From Northern Bass Strait, Southern Australia

Home , Carteriospongia, Corticata, Corticium candelabrum, East Island (Tasmania), Echinoclathria, Erith Island

31 August 1989 Memoirs of the Museum of Victoria 50(1): 1-242 (1989) ISSN 0814-1827 https://doi.org/10.24199/j.mmv.1989.50.01

DEMOSPONGIAE (PORIFERA) FROM NORTHERN BASS STRAIT, SOUTHERN AUSTRALIA

By Felix Wiedenmayer

Department of Invertebrate Zoology, Museum of Victoria, Swanston Street, Melbourne, Victoria 3000, Australia Present address: Naturhistorisches Museum Basel, Agustinergasse 2, 4001 Basel, Switzerland

Abstract

Wiedenmayer, F., 1989. Demospongiae from northern Bass Strait, southern Australia. Memoirs of the Museum of Victoria 50(1): 1-242. Eighty-four species (in 47 genera) in the Museum of Victoria, Melbourne, are described and illustrated. Of these, 21 species are described as new: Ancorina repens, A. suina, Stelletta arenitecta, Rhabdastrella cordata, R. intermedia, Tetilla praecipua, Latrunculia hallmanni, Pseudaxinella decipiens, Reniochalina sectilis, Rhaphoxya felina, Clathria wilsoni, Echinoclathria egena, Psammoclema bitextum, P. fissuratum, P. goniodes, P. radiatum, P. stipitatum, P. vansoesti, Callyspongia persculpta, C. toxifera, and Thorecta glomerosus. Eighteen records are new for the Maugean province, and three (Phorbas tenacior, Darwinella gardineri, and Gel- liodes incrustans) are new for the Australian fauna. The following revisions depart from those adopted in Wiedenmayer et al. (in press). The family Desmacididae is divided into Desmacidi- nae and Stylotellinae, and the genera Stylotella ( = Batzella), Phoriospongia ( = Chondropsis),

and Psammoclema ( = Psammopemma, Sarcocornea) are assigned to the latter. Dactylia,

Chalinopsilla and Arenosclera are synonymised with Callyspongia. Thorectandra is synonymised with Thorecta. Dendrilla cactos (Selenka) is a senior synonym of D. rosea Lendenfeld. The composition of this collection is even, with respect to the known demosponge fauna of Victoria and Tasmania. Its zoogeographic affinity is essentially Indo-West Pacific and relictic Tethyan,

its provincial endemism high, and its overlap with the Antarctic/Subantarctic fauna almost nil.

Contents Abstract j Introduction 5 Collections 6 List of collecting stations 6 Taxonomic procedure g Terminology and glossary ] j Size of specimens and its gradation , 12 Abbreviations 12 Class Demospongiae Sollas, 1885 13 Subclass Homoscleromorpha Levi, 1953 13 Order Homosclerophorida Dendy, 1905 13 Plakinidae Schulze, 1880 ...Ill ...... 13 Corticium Schmidt, 1 862 13 Corticium candelabrum Schmidt 13

Subclass Tetractinomorpha Levi, 1953 14 Order Astrophorida Sollas, 1887 14 Ancorinidae Schmidt, 1870 14 Ancorina Schmidt, 1862 14 Ancorina geodides Carter 14 Ancorina repens sp. nov lg Ancorina robusta Carter 17 Ancorina suina sp. nov 1 g Stelletta Schmidt, 1862 , . \9 Stelletta arenitecta sp. nov 20

1 F. WIEDENMAYER

2{ Rhabdastrella Thiele, 1903 2 ' Rhabdastrella cordatu sp. nov Rhabdastrella intermedia sp. nov

Geodiidae Gray, 1 867 ^3 Geodia Lamarck, 1815 Subgenus Geodia 24 Geodia (Geodia) punctata Hentschel 26 Coppatiidae Topsent, 1898 26 Jaspis Gray, 1867 26 Jaspis stellifera (Carter)

Order Spirophorida Brien, 1968 28 28 Tetillidae Sollas, 1886 28 Tetilla Schmidt, 1 868 28 Tetilla praecipua sp. nov

Order Hadromerida Topsent, 1894 29 Tethyidae Gray, 1867 29 Tethya Lamarck, 1814 29 Tethya ingalli Bowerbank 29

Polymastiidae Gray, 1867 32 Polymastia Bowerbank, 1863 32 Polymastia crassa Carter 32

Spirastrellidae Ridley and Dendy, 1886 34 Spirastrella Schmidt, 1868 34 Spirastrella papillosa Ridley and Dendy 35

Chondrillidae Gray, 1872 36

Chondrosia Nardo, 1 847 36 Chondrosia reticulata (Carter) 36

Latrunculiidae Topsent, 1922 37

Latrunculia Bocage, 1 869 40 Lutrunculia conulosa Hallmann 40 Latrunculia hallmanni sp. nov 41

Order Axinellida Levi, 1955 47 Axinellidae Ridley and Dendy, 1887 47 Pseudaxinella Schmidt, 1875 47 Pseudaxinella decipiens sp. nov 47

Reniochalina Lendenfeld, 1 888 48 Reniochalina sectilis sp. nov 49 Rhaphoxya Hallmann, 1917 50 Rhuphoxya cactiformis (Carter) 50 Rhaphoxya felina sp. nov 52

Trachycladidae Hallmann, 1917 53

Trachycladus Carter, 1 879 53 Trachycladus laevispirulifer Carter 53

Raspailiidae Hentschel, 1923 54 Clathriodendron Lendenfeld, 1888 55 Clathriodendren cacticutis (Carter) 55

Subclass Ceractinomorpha Levi, 1953 56 Order Poecilosclerida Topsent, 1928 56 Microcionidae Carter, 1875 .56 SPONGES FROM BASS STRAIT

Clathria Schmidt, 1862 56 Clathria transiens Hallmann 57 Clathria wilsoni sp. nov 57 Echinoclathria Carter, 1885 58 Echinoclathria carteri Ridley and Dendy 59 Echinoclathria favus Carter 59 Echinoclathria laminaefavosa (Carter) 60 Echinoclathria leporina (Lamarck) 61 Echinoclathria globosa (Lendenfeld) 63 Echinoclathria egena sp. nov 64 Echinoclathria tubulosa (Hallmann) 66

Desmacididae Schmidt, 1870 66 Desmacidinae 67 Strongylacidon Lendenfeld, 1897 67 Strongylacidon stelliderma (Carter) 68 Stylotellinae Lendenfeld, 1888 69 Stylotella Lendenfeld, 1888 69 Stylotella inaequalis (Hentschel) 69 71 Phoriospongia Marshall, 1 880 Phoriospongia argentea (Marshall) 72 Phoriospongia carcinophila (Lendenfeld) 72 Phoriospongia kirki (Bowerbank) 73 Psammoclema Marshall, 1880 75 Psammoclema bitextum sp. nov 75 Psammoclema callosum (Marshall) 76 Psammoclema densum (Marshall) 76 Psammoclema fissuratum sp. nov 77 Psammoclema goniodes sp. nov 78 Psammoclema nodosum (Carter) 79 Psammoclema radiatum sp. nov 80 Psammoclema ramosum Marshall 80 Psammoclema siipitatum sp. nov 81 Psammoclema vansoesti sp. nov 82

Mycalidae Lundbeck, 1905, sensu van Soest, 1984 83 Mycale Gray, 1867 83 Subgenus Arenochalina Lendenfeld, 1887 84 Mycale (Arenochalina) mirabilis Lendenfeld 84

Tedaniidae Ridley and Dendy, 1886 87 Tedania Gray, 1867 87 Tedania anhelans (Lieberkuhn) 87 92 Forcepia Carter, 1874 93 Forcepia biceps (Carter) 94 Lissodendoryx Topsent, 1 892 Lissodendoryx isodictyalis (Carter) 98 Ectyodoryx Lundbeck, 1909 J°0 Ectyodoryx maculata Hentschel I0°

Anchinoidae Topsent, 1928 102 Phorbas Duchassaing and Michelotti, 1864 102 102 Phorbas cf. tenacior (Topsent)

Order Haplosclerida Topsent, 1928 '03 103 Callyspongiidae de Laubenfels, 1936 103 Callyspongia Duchassaing and Michelotti, 1864 F. WIEDENMAYER

Callyspongia asparagus (Lamarck) 106 Callyspongia bilamellata (Lamarck) 106

Callyspongia diffusa (Ridley) 1 08 Callyspongia pergamentacea (Ridley) 109 Callyspongia persculpta sp. nov 110 Callyspongia ramosa (Gray) 112 Callyspongia serpentina (Lamarck) 115 Callyspongia toxifera sp. nov 117

Niphatidae van Soest, 1980 117 Gelliodes Ridley, 1884 117 Gelliodes incrustans Dendy 118

Oceanapiidae van Soest, 1980 119 Oceanapia Norman, 1869 120 Oceanapia putridosa (Lamarck) 120

Order Dictyoceratida Minchin, 1900 121

Spongiidae Gray, 1867 121 Spongia Linne, 1759 121 Spongia hispida Lamarck 121 Carteriospongia Hyatt, 1 877 123 Carteriospongia caliciformis Carter 124 Carteriospongia silicata (Lendenfeld) 126 Carteriospongia cf. vermicularis (Lendenfeld) 127 Coscinoderma Carter, 1883 128 Coscinoderma pesleonis (Lamarck) 128

Leiosella Lendenfeld, 1 888 1 30 Leiosella caliculata Lendenfeld 130 Leiosella levis (Lendenfeld) 132 Thorectidae Bergquist, 1978 134 ' Ircinia Nardo, 1 833 j 34 Ircinia caliculata Lendenfeld 135 Thorecta Lendenfeld, 1888 137 Thorecta choanoides (Bowerbank) 137 Thorecta glomerosus sp. nov 139 Thorecta tuberculatus (Carter) 140 Fasciospongia Burton, 1934 [ i 4 j Fasciospongia rimosa (Lamarck) \ \A\ Dysideidae Gray, 1867 144 Dysidea Johnston, 1842 ' 144 Dysidea avara (Schmidt) 144 Order Dendroceratida Minchin, 19 j 46 Darwinellidae Merejkowsky, 1879 \<\6 Aplysilla Schulze, 1878 145 Aplysilla rosea (Barrois) ] \A6 Darwinella Miiller, 1865 \\\ \Al Darwinella australiensis Carter \\\ 147 Darwinella gardineri Topsent ' ,[ 14 o Dendrilla Lendenfeld, 1883 /' ' ' 152 Dendrilla cactos (Selenka) " ,„

Order Verongiida Bergquist, 1978 ]54 Aplysinidae Hyatt, 1875 ' 154 SPONGES FROM BASS STRAIT

Aplysina Nardo, 1834 ]54 Aplysina ianthelliformis (Lendenfeld) 154 Aplysina lendenfeldi Bergquist 155 Druinellidae Lendenfeld, 1889 156 Pseudoceratina Carter, 1885 \%6 Pseudoceratina durissima Carter \S6 Druinella Lendenfeld, 1889 158 Druinella rotunda Lendenfeld 158 Notes on ecology j ^q Taxonomic diversity I ^q Zoogeography 163 Acknowledgements 1 70

References , 1 7 j Plates ' 1 g j '.'. '. '.'. '. '.'. '.'. '.'. '.'. '. ' Index '. '.'. '.'. '.'. '.'. '.'. 225

Introduction in concepts of genera and families, by authors In the initial stages of research on taxonomy, unfamiliar with the southern Australian fauna ecology and zoogeography of sponges inhabiting (except, after 1920, Bergquist, Burton, Dendy, Levi the continental shelf of southern Australia, at the et al., Topsent) has largely failed to incorporate this

Museum of Victoria (Melbourne, 1981-1985), it important fauna. became apparent that the basis for meaningful tax- To bring the literature up to date an annotated onomic work was very tenuous. The quality of bibliography and checklist of marine sponges of most published records of Australian species is Australia has been compiled (Wiedenmayer et al., inadequate for confident identifications, and the in press). The Bibliography contains 1520 anno- lack of illustrations and type data is almost general. tated references. The Checklist, which also covers For demosponges of the Maugean and Peronian Australia's overseas coasts and adjacent waters C provinces (Knox, 1960; defined for littoral faunas (Antarctica between 45 E and 160°E, including the of Victoria/Tasmania and New South Wales, French Adelie Coast; Heard and Macquarie respectively, here extended to the sublittoral fauna), Islands; Christmas Island) includes 1 173 valid spe- in particular, the primary literature on alpha- cies in 3 16 valid genera (993 species and 269 genera taxonomy dates mostly from before 1900 (Carter, of Demospongiae). The Checklist is based chiefly 1885-1887; Dendy, 1895-1897; Hyatt, 1877; on a survey of the literature, but includes some revi- Lamarck, 1813-1815; Lendenfeld, 1887a, 1888, sions based on type collections in Sydney, London, 1889b; Marshall, 1880; Polejaeff, 1884b; Ridley and East Germany. and Dendy, 1887; Selenka, 1867; Sollas, 1888). It The form of this report was dictated largely by is not only inadequate for identification, but its two constraints: (1) The aims of the research nomenclature (generic allocations) and synonymies project, as originally formulated, and (2) time and are badly out of date. availability of assistance with work in the labora- Subsequent revisions, some with redescriptions tory. A modern and thorough reappraisal, in pub- and illustrations chiefly of Lendenfeld's work lications, of the sponge fauna of the Maugean (Bergquist, 1980b; Burton, 1924a, b, 1927a, 1934a; province, even without the relatively well known Hallmann, 1912, 1914a-c, 1916a, b, 1917a, b, 1920; Calcarea (about 90 species recorded), would de Laubenfels, 1936a, 1948; Shaw, 1927a; Topsent, involve, in my estimate, at least 20 years of full- 1930a, 1932a, 1933; Vosmaer, 1911; Whitelegge, time work by a qualified taxonomist, under the best 1901-1907) alleviated the situation only in part, and circumstances. On the basis of data in Wieden- often brought new problems (especially with de mayer et al. (in press) and of extrapolation from Laubenfels). Besides these major revisions of the number of new species and new records in this demosponges of southern Australia, many isolated report I estimate that the number of species of revisions and relevant information (particularly re- demosponges inhabiting the Maugean province is examination of type specimens) are scattered in syn- about 500. onymies, discussions and footnotes, often in extra- It might have been preferable, for piecemeal limital literature. In any case, constant evolution coverage of the whole demosponge fauna, to pro- '

F. WIEDENMAYER

ceed by families or orders. But various circum- relevant data are indicated in the following list. stances (the author having started as a newcomer Field notes were taken as soon as practicable upon to the Australian sponge fauna; the state of previ- collection, but were sometimes delayed by sorting ous knowledge outlined above; lack of a complete of large volumes of sponges in dredge samples. set of micro-slides for the c. 1400 specimens from Colour photographs (Kodachrome) were taken of this province in the Museum of Victoria) suggested all specimens, and, whenever possible, notes on the present form as the only realistic one. Thus, colour using the Munsell Charts (edition for use a selection was made of newly collected specimens with plant tissues). Specimens were preserved in 10% from localities roughly aligned in a transect through formalin/seawater solution buffered with hexamine

northern Bass Strait (text-fig. 1). This had the (as described in Riitzler, 1978), and later transferred advantage that important field notes on fresh speci- to 70% alcohol at the Museum of Victoria. Riit- mens, particularly on colour, could be included in zler's method was also used for preparing spicule the descriptions. mounts and microslides with hand-sections. As Table 4 shows, the selection of material for description in this report is fairly representative of List of collecting stations this province. The most conspicuous gaps, in terms Text-figs 1, 2 of the relative importance of families and genera missed, are in the Suberitidae, Clionidae, Des- Tasmania, Kent Group (39°29'S, 147°20'E) macellidae, Halichondriidae, Hymeniacidonidae, between Wilsons Promontory and Flinders Island. Crellidae and Renieridae. The horny sponges are All specimens collected by divers (SCUBA). particularly well represented. The gaps are thought KG 1. 23 Mar 1981, Erith Island, West Cove, to reflect a very incomplete sampling of habitats. wreck of S.S. "Bullseye". Sponges among dense algal growth on hull, depth 10-12 m.—Jaspis stel- Collections lifera, Tethya ingalli, Rhaphoxya felina, The 151 specimens described in the following sys- Echinoclathria laminaefavosa, Mycale (Arenocha- tematic account were collected by, or in the lina) mirabilis, Ectyodoryx maculata, Callyspon- presence of the author. The collecting stations with gia pergamentacea, Amphimedon sp., Gelliodes - \a re

•

! .

