvertes aux terres Mayer, A.G. 1900. Some medusae from Tortugas, Australes, exécuté par ordre de sa Majeste l’Empereur Florida. Bulletin of the Museum of Comparative et Roi, sur les Corvettes le Geographe, le Naturaliste, et Zoology at Harvard College 37(2): 13–82, 44 pls. la Goelette le Casuarina, pendant les années 1800, 1912. Ctenophores of the Atlantic coast of North America 1801, 1802, 1803 et 1804. Historique: Tome 1 (de (Carnegie Inst. Washington Publ.: Washington D.C.). l’Imprimérie Impériale: Paris). McCrady, J. 1859a. Notes from the Meeting of the Péron, F. & Lesueur, C.A. 1810. Histoire de la famille Elliott Society, September 1st, 1857. Proceedings of des mollusques ptéropodes. Annales du Muséum the Elliott Society of Natural History of Charleston, d’histoire naturelle, Paris 15: 57–69, pl. 2, fig. 16. South Carolina 1: 239–240. Purcell, J.E. 1990. Soft-bodied zooplankton predators 1859b. On the development of two species of and competitors of larval herring (Clupea harengus Ctenophora found in Charleston Harbor. Proceed - pallasi) at herring spawning grounds in British ings of the Elliott Society of Natural History of Columbia [Canada]. Canadian Journal of Fisheries Charleston, South Carolina 1: 254–271, pl. 14. and Aquatic Sciences 47(3): 505–515.

44 Memoirs of the Queensland Museum — Nature Ÿ 2010 Ÿ 54(3) Ctenophores of Australia

1991. A review of cnidarians and ctenophores feed- Stephenson, T.A., Stephenson, A., Tandy, G. & ing on competitors in the plankton. Hydrobiologia Spender, M. 1931. The structure and ecology of 216/217: 335–342. Low Isles and other reefs. Scientific Reports of the Purcell, J.E. & Arai, M.N. 2001. Interactions of Great Barrier Reef Expedition 3: 17–112, 27 pls. pelagic cnidarians and ctenophores with fish: a Stiasny, G. 1931. Über einige coelenterata von Austra- review. Hydrobiologia 451: 27–44. lien. Zoölogische Mededelingen 14(1–2): 27–42. Quoy, J.R.C. & Gaimard, J.P. 1824. Voyage autour du Stretch, J.J. 1982. Observations on the abundance and monde ... executé sur l’Uranie et la Physicienne, feeding behavior of the cestid ctenophore, Velamen pendant ... 1817–20. Zoologie. 12 pp., plus Atlas. parallelum. Bulletin of Marine Science 32: 796–799. In, Freycinet, M.L.D. (Ed.) Voyage de l’Uranie (Paris). Tanaka, H. 1932. Coeloplana echinicola n. sp. Memoirs Ralph, P.M. 1950. Ctenophores from the waters of of the College of Science, University of Kyoto, Series B Cook Strait and Wellington Harbour. Transactions 7: 247–250, 1 pl. of the Royal Society of New Zealand 78(1): 70–82. Thiel, H. 1968. Coeloplana meteroris nov. spec. (Cteno - Ralph, P.M. & Kaberry, C. 1950. New Zealand Coelen - phora, Platyctenea): Beschreibung und systemat- terates: Ctenophores from Cook Strait. Zoology ische stellung mit einem vergleich der gastrovascul - Publications from Victoria University College 3: 1–11. arsysteme in dieser ordnung. Sonderdruck aus Rang, P.C.A.L. 1828a. Etablissement de la famille des “Meteor” Forschungsergebnisse, D 3: 1–13. Beroides dans l’ordre des Acalèphes libres, et Tokioka, T. 1964. Bolinopsis rubripunctata n. sp., a new description de deux genres nouveaus qui lui lobatean ctenophore from Seto. Publications of the appartiennent. Mémoires de la Société d’Histoire Seto Marine Biological Laboratory 12(1): 93–99. naturelle Paris 4: 166–173, pls 19, 20. Utinomi, H. 1963. Coeloplana komaii, a new creeping 1828b. Description d’un nouveau genre de la classe ctenophore from Sagami Bay. Japanese Journal of des acalèphes. Bulletin d’Histoire naturelle de la Zoology 14(1): 15–19. Société linnéenne de Bordeaux 1: 314–319. Uye, S.-I. & Kasuya, T. 1999. Functional roles of Reeve, M.R., Walter, M.A. & Ikeda, T. 1978. Labora- ctenophores in the marine coastal ecosystem. Pp. tory studies of ingestion and food utilization in 57–76 (Japanese with English Abstract) In, Okutani, lobate and tentaculate ctenophores. Limnology and T. et al. (Eds), Update Progress in Aquatic Invertebrate Oceanography 23(4): 740–751. Zoology (Tokai University Press: Tokyo) [not seen]. Robilliard, G.A. & Dayton, P.K. 1972. A new species Von Lendenfeld, R. 1885a. The metamorphosis of of platyctenean ctenophore, Lyrocteis flavopallidus Bolina chuni nov. spec. Proceedings of the Linnean sp. nov., from McMurdo Sound, Antarctica. Society of New South Wales 9: 929–931, pls 44, 45. Canadian Journal of Zoology 50: 47–52. 1885b. Notes on a beroid of Port Jackson. Proceedings Sars, M. 1835. Beskrivelser og iagttagelser over nogle of the Linnean Society of New South Wales 9: 968–976. mærkelige eller nye i havet ved den bergenske kyst 1885c. Uber coelenteraten der Sudsee, VI: Neis cor - levende dyr af polypernes, acalephernes, radiaternes, digera Lesson, eine australische Beroide. Zeitschrift annelidernes, og molluskernes classer, med en kort für Wissenschaftliche Zoologie 41: 673–682, pl. 33. oversigt over de hidtil af forfatteren sammesteds fundne Whitelegge, T. 1889. List of the marine and fresh- arter og deres forekommen. (T. Hallager: Bergen). water invertebrate fauna of Port Jackson and the Schulze-Röbbecke, A.C. 1984. Functional morphology neighborhood. Journal and Proceedings of the Royal of Bolinopsis infundibulum (Ctenophora). Helgo- Society of New South Wales 23: 163–324 [coelenter - länder Meeresuntersuchungen 38(1): 47–64. ates pp. 187–197, 306]. Shiganova, T.A. 1998. Invasion of the Black Sea by Willey, A. 1896. On Ctenoplana. Quarterly Journal of the ctenophore Mnemiopsis leidyi and recent Microscopical Science 39(155): 323–343, pl. 21. changes in pelagic community structure. Fisheries Wrobel, D. & Mills, C. 1998. Pacific Coast Pelagic Oceanography 7(3–4): 305–310. Invertebrates: A Guide to the Common Gelatinous Smith, B.J. & Plant, R.J. 1976. A creeping ctenophoran Animals (Sea Challengers and Monterey Bay (Platyctenea: Ctenophora) from Victoria, Australia. Aquarium: Monterey, CA). Memoirs of the National Museum of Victoria 37: 43–46. Zeidler, W. & Gowlett-Holmes, K.L. 1998. Confirm- Stechow, E. 1921. Neue genera und species von ation of the association of the hyperiidean amphipod hydrozen und anderen evertebraten. Archiv fur genus Hyperia (Crustacea: Amphipoda: Hyperiidea: Naturgeschichte Jahrg. 87, Sect. A (Part 3): 248–265. Hyperiidae) with ctenophores. Records of the South Steene, R. 1990. Coral Reefs: Nature’s richest realm Australian Museum 31(1): 117–118. (Crawford House Press: Bathurst, Australia).

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