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The lithistid Demospongiae in New Zealand: species composition and distribution

Kelly, Michelle; Ellwood, Michael; Lincoln, Tubbs; Buckeridge, John https://researchrepository.rmit.edu.au/discovery/delivery/61RMIT_INST:ResearchRepository/12247292150001341?l#13248428070001341

Kelly, M., Ellwood, M., Lincoln, T., & Buckeridge, J. (2006). The lithistid Demospongiae in New Zealand: species composition and distribution. Serie Livros 28, Porifera Research: Biodiversity, Innovation and Sustainability, 393–404. https://researchrepository.rmit.edu.au/discovery/fulldisplay/alma9921860555001341/61RMIT_INST:Resea rchRepository Document Version: Published Version

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Kelly, M, Ellwood, M, Lincoln, T and Buckeridge, J 2006, 'The lithistid Demospongiae in New Zealand: species composition and distribution', in Serie Livros 28, Porifera Research: Biodiversity, Innovation and Sustainability, Rio di Janeiro, Brasil, 7-13 May 2006, pp. 393-404.

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PLEASE DO NOT REMOVE THIS PAGE Porifera Research: Biodiversity, Innovation and Sustainability - 2007 393

The lithistid Demospongiae in New Zealand waters: species composition and distribution Michelle Kelly(1*), Michael Ellwood(2), Lincoln Tubbs(3), John Buckeridge(4)

(1) National Centre for Aquatic Biodiversity and Biosecurity, National Institute of Water and Atmospheric Research (NIWA) Ltd, Newmarket, Auckland, New Zealand. [email protected] (2) Department of Earth and Marine Sciences and Research School of Earth Sciences, The Australia National University, Canberra, ACT 0200, Australia. [email protected] (3) National Institute of Water and Atmospheric Research (NIWA) Ltd, Newmarket, Auckland, New Zealand. [email protected] (4) School of Civil, Environmental and Chemical Engineering, RMIT University, Melbourne, Victoria, Australia. [email protected]

Abstract: The lithistid Demospongiae fauna of New Zealand is reviewed here following the extensive inventory, documentation and revision of the fauna over the period 1991 to 2006. Examination of almost 300 specimens led to the discovery of 29 species (13 of which were new to science) in 9 families. New species of the poorly known phymatellid genera Neoaulaxinia Pisera and Lévi, 2002 and Neosiphonia Sollas, 1888 were described, and a new corallistid genus, Awhiowhio Kelly, 2007, was recognised. All specimens were dredged from between 80 and 1700 m, but were most common between c. 300-1100 m. With the exception of several specimens dredged from the Challenger Plateau to the west of New Zealand, the majority of specimens were found north of the Chatham Rise to the east, and in northern waters, contrasting markedly with what we know of the distribution of these in the past. The availability of hard substrate, ocean circulation patterns, silica availability, and past and present water temperature, are key factors constraining the present day distribution of lithistid sponges in New Zealand.