Itt

U< H IHJ \

''" x (i8or--

o,KG « v

4c • A r \

Figure 1. Map of Bass Strait with stations of the Museum of Victoria Bass Strait Survey sampled in 1979-83 The localities, from which the material described below has been obtained, are circled. The Kent Group with stations KG 1-10, is represented in detail in Figure 2. SPONGES FROM BASS STRAIT

incrustans, Clathrina sp., Aulorrhiza procumbens, KG 5. 25 Mar 1981 , Garden Cove, N side of Deal Sycon sp. Island. Habitat similar to KG 2-4. -Ancorina KG 2. 24 Mar 1981, Deal Island, Squally Cove robusta, Jaspis stellifera, Latrunculia hallmanni, (E of lighthouse). Slope with granite outcrops and Clathria wilsoni, Echinoclathria laminaefavosa, boulders to 10 m depth, algal forest, large ascid- Ircinia caliculata, Thorecta tuberculatus, Aplysina ians Herdmania, urchins, some encrusting corals lendenfeldi, Druinella rotunda, Aulorrhiza Plesiastrea, solitary corals Scolymia).-Geodia procumbens, Sycon sp. (Geodia) punctata, Psammoclema nodosum, Teda- KG 6.26 Mar 1981, Winter Cove, E side of Deal nia anhelans, Reniera sp., Fasciospongia rimosa, Island. Habitat similar to KG 2-5. Depth 3-6 m. - Dendrilla cactos. Ancorina repens, Ancorina robusta, Ancorina KG 3. 24 Mar 1981, Deal Island, East Cove, W suina, Rhabdastrella cordata, Rhabdastrella of jetty of settlement, some distance from shore. intermedia, Jaspis stellifera, Latrunculia conulosa, Outcrops and boulders, - habitat similar to KG 2. Echinoclathria leporina, Echinoclathria egena Latrunculia hallmanni, Psammoclema vansoesti, Echinoclathria tubulosa, Phoriospongia argentea, Carteriospongia caliciformis, Leiosella levis. Phoriospongia kirki, Psammoclema callosum, KG 4. 24 Mar 1981, Erith Island, West Cove, Psammoclema vansoesti, Tedania anhelans, Force- S of 1. Boulders covered KG with algae, much pia biceps, Coscinoderma pesleonis, Leiosella kelp.- Latrunculia hallmanni, Echinoclathria caliculata, Thorecta tuberculatus, Dysidea avara. globosa, Tedania anhelans, Coscinoderma pes- KG 7. 26 Mar 1981, North Point (rocky promon- leonis, Dysidea avara, Aplysilla rosea, Darwinella tory) of East Cove, Deal Island, depth 25-30 m, gardineri, Leucetta microrhaphis. rock face below algal forest. Sponges, ascidians and

NORTH-EAST ISLAND Wallabi Cove

Pulpit Rock

East Bluff

Figure 2. The Kent Group of islands, with stations KG 1-10. F. WIEDENMAYER

gorgonians dominate. Some specimens from kirki, Psammoclema fissuratum, Psammoclema adjoining sloping sand. — Polymastia crassa, goniodes, Psammoclema radiatum, Psammoclema Latrunculia conulosa, Trachycladus laevispirulifer, stipitatum, Callyspongia asparagus, Callyspongia Clathria transiens, Echinoclathria leporina, Psam- diffusa, Callyspongia persculpta, Oceanapia moclema ramosum, Mycale (Arenochalina) putridosa, Spongia hispida, Coscinoderma pes- mirabilis, Ectyodoryx maculata, Spongia hispida, leonis, Aplysina ianthelliformis, Pseudoceratina Thorecta glomerosus, Thorecta tuberculatus, Dar- durissima. winella gardineri, Dendrilla cactos, Aplysina Taxonomic procedure lendenfeldi. There is now a discrepancy, in several points, KG 8. 26 Mar 1981 , East Cove, Deal Island, half- between the formal, "monographic" style of this way between jetty and North Point. Rocky level report, and its provisional nature (as the "first bottom. Large sponges. — Ancorina geodides, instalment" of a continuing Latrunculia hallmanni. project), a discrepancy that might easily mislead newcomers to accept all KG 9. 27 Mar 1981, E side of low-tide isthmus parts of the systematic section (identification, between Erith and Dover Islands ("Swashway"). synonymies, diagnoses, descriptions, discussions) Habitat similar to KG 2-6. — Clathriodendron cac- as "definitive" for some time to come. This proclivity ticutis, Echinoclathria carteri, Callyspongia is conditioned by the bilamellata, Callyspongia serpentina, Thorecta particular situation in choanoides. southern Australia (as outlined above), where a desolate, stagnant state, since the turn of the KG 10. 27 Mar 1981, W side of Dover Island, cen- tury, is suddenly improved cove SW of isthmus, (a) Granite boulders with algal by a major effort. A qualification of all these points is therefore in forest, depth 10 m.-Spirastrella papillosa. (b) Sub- order. merged cave, depth c. 5 m. /. Higher iaxa. The reviewers of the first version Wilsons Promontory, Waterloo Bay. of this systematic account have justly criticised the WB 28. 28 Mar 1981. Cliffs, vertical "faces, cave, lack or deficiency of this sort of material (diagnoses depth 12 m, collected by divers. — Corticum can- and discussions of families and genera, keys to delabrum, Polymastia crassa, Echinoclathria favus, genera with several described species). A part of Carteriospongia silicata. this problem resides in Wiedenmayer et al. (in press) BSS 179. 18 Nov 1981. 39°03.2'S, 146°39.5'E (20 which I had hoped might be published now provid- km E of Station WB) depth 55 m, bottom muddy ing a complement in some of these respects. Its pub- sand. - Phoriospongia carcinophila, Psammoclema lication seems to be delayed. bitextum, Psammoclema densum, Callyspongia Diagnoses and discussions offamilies. Diagnoses toxifera. are included in Wiedenmayer et al. (in press). I have BSS 180. 18 Nov 1981. 39°12.9'S, 146°27.3'E (7 preferred not to repeat them here, which means km S of Wilsons Promontory), depth 65 m, bottom that, for this purpose, I mostly adhere to the defi- muddy sand, sand and shells, fine algae.— nitions given by Levi (1973), Bergquist (1978, Callyspongia ramosa, Carteriospongia cf. ver- 1980b), van Soest (1980, 1984), and Hartman micularis, Darwinella australiensis, Darwinella (1982). The family Desmacididae is revised, with gardineri. the two subfamilies Desmacidinae (Esperiopsidae BSS 181. 19Novl981.38°39.8'S, 144°18.2'E (30 of authors) and Stylotellinae, the latter compris- km SE of Lome), depth 79 m, bottom very fine ing the sandy genera Phoriospongia (Chondropsis sand with many branching, parchment-like worm and Psammascus of authors), and Psammoclema tubes (Chaetopteridae), on which most sponges col- ( = Psammopemma). Lengthy discussions are lected are encrusting or attached. -Strongylacidon provided for the Latrunculiidae and for the Myx- ste/liderma, Stylotella inaequalis, Phoriospongia illinae in the Tedaniidae (here chiefly regarding Lis- carcinophila, Psammoclema densum, Psam- sodendoryx). Several families in the Axinellida, and moclema goniodes, Tedania an he/cms, Lissodendo- Poecilosclerida (particularly the Microcionidae), ryx isodictyalis, Phorbas cf . tenacior, Callyspongia and genera in the Dictyoceratida (particularly Phvl- toxifera, Darwinella australiensis. lospongia/Carteriospongia, BSS and Coscinoderma) 187. 20 Nov 1981. 38°32.0'S, 142°28.6'E (17 need to be reappraised. But these happen to be km S of Warrnambool), depth 52 m, bottom coarse poorly represented in the material here covered, and sand and shells.— Ancorina robusta, Tetilla such revisions are better undertaken praecipua, in special Spirastrella papillosa, Chondrosia studies by my colleagues in northern Australia. reticulata, Pseudaxinella decipiens, Reniochalina Synonymies and diagnoses genera. sectilis, of Some diag- Rhaphoxya cacliformis, Phoriospongia noses are quoted from previous accounts by other SPONGES FROM BASS STRAIT

authors unchanged, some with additions, others stage. They would contradict the preliminary nature with alterations, and some are new. The variable of this report, by contributing essentially to a false

lengths tend to reflect the degree of difficulty at impression of definitiveness. It is imperative that this level. As a guide to the evolving concept of a reasonably complete sampling of the local fauna genera, I have adopted annotated synonymies simi- is worked up, whenever possible with more than lar in style to those used by me earlier (Wieden- one specimen per species, before keys are mening-

mayer, 1977a). ful here. I am convinced that many more new spe- Unavailable and unused names of the genus-and cies and records of Stylotellinae, Microcionidae species-groups (nomina nuda, nomina vana, (particularly Echinoclathria), and Callyspongiidae nomina nulla, nomina oblita, and junior primary will eventually emerge in southern Australia. In this homonyms) are cited in square brackets. Such Latin context, all relevant type slides in the AM, BMNH

terms for name categories denoting criteria of valid- and ZMB will have to be evaluated, which I could ity and availability, are used and defined in volumes not do.

of the Treatise of Invertebrate Paleontology (see Type species. All relevant information is in Loeblich and Tappan, 1964). Definitions of these Wiedenmayer et al. (in press). In some cases with terms are found again in the glossary below. They errors and misunderstandings, clarification are here are more amply defined and discussed, with respect added under Remarks following generic diagnoses. to Porifera, in Wiedenmayer et al. (in press). In

rewriting generic diagnoses, I often felt that old 2. Species. ones simply did not work with fresh material, and Identifications and synonymies. Some syn- that a modification or two in such diagnoses (and onymies are extremely long, and have as such raised an occasional merger of genera) were more sensi- doubts with the reviewers. There are two kinds of ble than trying to explain a "bad fit" or ambiva- such synonymies: (1) Those of cosmopolitan or lence of a little known or new species under widely disjunct species, headed as "Selected syn- "Remarks" following its description. It should be onymies" (I have avoided "restricted synonymy", considered that more time for collecting in the same as used by others, because this might imply a localities here covered would probably have deliberate confinement and revision with respect

produced dozens of other records, including sibling to earlier synonymies, while I give a broad sam- species of those described, which might in turn pling, including cross-references), and (2) those of necessitate further changes in generic diagnoses. polymorphic Australian or austral species, in which With only one-seventh of the estimated total most synonyms are names of Lendenfeld. provincial demosponge fauna covered in this In the first case, the doubts as to the reality of report, such speculations are legitimate. extremely wide distribution (as in Corticium can- Keys. These were suggested by the reviewers par- delabrum, Tedania anhelans, Lissodendoryx isodic- ticularly for Echinoclathria, Psammoclema, and tyalis, Dysidae avara, Aplysilla rosea, Darwinella Callyspongia. Keys are problematical with sponges, australiensis, D. gardineri) are probably tinted by mostly because, unlike other invertebrates, they are an exclusively actualistic outlook, by comparisons so difficult to put consistently into a hierarchic with biogeography of higher, more stenotopic framework, also because keys tend to mask the invertebrates, and by the argument that morpho- enormous differences in degree of variability, and logical characteristics in such species are too vague because several important regional faunas are too and variable, and therefore mask "real" biological poorly known for confidence (e.g., eastern tropi- differences. As long as such "differences" are not cal Pacific, South Atlantic, southern Australia). unequivocally demonstrated, it seems more sensi- Keys, for sponges have to rely only or chiefly on ble to regard such populations as conspecific, if not microscopical, skeletal traits, especially spicule panmictic, and not to use open nomenclature or morphology. They are therefore of little if any use even new species names for such austral records. to general biologists, who are not used, or unwill- It is quite plausible that those constitute relics of ing, to prepare two kinds of mounts in Canada late Tethyan distribution, more so than I argued balsam for every specimen (thick sections and dis- earlier (Wiedenmayer, 1974, 1977a). The disjunct sociated spicule mounts). Sandy sponges constitute Brazilian/ Central Atlantic/ South Atlantic/ Indian a special problem, because "teased" or sieved Ocean/ austral distribution of some stenotopic ele- mounts are often necessary. Nevertheless, keys are ments has palaeobiogeographic roots (e.g., the spe- desirable for newcommers to the sponge fauna of cies Callyspongia pergamentacea and the sibling southern Australia, particularly those with previ- pair Aplysina lendenfeldi/A. pedunculata in this ous experience in sponge taxonomy; but not at this report; Tedania commixta fide Topsent (1904); Ill F. WlbDI.NMAYIU

<". serpentina, all polymorphic and Hymedesmia mertoni I lentschel, 1912, fide Boury- C. ratnosa,