Keywords: Porifera, lithistid Demospongiae, sponges, taxonomy, New Zealand

Introduction southern locations, in the late Paleocene to early Oligocene (Kelly et al. 2003, Kelly and Buckeridge 2005). Lithistid are a polyphyletic group (de Two major regional faunas are known worldwide: the Laubenfels 1936, Reid 1963, 1970, Kelly-Borges and continental shelf and slope fauna of the tropical western Pomponi 1994), comprising 13 extant families; 36 genera Atlantic region (Schmidt 1870, 1880, van Soest and Stentoft are included in the most recent classification (Pisera and Lévi 1988, Kelly-Borges et al. 1994, Kelly-Borges and Pomponi 2002) but 5 genera remain of uncertain status. They differ 1994, Lehnert and van Soest 1996, Pisera 1999, Pomponi from other demosponges in that the dominant structural et al. 2001), and the seamount fauna of the southwest spicules (desmas) are articulated, forming in most species Pacific including the seamounts of the New Caledonian a solid, rigid, heavily siliceous skeleton. These desmas are Norfolk Ridge (Lévi and Lévi 1983, 1988, Lévi 1991, 1993, highly diverse morphologically; the overall architecture of Schlacher-Hoenlinger et al. 2005). In both locations lithistid the desma, the ornamentation of the desma surface, and the sponges dominate the fauna (Lévi 1991, Reed and Pomponi pattern of articulation with adjacent spicules, is diagnostically 1997, Richer de Forges et al. 2000), with Astrophorida, Halichondrida and Hexactinellida between 150-1800 m, but important (Schrammen 1910; see Kelly 2000a, Pisera and Lévi the structure and taxonomic composition of the communities 2002). An indication of the polyphyletic nature of ‘lithistid’ differ. Lithistid sponges are also common throughout the sponges is revealed in the wide range of microscleres and tropics in relatively shallow waters, and a significant number ectosomal megascleres, and desma axial geometries, which of species are known from South African waters (Kirkpatrick include tetraxial, monaxial, polyaxial and anaxial forms. 1902). Recently, Kelly (2007) recorded the first polar lithistid, A study of lithistid demosponges is of particular relevance Neoschrammeniella antarctica Kelly, 2007. in New Zealand because of their abundance in seamount and deep-sea faunas, and their high level of endemism at the Historical records regional and local scale (Kelly 2007). The past and present distributions of lithistid sponges in New Zealand waters is The earliest record of lithistid sponges in New Zealand disjunct, with lithistid species reported from what are now is to be found in the study of a species-rich assemblage 394 identified from siliceous microfossil spicules embedded in papers document biologically active compounds and their marine diatomaceous sediments from near Oamaru in North synthesis from southwest Pacific genera Aciculites Schmidt, Otago (Hinde and Holmes 1892). This formation is from the 1879, Pleroma, Callipelta Sollas, 1888, Microscleroderma early Runangan (late Eocene) horizon within the Oamaru Kirkpatrick, 1903, Reidispongia Lévi and Lévi, 1988, Diatomite Member of the basaltic Waireka Volcanics (c. 35 Discodermia, Neosiphonia, and Scleritoderma Sollas, 1888, million years ago) (Suggate et al. 1978, Edwards 1991). Hinde present in New Zealand, representing considerable potential and Holmes (1892) illustrated what appeared to be lithistid for biotechnological discovery and subsequent development. desmas amongst the numerous megascleres that were clearly of astrophorid, haplosclerid, halichondrid, poecilosclerid, Records post 1990 and hexactinellid species origin. These included illustrations of choanosomal desmas and ectosomal triaenes of what was For many years Aciculites pulchra and Lepidothenea thought to represent species of ‘Lyidium Schmidt, 1870’, incrustans (Dendy, 1924), were the only lithistid sponges ‘Corallistes Schmidt, 1870’, ‘Discodermia du Bocage, 1869’ known from New Zealand, both recorded during the non- and ‘Theonella Gray, 1868’. While Lyidium (= Pleroma Antarctic phase of the British Antarctic (Terra Nova) Sollas, 1888) and Discodermia are known from New Zealand Expedition of 1910 (Dendy 1924). While Bergquist (1968) waters today (Kelly 2007), Corallistes and Theonella are expanded locality records for A. pulchra and confirmed de restricted predominantly to tropical Pacific and southeast Laubenfels (1936) correction of the original occupied name Asian waters. of Lepiospongia incrustans to Lepidothenea incrustans, no A fuller understanding of the present day New Zealand further species were added. lithistid fauna enabled Kelly (2003), Kelly et al. (2003), and By the late 1990s many of the New Caledonian lithistid Kelly (2007) to increase the diversity of taxa illustrated as species first discovered by Lévi and Lévi (1983, 1988) and spicules in Hinde and Holmes (1892). Spicules identified Lévi (1993), and several new species, were progressively as being from species of Lyidium, were compared with discovered in New Zealand waters (Kelly et al. 1999, Pleroma turbinatum Sollas, 1888 and P. menoui Lévi and Kelly 2000b, Kelly 2001a, 2001b, Pomponi et al. 2001, Lévi, 1983, and the illustration of a discotriaene attributed to Kelly 2003, Kelly et al. 2003, 2004, Kelly and Buckeridge ‘Discodermia sinuosa Carter, 1881’ from the Indian Ocean 2005). In 2000, Homophymia stipitata Kelly, 2000 was by Hinde and Holmes (1892) was considered to be more like described from carbonate banks on the western continental Macandrewia spinifoliata Lévi and Lévi, 1983 present today shelf off Northland (Kelly 2000a). This species was the in the Bay of Plenty. second described within the genus Homophymia Vacelet and Illustrations of short-shafted dichotriaenes attributed to Vasseur, 1971, known previously only from Madagascar and species of Corallistes by Hinde and Holmes (1892) were La Réunion in the Western Indian Ocean. In 2003, a further compared by Kelly et al. (2003) to an undescribed species lithistid endemic to New Zealand was described from of the phymatellid genus Neosiphonia Sollas, 1888. Kelly the same location; Pleroma aotea Kelly, 2003 was only the (2007) expanded this possibility to include closely related third species in this genus to be described (Kelly 2003). genus Neoaulaxinia Pisera and Lévi, 2002 and corallistid A comprehensive inventory, redescription, and revision Awhiowhio Kelly, 2007, which also occur on the Chatham of the New Zealand lithistid Demospongiae, commenced in Rise today. Hinde and Holmes (1892) also described a 2004. Kelly (2007) recognised 9 families, 18 genera and 29 new species of Vetulina Schmidt, 1879, V. oamaruensis species (Table 1). Lithistid sponge specimens were recovered Hinde and Holmes, 1892 based upon what they identified from collections resulting from several hundred New Zealand as sphaerocladine desmas. However, these desmas strongly Oceanographic Institute (NZOI) and National Institute of resemble those found in a species of Crambe Vosmaer, 1880 Water and Atmospheric Research (NIWA) voyages around the from Spirits Bay, Northland; thus their conspecificity with New Zealand EEZ (Gordon 2000, Kelly 2007) (Fig. 1). The this latter species cannot be discounted (Kelly et al. 2003). area covered extends from 24° to 57°S and 155°E to 170°W, Although lithistid sponges have been of considerable covering parts of Lord Howe Seamount Chain, Norfolk Ridge interest to pharmacology over the last 50 years or so, due to to the north, Challenger Plateau to the southwest, Kermadec the broad range of biological activities that their chemical and Colville Ridges to the north east, the Bay of Plenty and extracts possess (see Bewley et al. 1998, Munro et al. 1999, East Cape region to the east, and the Chatham Rise and Pomponi 2001, Mayer and Hamann 2004, Piel et al. 2004), Subantarctic slope in the southeast. only a single paper refers to a New Zealand species. Crist et The systematics scheme used in Kelly (2007) followed al. (1983) reported the first example of double bioalkylation the recent revisions of Andrzej Pisera and Claude Lévi of the of the sterol side chain at position 26 in New Zealand endemic polyphyletic group ‘Lithistid’ Demospongiae (Pisera and Lévi Aciculites pulchra Dendy, 1924. 2002, and subsequent references on this group). The reader is New Caledonian specimens of Pleroma menoui, referred to this major publication for full family and genus- Reidispongia coerulea Lévi and Lévi, 1988 and Neosiphonia level diagnoses and histories, as none will be given here. superstes Sollas, 1888 from New Caledonia, have been the subject of considerable attention for their production of bromindoles (Guella et al. 1989), and antiviral (Laille The New Zealand lithistid sponge fauna et al. 1998) and antifungal (D’Auria et al. 1995) cytotoxic macrolides (D’Auria et al. 1996, Zampella et al. 1997, In the 1980s Professor Claude Lévi described numerous Carbonelli et al. 1999, Bassarello et al. 2000). Over 40 sponges from the seamounts of the Norfolk Ridge south of 395