I sufficient con- Esnaull and Lopes (198S); Asteropus simplex, superficially often similar , gained Tethya japonica, and three species of Phoriospon fidence from examining the lypc specimens; but the yja (as I'sununoclwhi and Psuininotoxu), fide ultimate control, with slides, had lo be foreclosed Boury-Esnaull (1973); furthfti species arc quoted due lo lack ot time (as with all other type material by Hechlel, 1976; 249). listed as examined). I therefore headed these syn- The annotations in these synonymies will serve onymies as "provisional", preferring not to mark as guides and aids In future revisions and evalua- all entries with question marks indiscriminately, tion of new records', and ilnis to better diagnoses, masking the really doubtful synonyms, of which by pointing <>ni the more relevant informations and I was unble to see type material. their nature. Unfortunately, many published In these cases, the annotations with most refer- records of such species, especially from I he type ences should be helpful lo future revisers, allow- province, lack supplementary information on mor- ing for rapid sorting out of old and new records, phology, which is probably due lo the assumption it-descriptions, type stains, etc, especially in con- thai they are sufficiently known from erlier nection with Wiedenmayer el al. (in press). descriptions. Diagnoses. These were added, somewhat reluc-

In the second case, I have been criticised lor rely tantly, al the suggestion of the reviewers and of the ing excessively on Bur Ion's revisions, which arc edilor. The provisional nature of this report is espe- notorious for supposedly indiscriminate lumping. cially conflicting with its style in the apparent con- This argument is far loo generalised. Burton some- sistency of these diagnoses. The most reliable ones times erred in the opposite direction (sec my com- will probably be those for endemic, sienotopic spe-

I ment on / issodendoryx ternatensis, . sinensis, and cies represented by several specimens in (he collec- /.. similis). Even in his most crilicised exercise in tion, and described previously. The least reliable lumping, his revision of all (alcarea (Burton, will be those for new species, and new records of 1963), the extensive mergers of species and genera rare species, with only one specimen in the collec- are largely theoretical, without loss to practice. In tion. Diagnoses for eurytopie and polymorphic spe- effect, all previously accepted species anil records cies (frequently parallel qualities, as with oilier are carefully redefined as "named forms" and syn- invertebrates), and those for cosmopolitan species, onyms, on I he basis of rc-cxaminalion of primary are, by nature, longer ami more likely to need and secondary types (including slides), for which Updating in details. The latter are obviously based a detailed list is given as an appendix. The same chiefly on various published sources, and will (espe- painstaking procedure, though less well docu- cially with Tedania anhelans and Lissodendoryx in mented prim, characterised the revisions of most isodictyalis) cover a broadei spectrum than is likclv "< of I endcnlcld's (1887a) halincac" by Bui Ion to be encountered in records from southern (1927a, 1934a). Australia.

Anyone thoroughly acquainted with I endcn- Descriptions. Many of these may be criticised as lcld's Australian type material (of which I have seen being loo long, and detailed. They can, however, virtually every specimen in the AM and BMNII) be justified with the excessive brevity o\' most of and with faulty his idiosyncratic and methods criti- < arter's and Dendy's descriptions for this area, the cised chiefly by Whitelegge and Dendy (Wiedcn Often misleading nature of I endenfeld's, (he rich- el at., in will have mayer press) come lo similar ness of this fauna, and the presence Of many sibling conclusions regarding his extreme proclivity for groups. ( )lher siblings of species here described arc splitting, and Ins negligence in observing oi and known lo occur in the area only through cailv describing microscopical features (though he could, descriptions (see Remarks following descriptions, at times, be accurate). A striking,, hitherto nine ami Wiedenmayer el al., in press), and remain to vised example of such negligence is Clniliiutla coxi be recollected and redescribed. Especially with

I cndenfeld (1885c), a junior synonym of I he chic- sibling pairs with several specimens in the collec- ft} estuarine Tetilla dactyloidea (Carter), of wide tion (as in Latrunculia), I preferred to err on the Indo Pacific distribution. side of verbosity, rather than suppressing details, With some o\' my synonymies involved because so little is still known on local-inlrnspecific. (Echinociathria lantinaefavosa, Phoriospongia biogeographicai and ecological differentiation of it kirki, Spongia hispida), isalreadj clear from I en- morphological characters, particularly outside the denfeld's descriptions, thai his distinctions do not best known provinces, such as the Western Mediter- hold up lo modern laxouonue standards. In (he ranean. Details, which might seem trivial now. may case of ( < 'allyspongia diffusa, '. pergamentacea, someday be evaluated in a different perspective. SPONGES I KOM BASS STRAIT II

Supplementary information. Dala on distribu- the same collection could be obtained For study (pi. tion are incorporated into the annotated syn- 38 RgS I 5). onymies. Synopses of world-wide distribution, by Terminology mid glossary provinces and localities For Australia, by regions Conventional morphological terms are mostly elsewhere, and depth ranges are in Wiedenmayer those explained in my illustrated glossary (Wieden- et al. (in press). mayer, 1977a; 30 51). I have, in addition, used some terms which have been criticised as old fashioned or otherwise rarely used. Some of these Other material examined. Virtually all Australian are revived from early descriptions (such as type specimens and hypotypes relevant to this C'arler's), from lime when spongologists were also report of the following authors and publications naturalists, publishing in several disciplines, and could be examined: I .eiuleufeld (1885f, h; 1886a had no qualms about using terms borrowed from e, f; 1887a, 1888, 1889b) in Sydney (AM) and botany. I find these apt ami implicitly descriptive London (BMNH); Carter (1885a-1886h, 1887a, ex lor recuriing features. Though most of these terms J.B. Wilson eollection; 1879b, 1881b, 1882a, aie defined in good standard English dictionaries, 1883b, c, mostly ex Bowerbank collection; all in 1 often use them in a more restricted and special BMNH); Dendy (1895-1897, NMV); Bowerbank sense. (1844b, 1872, 1873a; BMNH); Gray (1843, Alveolate, finely honeycombed on surface; in BMNH); Burton (1934a, BMNH); Whitelegge and clioanosome, regularly cavernous, With round oi Hallmann (AM); Marshall (1880, PM.I and /MB); rounded-polygonal interstices delimited by con some relevant ones of Polejaeff (1884b, BMNH), toiled Irabeculae or walls. Ridley and Dendy (1887, BMNH), Ridley (1884a c, ( (mediate. I alliced al right angles, cioss barred. BMNH), Kirkpatrick (1903b, BMNH), Shaw (erehroiil, Biain -like pattern of the surface. (1927a, BMNH), Sollas(l888, BMNH). At the time (lat/irale. I alliced. of ray survey of these type collections (AM in 1982; Concrescence. Lateral fusion, mostly ol BMNH, PM.I and /MB in 1483), which also branches, lamellae, or lubes. included species and papers no( covered in this CryptS. Widespiead interconnected vestibules report, most specimens were photographed. The between a thick, firm eclosome and the clioano- negatives and prims are now kept in the Museum some (subcortical), or within the inner portion of of Victoria Sponge Archive (Department of Inver- the eclosome (inlracorlical). tebrate /oology). Some of these are here Deadline vermiculale. Wrinkles al the surface reproduced in the plates (pi. 2 figs 6, 9; pi. 3 figs or subdermal canals have a continuously winding 2-6; pi. 4 figs 2-5, 7 12; pi. 5 figs 13; pi. 6 and blanching course. 4-7, pi. 7 ligs 11; 4 7, 10, 11; pi. 8 figs 4, 8 11; Diataxonic. Applies to synapoinorphies in higher pi. ligs 4-6, 9 13; pi. 10 figs 4-7, 10, II; pi. 1 1 ligs laxa, wilh accent on the uselessness of such cliaiac 1, 2, 7, 9, 10; pi. 12 figs 2, 3, II, 12; pi. 13 figs lers in taxonomy.

2, 3, 6, 7, II, 12; pi. 14 figs I 5. 10; pi. 15 ligs Excrescence. Outgrowth. 1-5, 7, 10; pi. 16 figs 2, 3, 6, 7; pi. 17 figs 2, 8). Fenestrate. Riddled; perforated by similar small Unfortunately, various circumstances prevented me gaps.

from examining the type slides involved in [he AM, Heteropolar, Denotes the general condition ol BMNH, and /MB. unequally ended pseudasters and ornamented The indications for each species, under the head- microrhabds (e.g., anisodiscorhabd). ing Material examined, following the new material, I.acinules. Small, often compressed and prolific now contain only valid species names and syno- lobes or digitations slicking oui from the surface, nyms in original combination, and usually type Lacuna. Oap, window, conspicuous hole other status. Other details (number ol specimens, register than an oscule oi pseudoscule. numbers, reference to film/frame numbers in the Lipogastric. Devoid ol atria. Museum of Victoria Sponge (Photo) Archive) are I. itlae. For incorporated fragments derived from found in Wiedenmayer et al. (in press). The type- rocks (through uptake from bottom sediment). specimens in East-Berlin (/MB) were nol available Monilijonn. Shaped like a siring ol beads. for study at the lime of my visit there, with two Nervines. Denotes I he pattern, m the dcimal exceptions: the holotypes of Phoriospongia retic- membrane, of mocroscopic collagenous ribs or ulum Marshall (pi. 6 fig. 6; pi. 24 fig. 3) and ol bands, often anastomosed, like the veins of a leaf. Psammopemma densum Marshall (p. 8 Rg. 4). One Nomen correetuin. Emendation Of a iii, men of I endenfeld's slides from Druinella rotunda in impel ledum. 12 F. W1EDENMAYER

Nomen dubium (genus dubium, species dubia). Spicular vicariance. The tendency, within some Name of unknown or doubtful application. families and genera (particularly in the Desmacid- Unrecognisably described species (type species) of idae and Tedaniidae), for pairs of principal which all type material is known to be lost, and megasclere types (diactines/monactines, acan- for which the fixation of a neotype is unadvisable. those/smooth), which are consistently segregated Nomen imperfectum. Valid name needing emen- in most other taxa, to pass from one state to the dation (when not meeting ICZN, Articles 26-29, other, mostly mutually exclusive between genera 31); particularly common with adjectival species or congeneric species, less often within populations, epithets and names of higher taxa deficient in or mixed in variable proportions, with transitions endings. (ambivalence). Spicular vicariance may be regarded

Nomen inquirendum (genus inquirendum, spe- as a special form of homology, but its phylogenetic cies inquirendd). Name of lower taxon of doubt- significance (especially in Tedaniidae) is not clear. ful application on the basis of description(s), but Stipitate. Stalked. of which type material (of type species) is known Stolon. A single repent (trailing) branch. to be accessible for redescription. Subtylote. Tylote with indistinct heads. Nomen negatum. Incorrect (unavailable) origi- Sulci. Furrows. nal spelling (ICZN Article 32c, d). Commonly Terete. Basically cylindrical, but commonly applied to those alternative forms in multiple origi- winding or crooked (when not applied to nal spellings eliminated by fixation of the correct megascleres).

original spelling by the first reviser. Trabecula. A choanosomal structural element in Nomen nullum. Incorrect subsequent spelling form of band or bar; an angular or compressed Article (ICZN 33c). tract, fibre, or portion of matrix, without definite Nomen oblitum. Forgotten name. Cited only as orientation. "a descriptive term" in the glossary of the ICZN Umbel. Term borrowed from botany, denoting (3rd edition, 1985: replaced 260), and in the main the pattern of multiple branches radiating from the text (Articles 23b, 79c) "unused by senior syno- end of a stalk (ascending fibre) at variable angles, nym." See Declaration 43, 1970, Bulletin Zoo- of occasionally subdivided, and ending more or less logical Nomenclature 27: 135-163, and comment in one plane. Here used for the peripheral skele- in Wiedenmayer (1977a: 52). ton of many Astrophorida and Hadromerida. Nomen vanum. Unjustified emendation (ICZN Vermiculate. Worm-like; winding but dis- Article 33 b(iii). continuous. Pedicel, peduncle. commonly small, short, A Verrucose. With scattered wart-like elevantions. stalk-or foot-like base. Size of Penicils. Brushes, especially of bundled or radi- specimens and its gradation ating megascleres, commonly not bound by spon- A gross subdivision into three categories is occa- gin, flush at or projecting from the surface. sionally used in diagnoses and descriptions. In my Polytylote. For megascleres having several annu- experience with statistics (most frequent sizes) of lar swellings along their shaft. Frequent in Bahamian sponges (Wiedenmayer, 1977a, most styles/tylostyles of Poecilosclerida, also in some tables in the systematic-descriptive part), the fol- Hadromerida. lowing subdivision seems appropriate (except for Rhaphidotoxon. An irregular, asymmetrical thinly encrusting forms): toxon, or irregularly sinuous rhaphide. Small. 2 Up to 5 cm in greatest diameter (25 cm Rhizome. Root-like structure. greatest projected area). Rugo-reticulate. The folds or wrinkles affecting Medium. Mostly in the order of first-size (100 the surface are combined into a net-like pattern, 2 2 cm ), up to 250 cm . with triangular, square, or polygonal depressed Large. Mostly in the order of head-size, and interstices. larger. Rugose. Of a surface provided with numerous wrinkles or folds (rugae). Rugo-vermiculate. The folds or wrinkles of the Abbreviations surface are discontinuous and have a winding AM, Australian Museum, Sydney. course. BMNH, British Museum (Natural History), Schizotype. Significant fragment or section of London. a type specimen. In 1977a, I attributed the term F51927, example of identifier (NMV register to W.D. Hartman. It was already used by Annan- number) for a new specimen described and or dale (1915c: 469). figured. -

SPONGES FROM BASS STRAIT 13

ICZN, International Code of Zoological Nomen- clature, third edition (1985). Mybp, million years before present. MJG, Landesmuseum Joanneum Graz (Austria), Abteilung fur Zoologie. MNHP, Museum National d'Histoire Naturelle, Paris. MZUS, Musee Zoologique de l'Universite Louis Pasteur et de la Ville de Strasbourg, France. NMV, Museum of Victoria, Division of Natural History (formerly National Museum of Victoria). Figure 3. Corticium candelabrum, spicules, x 320. a. Can- delabra, with range and mean of height, b. PMJ, Phyletisches Museum der Friedrich- Microcalthrops, with range and mean of ray lengths. Schiller-Universitat, Sektion Biologie, Jena, GDR. SMF, Natur-Museum Senckenberg, Frankfurt am Main, FRG. candelabra, less frequent simple microcalthrops. ZMB, Museum fur Naturkunde (Zoologisches Spicules packed below surface. Museum) der Humboldt-Universitat, East Berlin, Small compressed lobes with rounded GDR. Description. margins, sometimes coalescent, with irregular ZMH, Zoologisches Museum der Universitat, 2 specimens ear-shaped and -sized. In life Hamburg, FRG. folds; light brown (2.5 YR 8-9/4-8) with tinge of orange 2.5 YR 8/10, example of colour notation from in F51928 (2.5 YR 6-8/8). Firmly rubbery, smooth, the Munsell Colour Charts. slippery. Ostia regularly scattered on both faces; Italics in spicule measurements denote means. oscules on and near margins. Class Demospongiae Sollas, 1885 Atria and excurrent canals in mesial region; incurrent canals regularly parallel and radial. Subclass Homoscleromorpha Levi, 1953 Spicules packed in cortical region (250-500 ^m Order Homosclerophorida Dendy, 1905 thick); mostly scattered in choanosome, frequently aligned and more crowded along incurrent canals; Plakinidae Schulze, 1880 packed around excurrent canals. Microcalthrops Corticium Schmidt less frequent than candelabra; relatively rare in in F51928. Corticium Schmidt, 1862: 42.-Lendenfeld, 1903: F51927, more frequent 1 of rays in 12L-Thomas, 1970d: 260 (Indian Ocean records). Spicule measurements (F5 927): lengt Pulitzer-Finali, 1983: 448, 602 (Mediterranean species). microcalthrops, 28.5-52-38.2 \nm\ height of candelabra: 27.1-36. 7-42.1 fim. Few smaller, develop- Diagnosis. Plakinidae in which candelabra are mental forms and malformations occur. usually present, without diactines. Microcalthrops may be simple or lophate (equally or irregularly). Remarks. The other species of Corticium known from Australia, C. simplex Lendenfeld (1907a) Corticium candelabrum Schmidt from the North-West Shelf, is described as having

Plate 1 figure 1, plate 18 figure 1, text-figure 3 candelabra only and a peculiar vescicular structure of cortex and parts of the choanosome. A speci- Selected synonymy. men from the Northern Territory shown to me by Corticium candelabrum Schmidt, 1862: 42, pi. 3 fig. J.N. A. Hooper (Darwin) agrees in vescicular struc- 25 (Adriatic Sea).-Lendenfeld, 1903: 122 (synonymy, cortex and parts of the choanosome but diagnosis, distribution). -Burton, 1934a: 521 (synonymy, ture of distribution; Great Barrier Reef). - Rutzler, 1965: 15 (syn- contains simple microcalthrops. The other three onymy, ecology; Adriatic Sea). -Thomas, 1970d: 261 Mediterranean species known, C. bowerbankiSarh, (India). -Boury-Esnault, 1971: 295 (ecology; western C. topsenti Pouliquen, and C. reductum Pulitzer- ecol- Mediterranean). -Pulitzer-Finali, 1972: 350 (colour, Finali (see Pulitzer-Finali, 1983: 449) have more - 1972: 726, 746 ogy; western Mediterranean). Pouliquen, complex microcalthrops, and so has C. acant nos- -Pulitzer-Finali, 1983: 448 (western Mediterranean). trum Thomas, 1970d, from India. (ecology, colour; western Mediterranean). Burton's material of C. candelabrum from the specimens Material examined. Station KG 1 0b, three Great Barrier Reef came from an open-marine (F51927); station WB, one specimen (F51928). environment (presumably level bottom), 25-30 m

depth. It is interesting to note that our material Diagnosis. Small, comprcssed-lobate, light brown; from submerged caves, like many of the firmly rubbery, smooth, slippery. Oscules margi- came Mediterranean records. nal. Choanosome dense, with abundant scattered 14 F. WIEDENMAYER

In Mediterranean specimens, ray lengths I of Stellei 'inopsis Cater, 1879b: 348. -Bergquist. 1965: microcalthrops are given as 30-40 /xm, maximum 189. -Bergquist, 1968: 32. diameter (height) of candelabra as 27-32 ^m (Len- Psammastra Sollas, 1886: 194. -Sollas, 1888: cxli, denfeld, Pulitzer-Finali). Lendenfeld's description 174.-Thiele, 1900: 36.-Dcndv, 1905: 80. Algol (1903) does not significantly depart from mine, nor Sollas, 1888: cxli, 200 (objective svnonym of Stellettinopsis).- Bergquist, can I see any sensible discrepancies in published 1965: 189. Thalassomora figures of spicules of Mediterranean specimens. Lendenfeld, 1888: 40. -Lendenfeld, The 1903: 62 (synonymised with Ecionemia). disjunct distribution of C. candelabrum is Sanidastrella Topsent, 1892c: xviii.- Lendenfeld, gested by the presence of candelabra very similar 1903: (as 58 subgenus of A ncorina).- Levi (in Brien et al., syn- to Recent examples in the Upper Eocene-Lower onymised with (Ancorina). lar to Recent examples in the Upper Eocene-Lower Diagnosis. Oligocene Oamaru Diatomite of New Zealand Aneorinidae with microrhabds or sanidasters in addition (Hinde and Holmes, 1892, pi. 12 figs 12-15). Fur- to euasters. Microrhabds commonly minutely thermore simple and Iophate, regular and irregu- spined or roughened, forming lar fossil a dermal layer. Euasters may comprise microcalthrops (same plate) make it oxyasters. virtually certain that the candelabrum and bower- Remarks. Many authors have distinguished Anco- banki groups were already differentiated then rina and Ecionemia on the grounds of within cortical Coriiaum. As will be shown in the conclud- sanidasters in the former, cortical ing chapter, roughened the Oamaru fauna lived in shallow, microrhabds in the latter. I agree with warm water, contrary Dendy to previous opinions (includ- (1916a: 239; 1924: 296) and Bergquist (1968: ing mine), and the associated 32, Foraminifera point 33) that this difference is insignificant. Levi (in to immigration from the tropical Indo-West-Pacific Brien et al., 1973: 595) grants this basic region along the Papuan Geosyncline, similarity during a but uses (like Lendenfeld, period 1903) presence/absence of exceptional warming at higher latitudes. of oxyasters as diagnostic criterion. A centre of origin of Corticium Considering the in the Indo-West- variability and diversity of choanosomal euasters Pacific region is not likely, however. According to in Astrophorida, Hinde this criterion seems irrelevant and Holmes (1892: 228), candelabra "are (Wilson, 1925: 297). The already known from diagnosis is here revised the (Upper) Jurassic of accordingly. Poland. The genus may therefore have originated in the western Tethys and may have been affected Ancorina geodides (Carter) by later radiation in the eastern Tethys or Plate early 1 figure 2, plate 18 figures 2-3, Indian Ocean. As is the rule with long-lived, widely text-figure 4 distributed genera, Corticium was and is probably Steiktta geodides more eurytopic than Mediterranean records Carter, 1886b: 125 (Port Phillip sug- Heads). t gest (where dominance of inclined and recessed Psammastra geodides. -Sollas, 1888: rocky sublittoral substrates resulted 200 (redescrio from the par- tion, discussion). ticular Late Neogene tectonic setting of this region), Ancorina (Sanidas,rella) geodides. - Lendenfeld, 1903- though it may always have preferred shallow water.' 60 (resume). Stelletta geodides. -Dendy, Subclass Tetractinomorpha Levi, 1953 1905: 80 (discussion). Material examined. Station KG 8, one Order Astrophorida Sollas, 1887 specimen (F51929t only a segment kept. Sketch by Carter, in his Manuscript Aneorinidae Schmidt, Catalogue (BMNH). Unpublished 1870 specimen from M a fa sland, Tasmania, BMNH Synonym: 1925.11.1.1315 = RN XVIII Sicllctiidae Carter, 1875 (wet), .dentified by Dendy as Ancorina geodides. Aneorina Schmidt Diagnosis. Massive, cake-to dome-shaped, hard Ancorina Schmidt, 1862: 51.-Dendy, 1924: 296 Surface dark grey to black, (syn- may be faintlv tuber- onymised with Ecio temia).- Wilson, culate, 1925: 286 with minute acanthomicrorhabds:"oscules (resume). -Topsent, 1928: 35.- Bergquist, 1965: mconsptcuous. 189 Thick cortex detachable, (synonymised with Ecionemia). — internally Bergquist, 1968: 32 (syn- hght grey. Choanosome onymised with Ecionemia). dense, buff. -Levi (in Brien et al., 1973): 595 (retained as distinct genus). fefr&ion. massive, 15-20 cm in diameter, cake- Ecionemia Bowerbank, 1863c: to dome-shaped. HOL-Thiele, 1900: Lower half encrusted 28.-Lendenfeld, 1903:61 bv platelets (as subgenus at Ancorina) - Xt rnally y dark °- Dendy, 79. br ' '« 1905: -Wilson, 1925: 296 (resume, discussion, duckla f MpyxYR $?£1/3-4), also in alcohol.r probably a synonym of Ancorina). - Levi (in Brien et al.' Hard Surface 1973): 595 (retained smooth in life, now finely as distinct genus). and rugose microtuberculate with low magnificat SPONGES FROM BASS STRAIT 15

«^P yellowish grey (7.5 YR 9/1-4); easily detachable, with continuous zone of subcortical crypts. Inner cortex coarsely fibrous (collagenous), traversed by radial chones. Chones about 270 /tm wide, regularly spaced 0.7-1.5 mm, communicating with subcortical crypts, with sphincter somewhat below top of inner cortex. Lining of chones similar to outer cortex and to choanosome with many stron- glylasters and acanthomicrorhabds, usually in clusters. Acanthomicrorhabds scattered in fibrous cortex; asters rare here. Outer cortex strongly pigmented, about 500 ^m thick; with many strongylasters, and, in outermost portion, crowded acanthomicrorhabds. Choanosomal skeleton comprises megascleres mostly radially arranged, with cladomes of triaenes staggered in narrow zone below subcortical crypts. Deeper parts more confused, with many magascleres at various angles, singly and in bundles. Bundles traverse peripheral choanosome and cortex, usually expanding slightly in inner cortex forming narrow penicils ending at surface. Choanosomal aster, regularly scattered interstitially, numerous. Spiculation: (1) Oxea straight, mostly fusiform 1334-/775-2080 X 12.1-27-27.7 /xm; (2) orthotriaenes with small, often irregular cladomes (clads may be crooked, reduced, suppressed or follow slightly different angles), rhabds 662-/550-2026 x

9.5-2Z-26 urn, clads 39-46.7-58 ^m in length. (3) Choanosomal oxy-and strongylasters with 10-14 rays, diameter 12.7-/7-24.2 fim, often with a centrum occupying 15-33% of the diameter. (4) Cor- tical strongylasters (13. 8-16. 5-2\. 2 urn, 8-14 rays) intergrade with latter but usually lack centrum, frequently roughened and spined, especially distally. Some large ones have tylote to lanceolate (mucronate) rays. (5) Acanthomicrorhabds, transitional to sanidasters, 5.6-7.6-8.5 x 0.9-2.4-3.2 ^m.

Figure 4. Ancorina geodides, spicules. As with all fol- Remarks. The body of holotype must be regarded lowing figures of spicules, ranges (minimum, usually as lost. All that remains in the British Museum mean, and maximum dimensions) are indicted with (Natural History) is a flimsy sketch by Carter and separate scales for each category, a. Oxea (with example a slide made by Sollas, presumbably from the holo- length), Orthotriaenes, x68, withcla- of mean x68. b-c. type (BMNH 1/28). Sollas (1888) seems to have dome enlarged, x 263. d-f. Choanosomal oxyasters and based his redescription chiefly on "tracings illus- strongylaster, x 790. g. Cortical asters with lanceolate ray- trating the structure of this sponge obtained from tips, x790. h. Acanthomicrorhabds, x 1450. Mr. Carter." Except for the conflicting measure- Oscules inconspicuous, 0.2-0.3 mm in diameter, ments of thickness of the cortex and its parts, given round to elongate, irregularly scattered, some by Sollas, and the absence in the new specimen of clustered. the "low mulberry-like elevations", there is good Choanosome dense, buff-coloured (5 YR 7/6-8 agreement. Carter's specimen was much smaller to 5 YR 8-9/4), criss-crossed by excurrent canals than the new one and it seems likely that the ele- 1-2 mm wide. Cortex is 2-3 mm thick, of 2 dis- vations in question, if always present in young tinct layers, the outer almost black, the inner light specimens, disappear with growth. The BMNH 16 F. WIEDENMAYER

specimen, from Maria Island, Tasmania, agrees (1 a well with the present specimen in macroscopic- traits, and also lacks the regular round elevations. Ancorina geodides seems to be closest to A. tnur- rayi (Sollas), a species known only from Bass Strait, particularly in spiculation. As Sollas (1888) already

remarked, his species differs chiefly in its conulose surface, and in its megascleres being much larger.

Ancorina repens sp. nov.

Plate 1 figure 3, plate 18 figures 4-6,

plate 19 figures I, 2, text-figure 5

Holotype: NMV F51930, Station KG 6 (Tasmania, Winter Cove, E side of Deal Island, Kent Group; granite boulders with algae, depth 3-6 m).

Diagnosis. Slug-shaped with short pedicels, very firm. Surface rough, finely rugose, greyish brown, mottled. Oscules inconspicuous, scattered. Cortex thick, stratified, with abundant megascleres in crowded to overlapping umbels. Choanosome dense, light brown, with megascleres in confusion, with crowded euasters.