Fig. 1: New Zealand and the southwest Pacific region. A. Black dots represent NZOI (New Zealand Oceanographic Institute) collection localities from which lithistid sponges were recorded; B. Geographic names of the major marine ridges, seamount chains, basins, and plateaus in the New Zealand region. Depth contours are 500 m, 1000 m, 2000 m, and 4000 m.

New Caledonia, hailed as refuges for invertebrates whose and probably represents the southernmost reaches of the nearest relatives were Mesozoic fossils (c. 260-60 million species range. years ago) (Bourseau et al. 1987, Richer de Forges et al. Neosiphona superstes and Pleroma turbinatum were first 2000). The discovery of almost 20 lithistid species in almost as recorded from the Fiji Islands (Sollas 1888) (Table 2), but are many genera by Lévi and Lévi (1983, 1988) and Lévi (1991), now known to occur as far south as the south New Caledonia coupled with the observation that over 60% of New Zealand slope and New Zealand (Lévi and Lévi 1983, Kelly 2003, lithistid genera are also represented by only a single species 2007, Schlacher-Hoenlinger et al. 2005). (Kelly 2007), confirm the ‘relict’ nature of these southwest The New Zealand region shares 13 lithistid species with Pacific lithistid sponge species. The majority of non-lithistid the New Caledonian southern slope and northern Norfolk genera are represented by numerous species Ridge seamounts (Table 2, 3). The majority of these were (Kelly et al. 2007). found north of East Cape, but 60% were found in the Bay Several southwest Pacific lithistid genera with species in of Plenty, 54% in the Three Kings region, and 40% in the New Caledonia (Anaderma Lévi and Lévi, 1983, Corallistes, Cavalli Seamount region (Fig. 1). Isabella Schlacher-Hoenlinger et al. 2005, Jereicopsis Lévi and Lévi, 1983 and Neophrissospongia Pisera and Lévi, Endemic New Zealand lithistid species 2002) are not found in New Zealand, but are found elsewhere New Zealand waters harbour 15 endemic species of lithistid in southeast Asia and the west central Atlantic. Herengeria sponges and 2 endemic genera (Table 2, 3). Awhiowhio is Lévi and Lévi, 1988, Neoschrammeniella Pisera and Lévi, closely related to the corallistid genus Herengeria, with 2002 (Family Corallistidae), and Neoaulaxinia, Neosiphonia, three new species confirming the integrity of the genus and Reidispongia (Family Phymatellidae) are endemic to the (Kelly 2007). Unfortunately A. sepulchrum Kelly, 2007 was southwest Pacific region, and Lepidothenea de Laubenfels, described from an incomplete and macerated specimen, and 1936 and Awhiowhio are endemic to New Zealand (Table 2). requires further study when fresh material is obtained. The Scleritoderma flabelliformis Sollas, 1888 was found to be most common of the three species, A. osheai Kelly, 2007, has common to New Zealand, southeast Asia and the west central not been recorded outside the Bay of Plenty. Lepidothenea Pacific (Table 2). Despite this being the only species with incrustans was only recorded once from the North Cape and such a distribution, the species and megasclere morphology has not been seen since. (and ornamentation) were considered to be so similar that The description of two new species of the genus a new species could not be justified under those characters. Leiodermatium Schmidt, 1870 from the southwest Pacific There is some doubt, however, that these specimens comprise region in Kelly (2007) provided the first record of this genus the same species, due to the lack of overall characters for further south than the Philippines; Leiodermatium is typically differentiating species within Scleritoderma (Kelly 2007). present in the west central Pacific, extending into southeast Only a single specimen was found on the Three Kings Ridge Asia, but is also present on both sides of the tropical Atlantic 396 Table 1: Classification and checklist of lithistid Demospongiae Ocean. Two additional tropical west central Pacific and species from the New Zealand region (after Kelly 2007).