Description. Slightly compressed stolon of uneven

thickness, four points of attachment at base. 1 1 cm long, 2-2.5 cm high, 1.5 cm wide, bluntly pointed. in life Colour dark greyish brown, mottled . Very firm, little compressible. Surface slightly rough, potato-like. Faint, finely dendritic-vermiculate pattern on top. Oscules inconspicuous, numerous, 100-300 /im wide, regularly scattered on top. On upper portion, vermiculate-anastomosing, tangential canals below surface, 0.5-1 mm wide, roofed by smooth outer portion of cortex. These canals are not completely open, but contain narrow widely spaced penicils of mgascleres, supporting the dermis. Surface elsewhere regularly microtuberculate to rugo-reticulate, reflecting cortical penicils of magascleres. In few areas, tangetial magascleres or their bundles form subparallel lines. Ostia regu- Figure 5. Ancorina repens larly placed in depressed interstices of fine surface sp. nov., spicules, a. Oxea (with example of mean length), pattern. x 65. b. Triaenes, x65. c-e. Enlarged cladomes, x249. f. Peripheral strongyles, x65. Choanosome light brown, dense; irregularly g. Acanthomicrorhabds, x748. meandering and h. Small oxyasters of anastomosing canals below 100 /xm outer cortex, x748. i. Large oxyspherasters of cortex, wide. Megascleres mostly in confusion, occasion- X748. j. Choanosomal tylaslers, x748. ally arranged in loose sheafs. Triaenes with cladomes just below the cortex less numerous than at outer zone with croweded acanthomicrorhabds, surface. Cortex 2.5-3 mm thick, exceptionally rich numerous cladomes of triaenes and ends of oxea in magascleres arranged in narrow, mostly over- flush or slightly protruding; (b) inner zone with lapping, complex umbels, which begin c. 2 mm small oxyasters and scattered acanthomicrorhabds. below cortex and often branch once or twice. Inner (2) layer, very collagenous, coarsely fibrous Branches of umbels, 2-3 some with magascleres, (convolute fibres), 2 mm thick, off-white, with scat- interesect at angles 20-30°, some up to 60°. tered large multirayed oxyspherasters and acan- Cortex consists of: (1) Outer layer, 550-600 /xm thomicrorhabds. Below cortex and between bases thick, non-fibrous, in turn comprising: (a) Thin of the umbels are regular subcortical crypts, com- 1

SPONGES FROM BASS STRAIT 17

municating with ostia through few crooked chones, visible in section only where spicular umbels do not overlap. Chones 250-350 /*m wide, with thick sphincter below outer cortex. Chones lined by numerous multirayed oxyspherasters and scattered small oxyasters and acanthomicrohabds. Choanosome with conspicuous and crowded strongyl-and tylasters. Frequent sand grains strewn in choanosome and cortex. Spiculation: (1) oxea, mostly straight, fusiform (occasionally curved in choanosome),

927-/622-2037 x 1 1.3-25.9-44.2 ^m; (2) mostly plagiotriaenes, many mixed as orthoplagiotriaenes (clads at variable angles), asymmetrical also in partial reduction of clads or suppression (diaenes, monaenes, styles), particularly in choanosome, where rhabds are often curved: rhabds 841-/065-1333 x 15.6-20.5-28.6 pan, clads 17.3-40-62.4 ;im; (3) thin strongyles of peripheral choanosome (rare) 194-409 x 2.1-3.5 /im; (4) acanthomicrorhabds, 8. 6-//. 2-14.1 x 2-2.9-4 /mi; (5) small oxyasters of outer cortex, without or with a small centrum, 9-20 rays diameter 12.2- 14.3-11.4 fim; (6) large, densely rayed oxyspherasters of cortex, chiefly lining the chones; diameter I6.8-J/.5-39.4 /tm, centrum 38-57% of diameter; (7) strongylasters and tylasters (few oxyasters) of choanosome, with rays often distally roughened to spined, or roughened to crooked throughout, number of rays 6-12, diameter 11.8-2/.9-31.2 nm.

Remarks. The present species is most closely related to A ncorina acervus (Bowerbank). A. repens is distinguished by a thicker cortex, stunted triaenes (clad lengths generally equal to double rhabd diameter) absence of corti- with shorter and thinner rhabds, Figure 6. Ancorina robusta. spicules, a. Oxea, x68. b.

cal microxea, presence of subcortical thin stron- Range of orthotriaenes, x 68. c. Cladome of orthotriaene, gyles and presence of densely rayed oxyspherasters. x263. d. Range of anatriaenes, x68. e. Cladome of

The large, distally roughened and spined oxyasters analriaene, x263. f. Small triaenes of deeper choano- and tylasters of the choanosome may be homolo- some, x68. g. Flexuous strongyles, x68. h. Acan- thomicrorhabds, x 790. i. Cortical oxea, x 68. j. Tylaster, gous to the "anthasters" sensu Bergquist (1965: X790. 192f., description of A ncorina acervus).

Ancorina robusta (Carter) Psammastra gigas Lendenfeld, 1888: 39 (Port Jackson, NSW).-Whitclegge, 1889: 180 (off Green Point, Port

Plate 1 figure 4, plate 19 figure 3, Jackson).

Material examined. Station KG 5, one specimen (F51931); text-figure 6 station KG 6, one specimen (F51932); slation BSS 187,

Stelletta bacilli/era var. robusta Carter, 1883b: 351, pi. one specimen (F51933). Type specimens and hypotypes Stelletta bacilli/era var. robusta 14 fig. 3 (Port Elliot and Adelaide, SA). -Carter, 1886b: of (BMNH). Hololypc 123 (Port Phillip Heads, Vie., 34 m). (schizotypes) of Psammastra gigas (AM). Ecionema bacilliferum var. robustum.— Sollas, 1888: Diagnosis. Irregularly massive to bulbous, large, 197 (resume). Ancorina (Ecionemia) robusta. -Lendenfeld, 1903: 63 tending to expand from waisted base. Tough, (resume). slightly compressible. Surface red-brown to dark Psammastra bacilli/era var. robusta. - Dendy, 1905: 8 brown, smooth to faintly rugose. Oscules conspic-

(discussion). uous, often clustered. Cortex ill defined, of medium 18 F. W1EDENMAYER

thickness for genus, with megascleres in regular mediate length in inner cortex and peripheral umbels. Choanosome dense, cream to greenish choanosome, with ends slightly roughened. yellow. Large orthotriaenes with regularly curved Tylasters rare in choanosome. clads numerous, cladomes staggered around Spiculation: (1) Oxea, 903-/2/0-1412 x boundary choanosome/cortex and flush at surface. 6.2-/4.9-21.2 fim; (2) orthotrianes with gently Acanthomicrorhabds at surface exceptionally large curved clads: rhabds, 354-/295-1763 x for genus. 4.7-/7.5-26.9 /mi, length of clads 15.8-56.6-123 Description. /im; (3) anatriaenes (rare, often broken, more fre- Specimen F 51931 (pi. 1 fig. 4) bul- quent in F51933): bous, slightly compressed, expanding from narrow rhabds, 922-//27-1279 x 8.7-/5.6-19.4 urn, clads (chord perpendicular base, 7.5 X 7.5 x 5.5 cm. Apical, tilted depres- to rhabd) 26-55-44.7 ^m; few sion contains 8 oscules 2-4 mm wide. Coralline and (4) small ortho-and plagiotriaenes of the other algae encrusting on side. deeper choanosome, probably developmental: rhabds, 300-489 Specimen F 51932 swollen, expanding from base x 2.9-8 ^m, clads 7-16.5 of coalescent short pedicels. Main mass asymmetri- tim; (5) flexuous strongyles, 432-555-737 x 1 .5-2.5-3.5 cally bulbous, slightly compressed, top sloping jtm; (6) acanthomicrorhabds (width with spines), 14.7-59.9-52 lengthwise. On highest side two groups of oscules x 3.2-7.5-9.8 /xm; (7) small cortical oxea, 124.7-/65.2-188.5 (4 and 2), 2 to 9 mm wide. Greatest height 9.5 cm, x 1-2-2.8 width 9.5 x 11.5 cm. /an; (8) tylasters without centrum, knotty rays, diameter 8. 7-//. 6-14. Specimen F51933 compressed-bulbous with 6 ^m. narrow base, but more symmetrical, with crescent - Remarks. A ncorina robusta differs from most con- shaped, flattened top bearing scattered oscules generic Indo-Pacific and Australian species (A. 0.5-1.5 mm in diameter. Upper sides bear vague acervus (Bowerbank), A. murrayi (Sollas), A. aus- radial folds. Base 4x5 cm, height 10 cm, width Iraliensis (Carter), A. geodides (Carter), A. alata 9.5 x 7 cm. Dendy, A. progressa Lendenfcld, A. stalagmoides Colour in life red-brown to dark purplish brown; Dendy), and from the two new species by having dull cream to greenish yellow around base and neck much larger microrhbds. Only in A. corticata as for choanosome. (10 R6-7/4-8 and 10 R5/6 for (Carter), from South Australia, do microrhabds top of F51932.) In alcohol beige. Tough, slightly reach similar dimensions (40 x 4.2 compressible. jtm) according to the description (reported in Sollas, Surface 1888: 200; smooth except for vague rugose pattern Lendenfcld, 1903: 63). It differs from A. robusta with faint meandering depressions and pits. Rugae in its thinner cortex (500 /an), smaller orthotriaenes, and depressions 2-3 mm wide. Ostia scattered regu- lack of anatriaenes, of thin stronglyles of intermedi- larly except for some patches of irregular outline, ate size, and of small cortical oxea. without relation to rugose surface-pattern. Choanosome dense, except periphery with regu- Ancorina suina sp. now lar pararadial arrangement of incurrent canals (with Plate 1 figure 5, plate 19 intervening anastomoses) and of megascleres. figures 4-6, Deeper in choanosome conspicuous excurrent text-figure 7 canals with irregularly oblate cross-sections and lloloiype: NMV F51934. transverse fine ridges Station KG 6 (Tasmania, and folds in walls. Cortex ill Winter Cove, E side of Deal Island, Kent Group; defined, delimited below by staggered cladomes of granite boulders with algae, depth 3-6 m). loosely strewn, pararadially arranged trianenes of peripheral Diagnosis. Irregularly choanosome. From this zone, regular lobose, angular, with folds brushes of triaenes and lurrows, hard. Surface fan out to surface with little dull orange-brown to overlap, cladomes flush. cream. Cortex tough, 1-3 Width and individuality mm thick, smooth to of la.ntly rugose, internally brushes variable. Cortex 1.5 mm thick, with off-white, with irregularlv outer, scattered to bundled pigmented portion (1 mm thick) and mean- megascleres. Choanosome dering ectochones. dense, light tan, with Superficial zone of crowded cla- megascleres more abundant than in cortex, domes and acanthomicrorhabds. Lower part of in confusion. cortex fibrous, lighter. Intracortical crypts or Description. Dimensions 5.5 X 5 x 4 cm Irregu- anastomosing canals between bases of spicular larly lobose, angular, with folds and furrows in brushes, surrounded by outer pigmented layer. different directions. Several Intracortical small areas of attach- crypts and incurrent canals lined by ment on opposite sides. Colour in scattered tylasters and acanthomicrorhabds. life dull oranue brown, grading to pale fawn and cream. In Scattered small, thin oxea in cortex and outer- alco- hol, surlace beige, cortex off-white in section most choanosome. Flexuous strongyles of inter- choanosome light tan. SPONGES FROM BASS STRAIT 19

a b c crowded to packed acanthomicrorhabds. Collagen of cortex finely fibruous, with meandering and vaguely reticulate strands of fibres arranged in rounded packs; more stratified below. Few chones in deeper cortex, mostly oblique to meandering, narrow, with numerous strongylasters and acanthomicrorhabds in linings. Fine chones riddle portions of the outer cortex, 50 pta thick, where microrhabds are scanty at the surface and asters more abundant. Microscleres rare elsewhere in cortex below ectochrote. Below cortex zone of discontinuous but extensive crypts, lined by crowded asters. Tylasters abundant interstitially in choanosome. Some sand grains in cortex. In cortex and choanosome scattered balls with radial fibrous structure and clustered opaque granular matter in variable proportions, supposedly symbionts, with diameters 33-80 /xm; often crowded at surface, with outermost protruding portions frequently blackened or truncate. Spiculation: (1) Oxea, with tips often cylindrical or swollen, 1105-/257-1346 x 12.6-/5..?-! 8.8 ixm; (2) thin oxea of the choanosome, 541-703 x 2.8-3.8 /mi; (3) ortho and plagiotriaenes (often mal- formed, like dads at irregular angles, orthomesotri- aenes, stunted dichotriaene, style): rhabd 699-795-885 x 11.5-/5.5-21.8 fim, clads 25-40.4-53.8 (im long; (4) small and intermediate triaenes of choanosome: rhabd 190-499-693 x 2.9-7.0-1 1.2 /xm, clads 5.3-/J. 7-23.5 /on long; (5) tuberculate microrhabds, 5.3-6.7-8.3 x 1.9-2.9-3. 9 /tm; (6) tylasters and strongylasters with Figure 7. Ancorina stiina sp. nov., spicules, a. Oxea, x 68. 6-14 slightly rough to distally spined rays, diameter b. Thin choanosomal oxea, x68. c. Range of iriaenes, 8.9-/4.0-21.2 /*m. x68. d-h. Enlarged cladomes, x263. i. Small

choanosomal triaenes, x68. j. Microrhabds, x790. k-1. Remarks. Ancorina geodides has similar spicula- Tylasters and slrongylaslers, x 790. tion, with acanthomicrorhabds corresponding in size, but triaenes and oxea are almost double in size Consistency hard. Surface smooth in some areas, in A. geodides, and cortical and skeletal structure in others with finely dendritic depressions and are distinctive. Ancorina corlicata (Carter) agrees rugae, accentuated by colour. Oscules minute, in spicule measurements with A. suina, except for regularly scattered in furrow on one side. acanthomicrorhabds, which are larger (40 /an long) Cortex very collagenous, of variable thickness: and densely spiny in the former. A. corlicata, fur-

1 to 3 mm. Megascleres relatively scanty in thicker thermore, is round, with relatively large oscules, portions, either scattered at various angles, often and peculiar meandrine (brain-like) rugae, with crossed, some almost tangential, or arranged in ostia in the interstitial sulci. bundles gradually expanding towards surface. Many bundles in angular course in lower cortex, Stelletta Schmidt with many spicules repeatedly broken, often reaching surface obliquely. Many megascleres flexuous. Stelletta Schm\d[, 1862: 46. -Lendenfeld, 1903: 33 in part, (Myriasiru, Outermost triaenes have cladomes flush with sur- Pilochrota and Anlhastra synonymised).-Dendy, 1905: 77 (definition). face or slightly protruding, as for points of oxea. -Bergquisl, 1965: 195.- Bergquisl, 1968: 33, 44 (discussion). Megascleres in choanosome much more numerous, Myriastra Sollas, 1886: 187. — Sollas, 1888: exxxviif., chiefly in confusion, some in bundles, including dis- 112,-Dendy, 1905: 71 (definition). -Dendy 1916a: 235 tinctive infrequent small oxea and small inter- but (discussion). -Bergquist, 1965: 195. -Bergquisl 1968:44 cortex, mediate triaenes. Outermost zone of the (discussion, synonymiscd with Stelletta), - Wicdenmayer. 10-30 nm thick, occasionally to 250 /*m, contains 1977a: 177 (synonymy, references). 20 F. WIEDENMAYER