* southeast Asian species, L. intermedia Sollas, 1888, and L. colini Kelly, 2007, were recognised as part of the revision of Class Demospongiae Sollas, 1885 this group (Kelly 2007) (Table 3). Kelly (2007) also recognised and described several new Lithistid Demospongiae (formerly Order Lithistida Schmidt, 1870) species of the poorly known phymatellid genus Neoaulaxinia Family von Lendenfeld, 1903 and Neosiphonia (Table 1) previously only known from the Discodermia proliferans Lévi and Lévi, 1983 type species Neoaulaxinia clavata (Lévi and Lévi, 1988) and Neosiphonia superstes, respectively. In the genus Family Phymatellidae Schrammen, 1910 Neoaulaxinia, the species zingiberadix Kelly, 2007 was Neoaulaxinia clavata (Lévi and Lévi, 1988) described from a single specimen dredged off the western Neoaulaxinia zingiberadix Kelly, 2007 continental slope of Northland, while N. persicum Kelly, 2007 Neoaulaxinia persicum Kelly, 2007 is very common and widely spread from the Three Kings Neosiphonia superstes Sollas, 1888 region in the north to the northern edge of the Chatham Rise Neosiphonia motukawanui Kelly, 2007 in the southeast. The second known species of Neosiphonia, Reidispongia coerulea Lévi and Lévi, 1988 N. motukawanui Kelly, 2007, is only known from the West Cavalli Seamount (Table 3). Family Corallistidae Sollas, 1888 Significant levels of endemism in lithistid sponges occur on Neoschrammeniella fulvodesmus (Lévi and Lévi, 1983) the continental slope and carbonate banks west of North Cape Herengeria auriculata Lévi and Lévi, 1988 (80%), the Bay of Plenty region (43%), the Cavalli Seamounts Herengeria vasiformis Schlacher-Hoenlinger et al. 2005 region (44%), West Norfolk Ridge (40%), Kermadec Ridge Awhiowhio osheai Kelly, 2007 (40%), and the Three Kings Ridge (36%). Awhiowhio unda Kelly, 2007 Awhiowhio sepulchrum Kelly, 2007 Geographic distribution Family Neopeltidae Sollas, 1888 Homophymia stipitata Kelly, 2000 The majority of New Zealand lithistid demosponge species are found in two relatively distinct regions off northern and Callipelta punctata Lévi and Lévi, 1983 northeastern New Zealand (Table 3). The first region spans Neopelta pulvinus Kelly, 2007 northern New Zealand from the Lord Howe Seamount Chain Family Macandrewiidae Schrammen, 1924 and the Challenger Plateau in the north and west, across the Macandrewia spinifoliata Lévi and Lévi, 1983 West Norfolk Ridge east to the North Cape of New Zealand Family Pleromidae Sollas, 1888 (Fig. 1). This region includes the carbonate banks and Pleroma turbinatum Sollas, 1888 seamounts to the north and west of the Three Kings Islands and contains 61% of New Zealand’s lithistid species. Although Pleroma menoui Lévi and Lévi, 1983 the area of greatest diversity in this region is the Three Kings Pleroma aotea Kelly, 2003 Ridge and seamounts region, with 40% of New Zealand Family Isoraphiniidae Schrammen, 1910 lithistid species, only 1 species is endemic (Lepidothenea Costifer wilsoni Lévi, 1993 incrustans). Three species are endemic to the West Norfolk Family Scleritodermiidae Sollas, 1888 Ridge and slope region (Homophymia stipitata, Neoaulaxinia Microscleroderma novaezelandiae Kelly, 2007 zingiberadix, Aciculites manawatawhi Kelly, 2007) and one to the Lord Howe Seamount Chain (Leidermatium dampieri Scleritoderma flabelliformis Sollas, 1888 Kelly, 2007). Aciculites pulchra Dendy, 1924 The second region that harbours a majority of New Zealand Aciculites manawatawhi Kelly, 2007 lithistid sponges is to the northeast of New Zealand, from the Aciculites sulcus Kelly, 2007 Kermadec Ridge, south to the Cavalli Seamount region off Family Azoriciidae the northeast coast of Northland, and east to the volcanically Leiodermatium dampieri Kelly, 2007 active seamounts of the Bay of Plenty (Table 3, Fig. 1). This Leiodermatium linea Kelly, 2007 region contains 64% of New Zealand’s lithistid species. It is noteworthy that five species are endemic to this general Lithistid Demospongiae incertae sedis region and three are unique to the Bay of Plenty (Aciculites Lepidothenea incrustans (Dendy, 1924) sulcus Kelly, 2007, Awhiowhio osheai, and Leidermatium linea Kelly, 2007). *The systematics scheme used in Kelly (2007) followed the recent revisions The southeast New Zealand region (Raukumara Plain, of Andrzej Pisera and Claude Lévi of the polyphyletic group ‘Lithistid’ Hikurangi Plateau, south to the Chatham Rise) is depauperate Demospongiae in the Systema Porifera: a guide to the classification of lithistid demosponges compared to the northern regions; of sponges (Pisera 2002 and subsequent references on this group). The only 21% of New Zealand total lithistid fauna (6 species) reader is referred to this major publication for full family and genus-level diagnoses and histories, as none will be given here. were recorded from this region (Table 3, Fig. 1). Lithistid sponges were not recorded south of the Graveyard Seamount complex on the northern edge of the Chatham Rise. Only two species were recorded from the Graveyard seamount 397 Table 2: Checklist and geographic distribution of southwest Pacific Lithistid Demospongiae species.

Indian Southeast Fiji New New Ocean Asia Islands Caledonia Zealand

Scleritoderma flabelliformis Sollas, 1888 X X X Aciculites orientalis Dendy, 1905 X X Microscleroderma herdmani (Dendy, 1905) X X Neosiphonia superstes Sollas, 1888 X X X Pleroma turbinatum Sollas, 1888 X X X Aciculites oxytylota Lévi and Lévi, 1983 X Aciculites papillata Lévi and Lévi, 1983 X Anaderma rancureli Lévi and Lévi, 1983 X Corallistes australis Schlacher-Hoenlinger et al. 2005 X Corallistes multituberculatus Lévi and Lévi, 1983 X Corallistes undulatus Lévi and Lévi, 1983 X Homophymia pollubrum Schlacher-Hoenlinger et al. 2005 X Isabella mirabilis Schlacher-Hoenlinger et al. 2005 X Jereicopsis graphidophora Lévi and Lévi, 1983 X Microscleroderma stonae Lévi and Lévi, 1983 X Neopelta plinthosellina Lévi and Lévi, 1988 X Neophrissospongia microstylifer Lévi and Lévi, 1983 X Neoschrammeniella castrum Schlacher-Hoenlinger et al. 2005 X Neoschrammeniella moreti (Lévi and Lévi, 1988) X Neoschrammeniella norfolkii Schlacher-Hoenlinger et al. 2005 X Scleritoderma camusi Lévi and Lévi, 1983 X Callipelta punctata Lévi and Lévi, 1983 X X Costifer wilsoni Lévi, 1993 X X Discodermia proliferans Lévi and Lévi, 1983 X X Herengeria auriculata Lévi and Lévi, 1988 X X Herengeria vasiformis Schlacher-Hoenlinger et al. 2005 X X Macandrewia spinifoliata Lévi and Lévi, 1983 X X Neoaulaxinia clavata Lévi and Lévi, 1988 X X Neoschrammeniella fulvodesmus (Lévi and Lévi, 1983) X X Neosiphonia motukawanui Kelly, 2007 X X Pleroma menoui Lévi and Lévi, 1983 X X Reidispongia coerulea Lévi and Lévi, 1988 X X Aciculites manawatawhi Kelly, 2007 X Aciculites pulchra Dendy, 1924 X Aciculites sulcus Kelly, 2007 X Awhiowhio osheai Kelly, 2007 X Awhiowhio sepulchrum Kelly, 2007 X Awhiowhio unda Kelly, 2007 X Homophymia stipitata Kelly, 2000 X Leiodermatium dampieri Kelly, 2007 X Leiodermatium linea Kelly, 2007 X Lepidothenea incrustans (Dendy, 1924) X Microscleroderma novaezealandiae Kelly, 2007 X Neoaulaxinia persicum Kelly, 2007 X Neoaulaxinia zingiberadix Kelly, 2007 X Neopelta pulvinus Kelly, 2007 X Pleroma aotea Kelly, 2003 X

complex, one of which is endemic to the locality (Awhiowhio were collected between 80-1700 m, the water temperatures at sepulchrum), and only known from the type specimen. these sites ranging from less than 4°C to c. 18°C (Fig. 2A). Interestingly, the other species, Neoaulaxinia persicum, was The deepest sites (>1000 m) were on seamounts in the found in considerable numbers and many of the specimens Cavalli and Bay of Plenty regions (1040-1540 m), the North were quite small probably representing a recent settlement Chatham Rise, the South Norfolk Basin (1100-1680 m), from more northern sites. Ngatoro Ridge off East Cape (1050 m) and on the Louisville Seamount Chain (1050 m). Species found at the greatest Depth and temperature range depths and thus experienced the lowest temperatures of below 6°C, include Pleroma aotea in the Family Pleromidae, Lithistid sponges are found in most temperate and tropical Neoaulaxinia persicum, N. clavata and Reidispongia coerulea oceans in the world but are generally restricted to depths in the Family Phymatellidae, and Neoschrammeniella greater than 150 m and are rarely collected below 1700 m fulvodesmus, and Herengeria vasiformis in the Family (Pomponi et al. 2001). The New Zealand lithistid sponges Corallistidae. 398 Table 3: Geographic distribution of endemic lithistid Demospongiae species and those common to the New Zealand and New Caledonian Norfolk Ridge region. X = New Zealand endemic, ▲ = common species. Shaded columns link closely adjacent biodiverse regions. Challenger Plateau Lord Howe Seamount Chain Norfolk Ridge West South Norfolk Basin Continental Slope Western Three Kings Ridge South Fiji Basin Colville Ridge Kermadec Ridge Cavalli Seamount region Bay of Plenty Raukumara Plain East Cape Louisville Seamount Chain Hikurangi Plateau Chatham Rise of species number Total

Leiodermatium dampieri Kelly, 2007 X 1 Pleroma turbinatum Sollas, 1888 ▲ ▲ ▲ ▲ 3 Reidispongia coerulea Lévi and Lévi, 1988 ▲ ▲ 2 Neoschrammeniella fulvodesmus (Lévi and Lévi, 1983) ▲ ▲ ▲ ▲ ▲ 5 Homophymia stipitata Kelly, 2000 X X 2 Costifer wilsoni Lévi, 1993 ▲ ▲ ▲ ▲ 3 Pleroma aotea Kelly, 2003 X X X X X 4 Herengeria vasiformis Schlacher-Hoenlinger et al. 2005 ▲ ▲ ▲ ▲ ▲ 5 Pleroma menoui Lévi and Lévi, 1983 ▲ ▲ ▲ ▲ ▲ ▲ ▲ ▲ 8 Aciculites pulchra Dendy, 1924 X X X X X X X 8 Neoaulaxinia zingiberadix Kelly, 2007 X 1 Aciculites manawatawhi Kelly, 2007 X 1 Neoaulaxinia persicum Kelly, 2007 X X X X 4 Discodermia proliferans Lévi and Lévi, 1983 ▲ ▲ 2 Herengeria auriculata Lévi and Lévi, 1988 ▲ 1 Lepidothenea incrustans (Dendy, 1924) X 1 Scleritoderma flabelliformis Sollas, 1888 ▲ 1 Awhiowhio unda Kelly, 2007 X 1 Neopelta pulvinus Kelly, 2007 X 1 Neosiphonia superstes Sollas, 1888 ▲ ▲ ▲ 3 Neoaulaxinia clavata Lévi and Lévi, 1988 ▲ 1 Neosiphonia motukawanui Kelly, 2007 X 1 Aciculites sulcus Kelly, 2007 X 1 Awhiowhio osheai Kelly, 2007 X 1 Callipelta punctata Lévi and Lévi, 1983 ▲ 1 Leiodermatium linea Kelly, 2007 X 1 Macandrewia spinifoliata Lévi and Lévi, 1983 ▲ 1 Awhiowhio sepulchrum Kelly, 2007 X 1