Pilochrota Sollas, 1886: 189.-Sollas, 1888: cxxxviif., and tubercles on top and sides, separated by fur- 120.-Dendy, 1905: 74 (denfinition).-Dendy, 1916a: 235 rows and lacunae. Lobes and furrows 0.5-1 cm (synonymised with Myriastra). wide, lacunae up to 2.5 cm. Dimensions 8x10 A nt haslra Sollas, 1886: 191. -Sollas, 1888: cxxxviii, cm width, 4 cm height in holotype, 6 x 7 cm by 138. 3 cm in paratype. Thin plates of coralline algae on Diagnosis. Ancorinidae with one to three types of paratype. euasters, without microrhabds or sanidasters. Surface conspicuously sandy, red-brown (2.5 R5/8) in life, now greyish beige, choanosome light Remarks. Levi (1973: 595) retained Myriastra as beige. Firm, little compressible, tearing easily. a distinct genus, with only one category of asters, Except for few minute oscules in depressions, no versus 2-3 categories in Stelletta. Bergquist's argu- orifices. ments for merging the two genera are here Cortex 1-2 thick, packed with sand. accepted. They are corroborated by the following mm Small enclaves of sand in choanosome. Cortex with clear, species, wuh one sort of aster, being closely related finely fibrous structure without orientation, to two species with two sorts of asters. except for some fibres enveloping sand grains. Some oxea Stelletta arenitecta sp. nov. occur in sandy cortex, either scattered or in oblique bundles or brushes. Plate 1 figure 6, text-figure 8 Choanosome contains oxea in confusion, of vari- Holotype: NMV F51941, Station KG 5 (Tasmania, able abundance: scattered, occasionally in irregu- Garden Cove, N side of Deal Island, Kent Group; boul- lar bundles in paratype, frequently felted in ders with algae). holotype. Triaenes with atrophied cladomes scarce Paratype: NMV F51942, Station KG 6 (Tasmania, (below 2% of megascleres) and scattered. Small Winter Cove, E side of Deal Island, Kent Group; granite asters scarce, boulders with algae, depth 3-6 m). numerous in peripheral choanosome, also in cortex. Subcortical crypts irregularly dis- Diagnosis. Sprawling, irregularly tuberculate to tributed, narrow, slit-like, some well below cortex. lobulate; firm, but easily torn. Surface red-brown They communicate with oblique, twisted and in life. Cortex 1-2 mm thick, full of sand, with rare branching crevice-like incurrent canals. Excurrent spicules. Choanosome light beige, with megascleres canals round, up to 1.5 mm wide. Choanosome in confusion, triaenes rare and atrophied. Euasters almost packed with minute symbionts, in the shape scarce. of mulberries, with diameter as for asters. Some Description. Sprawling, capriciously shaped masses occur in cortex. Spiculation: (1) Oxea and stron- with lacunose outline at base; twisted ridges, lobes gyloxea transitional to strongyles and styles, mostly straight, hastate to fusiform, 286-645-873.5 X a b c 2.6-9.5-14.4 /mi. Thinnest oxea in paratype (absent in holotype) frequently flexuous, broken, with cor- (1 roded ends, possibly separate category (300-453 x 2.6-2.9 urn). (2) Plagio-to orthotriaenes with much reduced, regular or irregular cladomes, rhabd 503-625-704 x 5.3-9-10.3 Mm, clads 3.8-9.5-17.1 long. ^m (3) Strongylasters to tylasters (intergrad- mg), frail, without centrum, 6-9 very slender, smooth to slightly knotty, straight rays,' 6.5-/7.J-13.8 nm in diameter. Remarks. Stelletta arenitecta falls within a small group of species containing much foreign detritus and relatively rare, reduced triaenes of irregular distribution. Stelletta arenaria Bergquist (1968: 45) is comparable to S. arenitecta in its cortex, and in the rarity of its triaenes. It differs in shape, colour arrangement and size of megascleres, less reduced triaenes, and abundance of asters which fall into Figure 8. Stelletta arenitecta sp. nov., spicules, a-c. Oxea, two categories (tylasters and "chiasters" i e strongyles, and styles, x68. d. Triaenes, x68. e-g. anthasters sensu Wiedenmayer, 1977a, nee Berg- Enlarged cladomes, x263. h-i. Tylastcrs and stron- quist, 1965). Stelletta vestigium gylasters, x790. Dendy (1905- 78) resembles S. arenitecta in the shape, size 'and SPONGES FROM BASS STRAIT 21

arrangement of its megascleres, but differs in external shape, colour, lack of cortex, abundance of foreign matter in the choanosome, and in possessing two categories of asters (oxyspherasters and oxyasters). There seems to be a fourth, undrescribed sandy species of Stelletta, represented by a specimen from Port Phillip Bay (BMNH, Dendy Coll. RN 614, double-registered 1925.11.1.611 and — 438,

with 3 slides in cabinet 1 , trays 43, 44, which I have not examined), labelled with a manuscript name by Burton. It is massive, with almost smooth surface, no macroscopically visible cortex, and sand distributed throughout the sponge.

Rhabdastrella Thiele

[Aurora] Sollas, 1888: cxxxix, 187 (nee Aurora Rago- not, 1887). -Dendy, 1916a: 242 (definition, discussion, review). — Levi, 1967a: 240 (discussion, brief review), - Bergquist, 1968: 54 synonymised wilh Rhabdastrella). Rhabdastrella Thiele, 1903a: 934. -Bergquist, 1968:33, Figure 9. Rhabdastrella cordata sp. nov., spicules, a. 54 (discussion). Oxea, x68. b. Range of triaenes, x68. c-e. Enlarged Diastra Row, 191 1: 300. -Dendy, 1916a: 245 (discus- cladomes, x263. f. Ghoanosomal strongyles, x68. g. sion, synonymised with Aurora). — Bergquist, 1968: 54 Spherasters of inner cortex and choanosome, x263. h. (discussion, synonymised with Rhabdastrella). Anthospherasters of outer cortex, x263. i. Superficial Aurorella de Laubenfels, 1957: 245 (replacement name oxyspherasters, x790. j. Choanosomal strongvlasters, for A urora Sollas, nee Ragonot). -Hechtel, 1983: 80 (as x790. subgenus of Rhabdastrella).

Diagnosis. Ancortnidae with reduced triaenes or YR9/4, 5 R5/10, 2.5 R5/4, 5 R6/2, and neutral without triaenes; euasters comprise massive types black-grey. Firmly rubbery. Surface is partly (spherasters, anthospherasters, sterrospherasters), smooth, partly faintly rugo-vermiculate. Narrow commonly in cortical layer. zone on one side near point now finely wrinkled. Rhabdastrella cordata sp. nov. Broad side with deep oblong invagination. Numer- ous oscules, 0.2-1 mm wide in life, on top, in 5 Plate 1 figure 7, plate 20 figure 1, to 6 separate or confluent fields. Folds and invagi- text-figure 9 nations surround base. Cortex mostly 500-750 thick, Holotype: NMV F51935 Station KG 6 (Tasmania, Winter ^m finely fibrous Cove, E side of Deal Island, Kent Group; granite boul- and weakly pigmented almost throughout. Very ders with algae, depth 3-6 m). thin hyaline ectochrote. Choanosome dense and collagenous; boundary with cortex indistinct. Sub- Diagnosis. Massive-lobate, low, firmly rubbery; cortical crypts or canals infrequent and irregular. base slightly constricted, folded. Surface mottled, Megascleres in choanosome in widely spaced bun- dark grey-brown, purple, pale brown, and dull dles and loosely scattered interstitially. Orientation yellow. Surface smooth to faintly rugose. Oscules of the latter mostly random in deeper parts, more small, numerous, tending to cluster. Cortex parallel with bundles in periphery. Bundles below 500-750 /mi thick, light, collagenous, lower bound- cortex usually oblique, some almost tangential, ary indistinct, with megascleres in vague umbels, branching irregularly, expanding into brushes well and spherasters of 2 sizes in thin crust at surface. below cortex. Contiguous brushes separated or Choanosome dense, with megascleres scattered or overlapping irregularly. Their apices uneven, with loosely bundled. Triaenes often irregular. cladomes of triaenes commonly placed in inner Description. Like a heart lying on its side. Height third-half of cortex, oxea frequently reaching 3.5 cm, width 5 x 5.5 cm. Base of attachment higher. Large spherasters (mostly oxyspherasters slightly contracted. Colour in life mottled: dark grey- with blunt rays) scattered in choanosome and brown, mixed with purple, pale brown, dull yellow. cortex. Small asters of various shapes rare in Sides finely speckled purple. Base, like choano- choanosome. In outer quarter-two-fifths of cortex, some, light beige to cream, with sharp boundary spherasters are crowded to packed in clusters, to darker top and sides: 5 RP3/2, 10 R8/4, 7.5 mostly modified as anthospherasters. Conspicuous 22 F. WIEDENMAYER

zone free of spicules (except for occasional oxea and scattered spherasters) between ends of megasclere-brushes and the crowded spherasters of outer cortex. In hyaline ectochrote and between outermost anthospherasters infrequent small spher-

asters of a distinct category. Spiculation: (1) Oxea, straight to curved or bent, fusiform to hastate, mostly with blunt, conical, or mucronate tips, 387-526-657 x 3.2-9.3-15 pm; (2) triaenes, often irregular, with variable directions of single clads (mixed ortho-, plagio-and pro-types), one, two, or all clads stunted or suppressed (diaenes, monaenes, tylostyles), occasionally as mesotriaenes, but with orthotriaenes dominating in periphery and cortex; rhabd 207-2^-342 x 5.4-9-13.2 pm, length of clads, 9.7-57.4-90.3 pm; (3) thin strongyles of choanosome, 150-250-294 x 2 pm; (4) large spherasters, with blunt, conical to terete rays (mostly in choanosome and inner portion of cortex) and anthospherasters with truncate rays (near surface), diameter 18.6-22. /-25.5 pin, centrum 54-64-73%,' (5) small oxy-and strongylospherasters at the surface, diameter 7.0-70.5-13.4 pm, centrum 34-46-66%; (6) small asters of choanosome, of variable types (oxy-, strongyl-and tylasters), mostly without or with small centrum (rarely around 35%), and 6-18 rays, diameter 7.8-77.9-19.4 Figure pm. 10. Rhabdastrella intermedia sp. nov., spicules, a. Range of oxea, x68. b-c. Remarks. See next species. Enlarged ends of oxea, X263. d. Range of triaenes, x68. e-g. Enlarged cladomes with range of widths, X263. h. Rhabdastrella intermedia sp. nov. Choanosomal thin oxea, x68. i-j. Cortical spherasters, x790, k-1. Choanosomai Plate oxyasters, 790. 1 figure 8, plate 20 figure 2, x

text- figure 10 black, tinged purple to red (5-10 RP3/T to 2.5 R5/2 Holotype: NMV F51936, station KG 6 in life, (Tasmania, Winter N3/R2 in alcohol); bulging rim slightly Cove, E side of Deal Island, Kent Group; granite boul- lighter. Eirmly rubbery in life. Choanosome light ders with algae, depth 3-6 m). cream Y9/2). (2.5 Depression, with fine folds along Diagnosis. axis and round Irregularly cake-shaped, with truncate, invagination near the lower end hollow top, firmly riddled by minutes rubbery. Surface purplish black," oscules, 100-200 /

choanosome. Matrix generally dense, lightly col- 3, and R. fibrosa Hechtel (1983: 81) to group 1 of lagenous, almost packed with clustering unicellu- Levi. Neither of the new species falls comfortably lar symbionts (diameters around 12 /itn). In into one of these groups, though R. cordata might peripheral choanosome, with filamentous algae, be included in the first on account of the dominance just below zone of subcortical crypts, principal of orthotriaenes. Of the species included by Levi spicule-bundles expand into wide, complex umbels, in this first group, R. primitiva (Burton, 1926c) commonly erect and much overlapping. Other shows some similarities in spiculation with R. cor- umbels narrow, not overlapping, oblique. All data (distinct category of small cortical spherasters; umbels end at surface. tylasters and microxea of the choanosome), but the Cortex 460 to 650 /mi, in two layers: (1) Outer megascleres are distinctly larger, with only regu- cortex, heavily pigmented, 1 10-250 /mi, with spher- lar othotriaenes. R. globostellata (Carter), with syn- asters packed between ectochones: (2) Inner cortex, onyms Aurora providentiae Dendy (1916a) and less pigmented, fibrinous, with scattered spherasters Stelletta disco/or Bosraug (1913), has cortical spher- (rare in the basal part), 350-400 /un. Immediately asters and choanosomal oxy-to strongylasters (no below is a zone, 250 /mi, of regular subcortical centrum, slender rays) of much larger dimensions crypts, between bases of umbels, with matrix lighter than in either of the new species. Rhabdaslrella than in subjacent choanosome, containing filamen- reticulata (Carter, 1883b; see Sollas, 1888: 188), has tous algae. Ectochones narrow, meandering to cortical spherasters and choanosomal stringlylasters oblique, wider endochones lined by small asters, comparable to those in R. cordata, but lacks special leading into subcortical crypts. Spiculation: (1) small spherasters, thin choanosomal strongyles, Fusiform and hastate oxea and strongyloxea, fre- and has larger magascleres. R. row/ (Dendy, 1916a) quently slightly bent, 494-603-618 X 5.6-/2.4-15 has larger megascleres, and distinctive cortical ster- urn. (2) Triaenes, of different types, rare, stunted rospherasters. Rhabdaslrella spinosa (Levi, 1967a) in various ways (ortho-, ana-, plagiotriaenes to has again larger megascleres, and cortical spher- -monaenes); types with one or two dads much asters up to 45 /mi in diameter, with a centrum not reduced or suppressed most common, some with exceeding 50%. Rhabdaslrella aurora (Hentschel, all clads suppressed (subtylostyles); no regular 1909; see Bergquist, 1968: 54) is comparable to R. types; some have subterminal aborted clads in addi- intermedia in the frequent irregularity of cladomes, tion; rhabds 520-638 x 10.6-14.1 /un, clads and to R. cordata in the presence of special small 5.3-70.2 urn in length. (3) Thin oxea and strongyles spherasters at the surface. But the megascleres are of choanosome, some (especially the thinnest) very again larger than in both new species, and the large flexuous, 295-404-549 x 2-5.5-5.7 /mi. (4) Cor- spherasters are comparatively scarce, being largely tical oxy-and anthospherasters, the latter often replaced by foreign detritus in the cortex. The spirit mixed; with rays conical to blunt, smooth to dis- collection in the British Museum (Natural History) tally acanthose, and truncate in same spheraster, contains 3 specimens (Dendy's RN 739, 745, 1200, diameter 15.4-20.9-24.3 /mi; centrum 42-49-57%; with 16 slides, BMNH cabinet 1, tray 24) of yet species, (5) choanosomal oxyasters, diameter 7.3-/7.0-28 .7 another undescribed with a manuscript /*m, centrum 19-29-38%. name by Burton. 1 have not examined the slides, but one of the specimens differs from both new Despite similarities in spicule types Remarks. species by having a thicker cortex (1.3 mm) and a there are enough present and their measurements, dark brown choanosome. differences to regard Rhabclastrella intermedia and R. cordata as distinct species and not as extreme forms of a polymorphic species. The chief differ- Geodiidae Gray, 1867 ences are in cortical and skeletal structure, in morphology, relative abundance and distribution of Geodia Lamarck choanosomal and cortical asters, in the absence of Geociia Lamarck, 1815 (1814-1815): 333. -Hechtel, regular orthotriaenes in R. intermedia, and in tri- 1965: 68, 70 (subdivision in subgenera). — Levi in Brien aenes having much shorter rhabds in R. cordata. et al., 1973: 595 (subdivision). In his brief review of 14 species then known in subgenus, see Wieden- the genus, Levi (1967a: 240) distinguished three Subgenera. Nominotypical 1977a: 178 (synonymy, references). groups: (1) with orthotriaenes, (2) with plagiotri- mayer, Fleming, 1828: 516. -Sollas, 1888: aenes, and (3) without triaenes. Rhabdaslrella Cydonium 254.-Hechtel, 1965: 70. tenuis (Lindgren) should be added to group 2. Of CXLVIII, 218, 1880: 396.-Lendenfeld, 1903: 93 the two Brazilian species described more recently, Isops Sollas, key, review). R. virgula Boury-Esnault (1973a: 270) falls to group (diagnosis, 24 F. WIEDENMAYER