Total number of species 1 2 5 2 5 11 2 1 5 9 14 1 1 1 2 2

Some of the shallower sites were volcanically active considerable depth and thus temperature ranges. Aciculites seamounts and knolls in the Bay of Plenty (Rungapapa and pulchra was found between 80 and 1100 m (5-17°C), and Tuatoru Knolls), and on the Cavalli Seamount Ridge (140- Neoschrammeniella fulvodesmus (Lévi and Lévi, 1983) had 200 m). The shallowest sites were just off Ngunguru Bay the greatest depth range of 100-1680 m (5-15°C) (Fig. 2). (80 m) and on Cavalli Seamount (103 m), Northland. Over 60% of species were found in waters shallower than c. 300 m Species displaying very narrow depth and temperature ranges, (experiencing water around c. 14-18°C). such as Neoaulaxinia zingiberadix, A. sulcus, Leiodermatium While the majority of specimens were collected between dampieri, L. linea, and Lepidothenea incrustans, were only 200 and 1100 m (Fig. 2A), many of these species had recorded from a single location. 399 Discussion Ocean circulation – While the distribution of lithistid sponges coincides with the distribution of major hard grounds such Lithistid demosponges are absent south of the Subtropical as seamount chains and banks across northern New Zealand, Convergence just below the Chatham Rise (Shackelton and they are absent from the hard grounds on seamounts and Kennett 1975, Nelson and Cooke 2001), and restricted to continental margins south of the Subtropical Convergence ‘warmer’ waters in northern New Zealand, extending from below the Chatham Rise. What factors contribute to this the Lord Howe Seamount Chain in the west through the West disjunct distribution? Norfolk Ridge, western continental margin off North Cape, The major current systems around New Zealand (Fig. 3) Three Kings Ridge, east to the Cavalli Seamounts, Kermadec coincide with areas of high diversity of lithistid sponges, and volcanic arc, and Bay of Plenty. Several sites within the latter other sponges and invertebrates; the East Auckland Current, three regions have relatively high diversity and abundance of fed by the Tasman Front (Stanton 1969, Carter et al. 1998), is lithistid sponges; these sites are relatively shallow (100-200 reknowned for its contribution of subtropical taxa to eastern m) compared to the majority of sites, where lithistid sponges New Zealand locations (Squires 1964, Yaldwyn 1968, typically occur much deeper. What are some of the factors Russell and Ayling 1976, Powell 1976, Ayling 1982, see also that effect the distribution of lithistid sponges today? Kelly 1983) and may explain the rather high incidence of Availability of hard substrate – Lithistid sponges require New Caledonian lithistid species on the West Norfolk Ridge, hard rocky substrate for settlement (Pomponi et al. 2001). Three Kings Ridge and Kermadec Ridge. Major eddies and With the exceptions of free-living Herengeria auriculata gyres from the East Auckland current dominate the Kermadec Lévi and Lévi, 1988, and Discodermia proliferans Lévi and and Colville Ridges, sustaining the high population levels in Lévi, 1983, the vast majority of lithistids are found attached these regions. The major current systems on the west coast to rock, e.g., carbonate reef rubble, mudstone, or basalt, in of New Zealand are the West Auckland Current in the north temperate and tropical regions. As for most sponges and and the Westland Current to the south. The West Auckland many other invertebrates, larval settlement is facilitated by Current, which diverges from the East Auckland Current to hard surfaces devoid of macroinvertebrate competitors and move south along the west coast of the North Island, may heavy sediment loads. Lithistid sponges are thus restricted however explain the occurrence of a few rare specimens on to deep and shallow sites that have an abundance of hard the northern edge of the Challenger Plateau (Fig. 1). sediment-free substrate such as on steep-sided seamounts, The East Cape Current and Wairarapa Eddy, which touch and ledges on continental margins. This is clearly illustrated the northern boundary of Chatham Rise, may be a source of in the restriction of lithistid sponges to the Graveyard larvae of the single dominant lithistid species, Neoaulaxinia Seamount complex on the North Chatham Rise; no lithistids persicum, on the isolated Graveyard Seamount complex have been recorded at other sites due, we believe, to their situated on the northern edge of the Chatham Rise, In this sediment loading. The sediments on the Chatham Rise are a location specimens are as numerous as in the Bay of Plenty and combination of relic authigenic sediment, whereas towards the Cavalli Seamount region further north and east. The second south the seafloor is draped with a fine-grained combination species known from the Graveyard Seamounts is only known of pelagic, hemipelagic sediment (Carter et al. 2000). from a single specimen, which was dead on collection. Carbonate banks such as Pandora and Wanganella to the west Finally, the Chatham Rise constrains the Subtropical of North Cape also provide an abundance of clean carbonate Convergence thereby forming a strong barrier to the transport rubble for settlement of lithistid sponges and numerous other of larvae southwards over the Chatham Rise, perhaps invertebrates (Kelly 2000a). contributing to the absence of sponges below the Subtropical The extent of settlement success of lithistid sponges in some Convergence. locations where hard substrate is available was illustrated by Kelly et al. (2003, Fig. 10), in which the basalt substrate, Historical patterns of distribution – As noted previously, viewed using an epibenthic sled, was closely covered with microfossil spicules very similar to those of living lithistid immature sponges. The site was relatively shallow at 160 m sponges were recorded further south than the present day and was situated on Rungapapa Knoll in the Bay of Plenty. This Subtropical Convergence. Hinde and Holmes (1892) recorded settlement success also appears in at least one population of diverse spicules in the Oamaru Diatomite (c. 35 million years lithistid sponges living on a shallow (50-150 m) volcanic tuff ago) and Kelly and Buckeridge (2005) recorded microfossil cone (Lee et al. 1997) off the South Island in the late Eocene lithistid desmas in the Tutuiri Greensand of Chatham Island (35-33 million years ago) (Kelly et al. 2003). The sponge- (c. 56-53 million years ago). Sponge body fossils attributed bearing horizon contained extremely numerous sponge body to the extant lithistid Pleroma aotea were recorded from the fossils with a ‘normal’ bell-shaped size frequency distribution Ototara Limestone of Kakanui (c. 33.7 million years ago). population curve (Kelly et al. 2003: Fig. 8). The stratigraphy Campbell et al. (1993) contend that water temperatures in the and palaeoecology of the horizon indicated that the sponges southern parts of the New Zealand region were sufficiently were attached to volcanically derived Mineral Breccia during warm during periods of the Cainozoic to permit the establish- their life, and the uniform distribution of the fossils throughout ment of sponges, cirripedes (Buckeridge 1993, 1996, 1999), the sponge-bearing horizon support the hypothesis that the deep-sea bryozoans (Gordon 1984), brachiopods (Lee et al. live sponges were simultaneously detached by a submarine 1997), crinoids (Lee 1987), and molluscs (Beu and Maxwell flow down the flanks of the tuff cone, transported down slope 1990), that today are generally found in waters north of the and rapidly buried. Chatham Rise. 400