Sidonops Sollas, 1889: 277.- Lendenfeld, 1903: Geodia (Geodia) punctata Hentschel 99 (diagnosis, key, review). - Wiedenmayer, 1977a: Plate 2 figure 1, plate 20 figures 179 (type species redescribed). 3, 4, text-figures 11-13 Diagnosis: (Genus and subgenera) Geodiidae with Geodia punctata Hentschel, 1909: 366, text-fig. 10' pi sterrasters only (no small euasters, spheres, 23, fig. 25 (Shark Bay, WA). microrhabds) in the cortex. Subgenera distin- Material examined. guished by distribution and relative size of incur- Station KG 2, one specimen (F51937). rent and excurrent openings. In the nominotypical Diagnosis. Stump-shaped, truncate top with con- subgenus, chones have cribriporal roofs and oscules fluent depressions containing oscules; firm. Sur- are clustered. In Cydonium, incurrent and excur- face smooth, slippery in life, with minutely rent orifices are cribriporal and similar in size. Isops reticulate pattern, greyish brown mottled with dull has solitary ostia and distinct solitary oscules. In purple and yellowish cream. Cortex 300-400 Mm Sidonops, chones are cribriporal and oscules single, thick, often lacunose with conspicuous chones. in depressions or deep invaginations. Sterrasters packed except at base of cortex.

Figures 11, 12. Geodia (Geodia) punctata. Details in reflected light. Figure 11. Perpendicular section, x9.8 showing, from top, oblique view of surface, mottled cortex, peripheral choanosome with brushes of megascleres, and irregular canals of deeper choanosome Figure 12. Reticulate pattern of surface, x 11.7. SPONGES FROM BASS STRAIT 25

a b c greyish brown with purple tinges on top (now greenish grey, 5 Y5-6/2), dull purple (7.5 RP4-6/8) mottled with yellowish cream (5 Y9/4), the latter also in choanosome. Surface in life smooth, slippery. Firm but somewhat compressible. Surface in purple areas of sides with fine reticulate pattern, like giraffe-skin when magnified, slightly in relief, with fine ostia scattered and clustered (text-fig. 12). Choanosome dense, collagenous; excurrent canals commonly compressed. Megascleres mostly loosely scattered in deeper

choanosome; some arranged in bundles with little orientation, or subparallel and oblique to the surface. Small asters of various sizes numerous throughout choanosome, but irregularly distributed, most abundant around canals. Sterrasters scarce in choanosome. Peripheral choanosome 800 fim thick, with brushes of megascleres, some perpendicular and isolated, most oblique, overlapping and crossing each other. Subcortical crypts and canals irregularly and widely spaced, often continuing radially or obliquely below, confluent or connected by fine, irregular anastomoses. Cortex mostly 300-400 /uu thick. Below very thin hyaline eetochrote, with scattered small spherasters, a weakly and finely fibrous main layer. Fibres more distinct and often stratified in inner portion. Ster- rasters crowded to packed between chones, except for a thin basal layer of scattered sterrasters. Inner boundary of cortex indistinct in collagenous structure, but usually marked by tangential magascleres. Cladomes of scarce triaenes commonly flush with this base, but tips of outermost oxea often reach somewhat higher. Chones 80-100 /un wide in outer Figure 13. Geodia (Geodia) punctata, spicules, a. Range half, frequently oblique, widening and confluent ofoxea, X 68. b. Enlarged ends of strongyloxeon, X263. below. c. Range of triaenes, x68. d-g. Enlarged cladomes, Spiculation: (1) Oxea, strongyloxea, strongyles, X263. h-j. Sterrasters, x790. h. Earliest developmen- rarely anisoxea, straight or weakly bent, frequently tal stage, i. Intermediate stage, j. Fully grown stage, k-1. with telescoped or mucronate ends, 504-667-806 C'hoanosmal asters, x 790. k. Oxyaster. I. Strongylaster. x 3.8-9.4-14.1 /un; (2) ortho-to plagiotriaenes, m. Small Strongylosphcrasters of eetochrote, x790. often with irregular cladomes and atrophied dads,

Choanosome yellowish cream, its periphery with rhabds frequently telescoped, strongylote or megascleres in distinct, often oblique umbels, in swollen, rhabds 334-423. 7-567. 6 x 7.6-/0./-12.4 deeper parts mostly confused, with numerous small /im, clads 22.6-69.7-110.9 /on; (3) graceful ster- asters. Ortho-to plagiotriaenes often with irregu- rasters with relatively long rays, spacing of trun- lar cladomes. cate tips mostly equal to half their width, occasionally equal to their full width, diameter Description. Like a tree-stump, 7.5 cm high, 6 x 22.6-29J-33.8 /un; (4) choanosomal oxy-to stron- 7 flat, truncate top, slightly con- cm wide, with gylasters with no or small centrum, 6-12 rays, often stricted 3.5 x 4.5 cm wide. Base of attach- base roughened or finely spined in larger ones, diameter oblique, extending in patches to two-thirds ment 8.4-/7.7-29.7 ^m; (5) small strongylo-and partly smooth, partly thrown into of height. Sides tylospherasters of eetochrote, rare in choanosome, broad lobes. Top bears shallow furrows and low, diameter 6.8-9.5-12.5 /mi, centrum 38-50%. 8-9 partly confluent depressions of rounded- polygonal outline now shrivelled, containing scat- Remarks. In the original description, Hentschel infrequent, flexuous, tered oscules 0.3-0.5 mm wide. Colour in life mentions thin oxea, rate 26 P. wildlnmaylr

styles (derived from oxca), and very rare anatri- 1954a: 2^5, text-fig. 155 (Caroline Islands). - Bcrgum'sl, aenes, none of which were seen in my preparations. 1969: 69, lexl-lig. 2 (Heron Island). Ridley, 1884a: pi. tigs The measurements given for oxea, triaenes, ster- Steltettinopsis carteri 476, 43, n, n' (Torres Strait). rasters and elioanosoinal oxyaslers are all some- Coppatias carteri. -Sollas, 1888: 208 (resume, what above mine bnl whether this is of discussion). biogeographic or ecological significance would be Stellettinopsis tubercuiata Carter, 1886b: 126 (Port premature to decide. Phillip Heads). ( >f the oilier three species with Australian records Coppatias tuberculatus,— Sollas, 1888: 207 here accepted in the nominate subgenus, (1. ((1.) (resume). Deucly, 1905: 109 (in discussion). carteri Sottas has generally larger spicules, includ- Jaspis tubercuiata. -de Laubenfels, 1954a: 224, text- dieholriaenes. (i. fig, ing (G. ) globostellifera Carter has 154, (Marshall Islands). larger slerraslers and niueh largei megaseleres, Stellettinopsis lutes Carter, I886h: 459 (Westernpon Hay). including protriaenes. G. (G.) nigra Lendenfeld is

Coppatias /ulcus . Sollas, 1888: 207 (resume). black, has large lyloslyles, eortieal styles, and plagiolriaenes with longer and thicker rbabds than Material examined. StaU'on KG I. one specimen (1-5 1938). in G. punctata. Type specimens of Amorphina stellifera. Stellettinopsis tubercuiata, ami Stellettinopsis lutea (BMNH). Coppatiidae Topsent, 1898 Diagnosis, Encrusting to massive, irregularly Synonyms. Eplpolasidae Sollas, 1888 (nomen nudum; lobale, commonly enveloping parts of algae, shells, used by I evi [in Brien el al., 1973}: 598; Vacelct, Vas- detritus and other foreign objects; leathery, but seur and l.vi, 1976: 23). easily lorn; dull purple Io brownish drab, or golden Jaspidae ilc I aiiix-nk'ls, 1936a. yellow throughout, but deeper al surface. Surface Remarks. I have already pointed out (Wieden- smooth to microhispid, vaguely rugose, occasion- mayer, 1977a: 172) thai the priority of Coppatii- ally (uberculate. Cortex indistinct, about 0.5 mm dae is not affected by the obsolescence of the type thick, with matted oxea, paralangential or in con- genus. Though Jaspidae was introduced before fusion. Oscules indistinct. Choanosome with abun- 1961, and was increasingly accepted following dant, mostly confused, bent oxea of extreme Size Hcrgquisl (1968: 30) it cannot be said to have won range, and numerous minute euasters. general acceptance in Ihe sense of the tC 'XN, Article Description. 40b. Levi et al. have preferred to use Lpipolasidae Small, partly enveloping 2-3 empty vermetid (see synonymy above) and other authors have shells, together with two smaller sponges (Celllodes avoided family names within Ihe order or suborder incrustans and Aulorrhiza protumbens). I'pipolasida. Vermetid shells partly coaled inside, partly penetrated between layers, by Ihe sponge. Colour dull purple to Jaspis Gray brownish drab on surface (5 RP5/6-8). Leathery on surface, easily JaspisQt&y, lorn to I 867a: 526. Wilson, 1925: 329 (resume, crumbly internally. Indistinct cortex of variable in pari, not Rhabdastreila). Bergqujst, 1968: 33, 55 (dis- thickness, mostly 400-500 /t m, not eussioii). Wicdcnniayer, 1977a: 172 (synonymy, easily detachable, sometimes references). discontinuous, or layered. Outer, deeply pigmented layer. Whole cortex commonly Diagnosis. Coppatiidae with euasters only as packed with oxea in confusion. Oxea at microsclercs. surface either chiefly tangential or in a rugose pattern of mailed brushes Jaspis stellifera (Carter) (text-fig. 14). Lining similar to cortex surrounds larger cxciu rent canals in choauo- Plate 2 figure 2, plate 20 figures 5, 6, some, about 1.5 mm wide. Smaller canals, 50-1 50 ,im wide, Icxi-I'igures 14 17 riddle the choanosome. No oscules Clusters of oslia (eribriporal groups) in slight AmOrphinasteliiferaQa.net, 1879b: .144, pi. 24, fig, saucer-like depressions (text-fig. io (South Australia), 14). Single ostia' 50-100 ^em wide, lead to oblique, Stellettinopsis Stellifera, Ridley, 1884a: 477 meandering, often confluent ehoncs (discussion). of similar diameter, confluent with numerous subcortical Coppatias Stellifera, Sollas, t888: 208 (resume, crypts (text-fig 15) discussion). commonly arranged in several equidistant layers' Jaspis Stellifera. Shaw, l')27a: Arrangement of 422 (excessive syn- megaseleres in choanosome mostly onymy, description from Tasmania, discussion, confused, distribu- scattered or in disoriented bundles tion; possibly Jaspis coriacea). -- Burton, 1934a; 52: (now ( hoanosome with pigmented cells and record from Ciieal llaniei scattered to Reel Duly), de I aubeiilels. packed elhpsoidal bodies, 20 30 /tm long, with cen- SPONGES FROM BASS STRAIT 27

W^i.

\r- ^ =

'5 v ' '' ^2t tr^^iri'6dtliii- .ill* 14 *MT

Figures 14-16. Jaspis stellifera, details in reflected light, x 16.5. Figure 14. Smooth depression with cribriporal groups surrounded by brushes and felt of megascleres. Figure 15. Perpendicular section through cortex with a cribriporal group, intracortical and subcortical crypts. Figure 16. Perpendicular section showing irregular cortex, small and large canals in choanosome. 28 F. WIEDENMAYER

based on the discussion therein, from which Jaspis slellifera emerges as a rather polymorphic species. Some of Bergquist's points need to be corrected or clarified. Neither Carter (1879b) nor Sollas (1888) mentioned that the asters in /. slellifera are tylasters. Carter's primitive figure can only be interpreted as a strongylaster. Stellettinopsis carteri, as described by Ridley (1884a), does not have identical microscleres: Carter and Sollas give 17 fim diameter for J. slellifera, Ridley gives 50 ftm for 5. carteri. Diameters of asters given by Carter and Sollas are 8.3 /xm for S. tuberculata, and 18 //m for 5. lutea, hence the latter are not "much smaller" than in J. slellifera, and those of S. tuberculata are near the lower extreme in the specimen examined. The Australian species of Jaspis urgently need revision, as none of the primary types involved have been re-examined since the original descriptions. Sollas merely gave resumes with spicule dimensions converted to metric. Carter gave only single measurements (commonly the largest observed) with no or little regard for ranges. Since the smallest oxea in the specimen examined fall into the range of the "microxea" in Stellettinopsis coriacea and S. purpurea Carter, it is not unlikely that Carter overlooked intermediate sizes, that the latter two species might have to be merged in Jaspis slellifera, and that this species is perhaps more polymorphic in southern than in northern Australia.