Fig. 2: Box and Whisker plots of depth distribution and bottom-water temperatures for New Zealand lithistid species. A. Depth range for combined specimens collected within each lithistid species from stations listed by Kelly et al. (2007); B. Estimated bottom-water temperature at sponge collection station. Bottom water temperatures were estimated from nearby WOCE stations where standard hydrographic data have been collected (www.clivar.org). Upper and lower bound of box represents 75% and 25% of range, mid line in box is the median, upper and lower bounds of whiskers are maxima and minima.

It is thought that around the end of the Eocene major development of cold bottom waters (Shackleton and Kennett changes in Southern Ocean circulation occurred as the Drake 1975). The amount of biologically-available silica in shallow Passage between South America and Antarctica opened environments also decreased markedly after the Eocene, (Flower et al. 1997). The formation of the Circum Antarctic because of the increase in diatom diversity and abundance Current lead to cooling of the Southern Ocean and the (Maliva et al. 1989); only the deepest waters around New 401

Fig. 3: Modern-day circulation patterns within the New Zealand region. Abbreviations for fronts features: TF = Tasman, STF = Subtropical, SAF = Subantarctic; Abbreviations for surface currents: WAUC = West Auckland, EAUC = East Auckland, ECC = East Cape, DC = D’Urville, WC = Westland, SC = Southland, ACC = Antarctic Circumpolar Current. This figure taken from Carter (2001), but based on Carter et al. (1998).

Zealand are silica-rich today (Vincent et al. 1991, Frew and deeper (Kelly 2007) (Fig. 1). The broad Bay of Plenty region Hunter 1995). These events contribute to our understanding of covers less than 15% of the marine region studied yet it har- the extinction of lithistid sponges and some other invertebrate bours over half of New Zealand lithistid species recorded and species from southern New Zealand waters after the Eocene, has several endemic species. What factors facilitate higher as evidenced by Gordon (1984), Hinde and Holmes (1892), settlement rates and greater diversity in these relatively shal- Lee et al. (1997), Buckeridge (1996), Kelly et al. (2003), and low sites? Kelly and Buckeridge (2005), and the present day restriction The Bay of Plenty and southern Kermadec Volcanic Arc of lithistids to northern ‘warmer’, deeper waters. regions are hydrothermally active with a number of submarine Silica availability – Several sites within the Cavalli Seamounts hydrothermal plumes releasing hydrothermal fluids into the region (NZOI I014: 103 m), the Bay of Plenty (Rungapapa water column. Basic chemistry measurements of these fluids and Tuatoru Knolls: 136+ m), on Kermadec Ridge Volcano indicate that they are rich in silica and heavy metals (Pantin L: 154+ m), and on carbonate banks off North Cape (192+ and Wright 1994, Tarasov 2006). Primary phytoplankton m) have relatively high diversity and abundance of lithistid production in the waters near fluid discharges was found to sponges; these sites are relatively shallow compared to the be significantly greater than in surrounding oceanic waters, majority of sites where lithistid sponges typically occur much thereby indicating release of increased nutrients to fuel 402 primary and secondary production (Sorokin et al. 1998). Bergquist PR (1968) The marine fauna of New Zealand: Porifera, Lithistid sponges living in the waters near these vents would Demospongiae, Part 1 (Tetractinomorpha and Lithistida). Mem benefit greatly from the increased silicon concentration within New Zeal Oceanogr Inst 37: 1-106 the water column and the increases in food supply linked to Beu A, Maxwell PA (1990) Cenozoic Mollusca of New Zealand. increased primary production. New Zeal Geol Surv Paleontol Bull 58: 1-518 A recent geochemical study has shown considerable Bewley CA, Faulkner D, Angew J (1998) Lithistid sponges - star hydrothermal fluid release from a number of submarine performers or hosts to the stars [review]. Angewanta Chemi volcanoes along the southern Kermadec Ridge system (de International Edition English 37: 2163-2178 Ronde et al. 2001). Such fluid releases are rich in heavy Bourseau JP, Ameziane-Cominardi N, Roux M (1987) Un Crinoïde metals and are likely, although no silicon measurements were pédonculé nouveau (Echinodermes) représentant actuel de la made, to enrich in silica which would explain shallow depth famille jurassique de Hemicrinidae: Gymnocrinus richeri nov. distribution of lithistid sponges along the Kermadec volcano sp. des fonds bathyaux de Nouvelle-Calédonie (SW Pacific). C R system. Hebd Acad Sci Paris 305(3): 595-599 The carbonate banks west of North Cape are an interesting Bradford JM, Roberts P (1978) Distribution of reactive phosphorus exception as they are not presently hydrothermally active. and plankton in relation to upwelling and surface circulation Wanganella Bank may have had localised phases of volcanism, around New Zealand. NZ J Mar Fr Res 12(1): 1-15 however, with hydrothermal activity associated with older Buckeridge JS (1983) The fossil barnacles (Cirripedia: Thoracica) plate boundary positions to the north of New Zealand (Herzer of New Zealand and Australia. New Zeal Geol Surv Paleontol Bull and Mascle 1996, Balance 1999). However, these banks are 50: 1-151 in regions of strong upwelling (Stanton 1976) and primary Buckeridge JS (1996) Phylogeny and biogeography of the primitive and secondary productivity are elevated as a result (Bradford Sessilia and a consideration of a Tethyan origin for the group. and Roberts 1978). Crustacean Issues 10: 225-267 We conclude that the disjunct distribution of lithistid Buckeridge JS (1999) Post Cretaceous biotic recovery: a case sponges, both geographically and in depth, is due to multiple study on (Crustacea: Cirripedia) from the Chatham Islands, New factors. The most important of these factors include major Zealand. Rec Cant Mus 13: 43-51 changes in Southern Ocean circulation patterns coupled Campbell HJ, Andrews PB, Beu AG, Maxwell PA, Edwards AR, with global reduction in bioavailable silica in shallow water Laird MG, Hornibrook NdeB, Mildenhall DC, Watters WA, since the Palaeogene, overlayed by changing patterns of Buckeridge JS, Lee DE, Strong CP, Wilson GJ, Hayward BW active submarine volcanism, and maintained in the present (1993) Cretaceous-Cainozoic geology and biostratigraphy of the day by ocean currents, high primary productivity in areas of Chatham Islands. New Zeal Inst Geol Nucl Sci Monogr 2: 1-240 upwelling and hydrothermal activity, and determined overall Carbonelli S, Zampella A, Randazzo A, Debitus C, Gomez-Paloma by the availability of hard substrate. L (1999) Sphinxolides E-G and reidispongiolide C: four new cytotoxic macrolides from the New Caledonian lithistida sponges Acknowledgements N. superstes and R. coerulea. Tetrahedron 55: 14665-14674 Carter L (2001) Currents of change: the ocean flow in a changing Research support and fieldwork for aspects of the research leading world. Water and Atmosphere 9: 15-17 to this review were funded by Biological and Biotechnological Carter L, Garlick RG, Sutton P, Chiswell SM, Oien NA, Stanton Research Council (BBSRC), UK, and British Airways Conservation BR (1998) Ocean circulation New Zealand. NIWA Chart. Travel, while the senior author was employed at the Natural History Miscellaneous Series 76 Museum, London (1992-1996). Support in New Zealand was given Carter L, Neil HL, Mccave IN (2000) Glacial to interglacial changes by a New Zealand Lotteries Grants Board Repatriation Fellowship in non-carbonate and carbonate accumulation in the SW Pacific to MK (1997-1999), a Royal Society of New Zealand Marsden Ocean, New Zealand. Palaeogeography Palaeoclimatology Fund Grant to MK (2000-2002), and a French Foreign Affair Ministry travel grant to work with IRD (Institut de Recherche pour Palaeoecology 162: 333-356 le Developement) in New Caledonia (1999). In later years this work Crist BV, Li X, Bergquist PR, Djerassi C (1983) Sterols of marine was supported by the New Zealand Foundation for Research Science invertebrates. 44. Isolation, structure elucidation, partial synthesis, and Technology (Contracts C01X0028 (Seamounts) and C01X0219 and determination of absolute configuration of pulchrasterol. The (Biodiversity)) to NIWA. first example of double bioalkylation of the sterol side chain at position 26. 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