Order Spirophorida Brien, 1968 Figure 17. Jaspis slellifera, spicules, a. Range of oxea, Tetillidae Sollas, 1886 x68. b. Oxea, enlarged, x263. c. Style (rare), X263* d-f. Asters, x790. Tetilla Schmidt

Tetilla Schmidt, 1868: 40. -Sollas, 1888: cxxiv 1 (defi- tral clear core surrounded by equal cells. nition). Such -Lendenfeld, 1903: 16 (review with resumes) - bodies were originally described by Carter, Dendy, 1905: (definition). and are 89 -Wilson, 1925: 355 (defi- probably algal symbionts. nition, resume, Scattered foreign debris, discussion, list of species). -Bergquist 1965: 197f throughout choanosome, some quite large. Asters (discussion). -Bergquist, 1968: 60 (Southern numerous. Ocean species). Chrotella Sollas, 1886: Spicululation: (1) Oxea (pi. 20 fig. 6) hastate to I80.-Sollas, 1888- cxxv 17 (definition).- Lendenfeld, 1903: fusiform, commonly symmetrically bent, 16 (synonymised with occasion- Tetilla).- Burton, 1937: 11 (synonymised ally modified to anisoxea, strongyles and styles, with Cmcckyra) -Burton, graded 1959: 200 (synonymised with size range with little if any relationship Cmacnyra). between length and thickness, 89-564. £-682. 8 x Diagnosis. Hemispherical 2.6-5.7-17.6 /tm; (2) oxyasters with about 10-20 to globular Spirophorida without vestibular roughened to irregularly spined rays, centrum calices, and without modified tnaenes. Small oxea, mostly below 24%, with relatively rare stron- if present, not spiny and not in cortical layer. gylasters of similar size, with some rays tylote, cen- Sigmaspires present. trum occasionally up to 45%, diameter 8.9-72-14.6 Tetilla praecipua ^m. sp. nov. Plate 2 figures 3, 4, plate 21 figure 1, Remarks. It is possible that the partly corroded ver- text-figure 18 metid shells functioned as excurrent canals and Molotype. NM\ F51943, oscules. My synonymy is based on that suggested Bass Strait, 17 km S of Warr namboo 38°32.0

~S~

Figure 18. Tetil/a praecipua sp. ft., spicules, a. Range of oxea, x 34. b. Enlarged ends of oxea and range of thickness, x 329. c. Range of triaenes, 34. d. x Enlarged cladome, x 329. e. Microxea, x 329. f. Sigmaspires, x 329.

Diagnosis. Small, roughly hemispherical, firm. Sur- 104.7-757.2-176.5 x 1.0-/. 6-2.0 pm; (4) sig- face hispid, orange-yellow with greenish grey on maspires, 9.5-/7.2-24.4 ^m chord, about 0.8 fxm sides. Small, clustered oscules. No distinct cortex. thick. Radial choanosome with conspicuous terete canals Remarks. Tetilla praecipua falls within the small regularly arranged, thick megascleres in bundles, group of species having small oxea scattered interstitial microxea in confusion, and faint sig- throughout the sponge. The three species of this maspires. Protriaenes rare, atrophied; no group then known, Tetilla monodi Burton anatriaenes. (1956) from West Africa, T. oxeata Burton (1959), from Description. Torn off from base, slightly damaged South Arabia, and Tetilla microxea Bergquist, top. Dome-shaped, with humped top. Diameter at from the Caroline Islands, were discussed in Berg-

base 6 cm, height 4 cm. Firm, moderately compres- quist (1965: 197). Tetilla praecipua is closest to T. sible. In life dull orange-yellow with mixture of microxea in spicule measurements, rarity of triaenes greenish-gray on sides (7.5 YR6-7/10). Surface and atrophied cladomes of protriaenes. Tetilla from faintly hispid on top, to distinctly hispid (spic- microxea differs from T. preaecipua in colour, in ule plush 0.5-1 mm) on sides, with foreign debris, possessing conules and long spines of oxea project- small algae and hydroids. Two symbiotic barna- ing from the surface, and in the presence of cles imbedded on one side. Small oscules, 0.2-0.5 anatriaenes. mm wide, clustered in irregular fields, chiefly on Order Hadromerida Topsent, 1894 one side of apical hump. Excurrent canals terete,

straight, annualate, mostly around 1 mm wide; Tethyidae Gray, 1867 radial and oblique, regularly spaced. No distinct Tethya cortex. Radial arrangement of megascleres. Thin- Lamarck ner megascleres (oxea) commonly single, slightly Tethya Lamarck, 1814 (1814-1815): 69 (in part).- spaced; thicker ones mostly in bundles 140-350 /*m Lindgren, 1898: 358f (revision). -Burton, 1956: 120.- wide, between radial canals. Throughout choano- Burton, 1959: 214 (synonymy, discussion). — Bergquist, 1968: 37 (discussion). — Wiedenmayer, 1977a: 170 some scattered to felted microxea or rhaphides. (synonymy, references). Thin sigmaspires numerous. Triaenes rare, cladomes atrophied, commonly as promonaenes. No Diagnosis. Commonly globular Tethyidae, with triaenes in inner (focal) choanosome; no anatri- distinct cortical and choanosomal types of euasters. aenes anywhere. Spiculation: (1) Oxea, straight or bent in the Tethya ingalli Bowerbank middle, regular, with blunt, mucronate or tele- Plate 2 figures 5, 6, 9, plate 21 figure 2, scoped tips, 91 0-3065-4 182 x 4.7-26. /-41. 5 jtm; (2) triaenes, much atrophied, mostly as text-figure 19 promonaenes, occasionally as tylostyles, with fusi- Provisional synonymy. form rhabds thickest in the middle, clads occasion- Tethea ingalli Bowerbank, 1859: 307, pi. 25 figs 12, ally subterminally irregularly polytylote, rhabds 14. -Bowerbank, 1872: 119, pi. 5 figs 11-17 (Fremantle, 30.7-67.6 jun; 1093-2409 x 2.8-4.1 pm, dads (3) WA)-Vosmaer, 1932-1935: 192 (resume). roughened, straight or microxea or rhaphides, Tethya ingalli. - Ridley, 1 884b: 624 (discussion). -Sol las gently curved to slightly flexuous, 1888 431, pi. 44, figs 15, 16 (Bass Strait, Port Jack- -.

30 F. W1EDENMAYER

- Tethya inflata Lendenfeld, 1 888: 49 (Port Jackson). Lindgren, 1898: 360 (discussion). -Hallmann, 1914a: 275 (syntypes redescribed, as Donatio ingalli var. laevis (Len- denfeld). — Topsent, 1918: 590 (discussion). — Vosmaer, 1932-1935: 197 (resume, discussion). Tethya phillipensis Lendenfeld, 1888: 50 (Port Phillip Bay, Vic.). — Lindgren, 1898: 360 (discussion, synonymised with T. ingalli). — Hallmann, 1914a: 277, pi.

1 5, fig. 4 (syntypes redescribed as Donatio phillipensis). — Topsent, 1918: 579, 590, 650, 601 (resume, discussion, synonymised with T. diploderma). — Burton, 1924b: 1039 (syntype in BMNH re-examined, synonymised with Dona- tio diploderma). -Vosmaer. 1932-1935: 198 (resume). Tethya laevis Lendenfeld, 1888: 51 (Port Jackson).— Lindgren, 1898: 360 (discussion, synonymised with T. ingalli). —Hentschel, 1909: 372 (remark). -Hallmann, 1914a: 281 (syntypes redescribed, as Donatio ingalli var. laevis, includes Tethya ingalli: Sollas). —Topsent, 1918: 591 (discussion, synonymised with T. diploderma). Burton, 1924b: 1039 (syntype in BMNH re-examined, synonymised with Donatio diploderma). -Vosmaer 1932-1935: 197 (resume, discussion).

Wonatia parvistella Baer, 1906: 31, pi. 2 fig. 9; pi. 5 figs 35-38 (Zanzibar).—Dendy, 1916a: 262 (synonymised with Donatio japonica). -Topsent, 1918: 598, 601 (discussion, synonymised with Donatio diploderma). Burton, 1924b: 1039 (synonymised with Donatio Figure 19. Tethya ingalli, spicules, a. Range of principal diploderma). mcgascleres., x27. b-d. Enlarged tips of principal Donatio japonica varr, mtcleata and glohosa Hentschel, megascleres, x 263. e. Cortical styles, x68. f. Oxyspher- 1909: 372, text-fig. 1 ; 373, text-fig. 1 12, pi. 22 fig. 8 (WA asters, x263. Tylasters, x 790. h-j. Choanosomal g. between Shark Bay and Albany). -Hentschel, 1912: 317 asters, x790. (types re-examined, synonymised with Donatio ingalli). - Topsent, son). Lendenfeld, 1897a: 18 (synonymy). — ?Kirkpatrick, 1918: 598 (resume, discussion). Donatio diploderma. 1900a: 132 (Christmas Island). -Topsent, 1918: 578, 579, -Topsent, 1918: 574, 600, text- figs 22-24 600 (resume, discussion, synonymised with Donatio (part, discussion, description of new records; Gulf of Aden?; diploderma (Schmidt), other synonyms). — Burton, 1924b: Ambon, Indonesia?). — Burton, 1924b: 1039 (part). -?de 1039 (holotype re-examined, synonymised with Donatio Laubenfels, 1936b: 451 (Pacific end of Panama Canal). -Guiler, 1950: diploderma). -Bergquist, 1968: 36, pis 5b; 13c, e (syn- 9 (Tasmania, identified by Burton). onymy, description, discussion; New Zealand). -?de Laubenfels, 1950a: 30, text-fig. 20 (Hawaii). -?de Laubenfels, nee Tethya ingalli. -Lindgren, 1898: 317, 360 1954a: 232, text-fig. 160 (Marshall Islands). (= Tethya seychellensis fide Hechtel, 1965: 65). -?de Laubenfels, 1955a: 141 (Gilbert Islands). -Levi, 1956b: 7, text-fig. 4 (fide 1 Donatio ingalli. -Hentschel, 1909: 371 ( = ? Tethya Bergquist, 1968; seychellensis). Madagascar). Tethya multislella.— Donatio ingalli.— Hentschel, 1912: 316 (Aru Islands, Bergquist, 1961b: 193, text-fig. Indonesia). 16a, b (nee Lendenfeld; New Zealand).

nee Donatio ingalli. - Dendy, 1916a pi. 264, 48 fig. 3 Material examined. Station KG 1 , one specimen (F51944). = Tethya ( robusta Bowerbank, fide Burton, 1924b). Type specimens of Tethea ingalli, T. cliftoni (BMNH); Donatio ingalli var. maxima Hentschel, 1909: 372 of Tethya inflata, T. phillipensis, T. laevis (AM). (Shark Bay, WA).- Topsent, 1918: 598 (discussion). - Burton, 1924b: 1039 (synonymised with Donatio diploderma). Diagnosis. Globular, small to medium-sized; Tethea cliftoni Bowerbank, com- 1873a: 16, pi. 3, figs 14-18 monly thin pedicels variable in number (up to (Fremantle, WA).-?Carter, 1886b: 122 (Port" Phillip 10) and length. Very firm.'Ieathery. Heads, Vic.).-Sollas, 1888:431 (holotype re-examined, Surface smooth to moderately but irregularly synonymised with T. ingalli). -Topsent, 1918: 583 tabuerculate, pale red to pink or dull orange. (resume, discussion, synonymised with Donatio Cortex thick (2 5-3 mm) diploderma). — Burton, dense, orange-buff, traversed 1924b: 1039 (holotype re- by regular radial pillars of megasclere examined, synonymised with Donatio diploderma).— bundles and of oxyspherasters Vosmaer, 1932-1935: 195 (resume). expanding into outer cortex with packed oxyspher- ''Tethya cliftoni. -Ridley, 1884b: 624 (discussion; asters; hence regular arcade-like cortical structure Seychelles). in sections. Large oxyspheraster of cortex and SPONGES FROM BASS STRAIT 31

peripheral choanosome regular, massive, mostly merge, overlap, and end below cortex, remainder with 12 conical rays. traverses cortex. Spiculation: Oxea, Description. Globular, slightly ellipsoidal, 4-4.5 cm (1) fusiform, straight, mostly as anisostrongyloxea, in diameter. 10 pedicels around base, 3-20 mm with blunt tips, some modified to styloids, strongyles, long, 1-4 mm thick. Surface in life pale red to pink rarely styles, 310-/045-1733 (5-7.5 R8/10). After preservation dull red-orange X 4.1-7/.7-17.4^m.(2)Stylesof cortex, variably flexuous, (10 R6/10 to 2.5 YR7-8/6) top and outer cortex, 249-425-548 x 4.2-6.4-8.2 /on. Oxyspherasters 12-16 orange-buff (5 YR9/4) base and inner cortex. (3) with massive, conical rays; in spicular slides, most Choanosome in first alcohol dull orange-yellow (7.5 common meristic form has six rays in focal plane YR7/10 to 2.5 Y8/8). Very firm, little compress- and three oblique rays ible, leathery. Surface mostly smooth, except one on top; diameter 20.9-60.5-72.4 side and top, with either faint polygonal tubercles ^m, centrum 26-52-40%. (4) Tylasters with 6-12 rays, mostly (2 mm wide) separated by narrow grooves, or a stocky occasionally slender, more numerous in pitted and rugo-reticulate pattern (pits and rugae cortex; diameter 9.9-/5.6-16.1 urn, centrum 0.5-1 mm wide). No oscules. 17-28%. (5) Oxyasters of the choanosome, centrum small or absent, with Cortex 2.5-3 mm thick, of two layers. Outermost 6-10 slender, commonly thin hyaline ectochrote, 30-35 /*m thick, with crooked and distally spined rays: proximal parts of rays frequently packed tylasters. Outer cortex 2 mm thick, very col- straight, distal, spined parts deviate at a lagenous, coarsely but vaguely fibrous. Few radi- low angle, or curved. Many forms show spines ally oriented fibres clearly visible. Inner cortex decreasing in length form middle of ray; initial, strongest spines radially traversed by pillars of crowded to packed may coincide with a tyle. rays oxyspherasters. Pillars mostly 300-350 /tm wide Some branch at this point; branches mostly uneven in length and thick- proximally, expand distally, in joining outer cortex. ness. Some transitional forms tylasters, General aspect of an arcade. Equally frequent, but to with stocky rays; some or all tyles in shape of flowers more irregularly spaced between pillars of spher- (anthasters), from which one stronger spine or asters, also radial, are columns of oxea, in width spiny actinal branch mav issue. Diameter generally inferior to pillars of spherasters. Oxea- 9.6-29.5-47 /tm. columns issue from choanosome, surrounded in

periphery by receding sheaths of cortical matrix. Remarks. The synonymy given above is tentative Columns compact through choanosome and most and follows that suggested and discussed by Berg- of inner cortex; expanding towards outer cortex, quist (1968: 36). But some further comments seem then often looser, branching through outer cortex, appropriate. Donatio ingalli sensu Dendy (1916a: ending flush with surface. Interstices of inner cortex 264) is a misidentification of Tethya robusta contain some radial lines of crowded spherasters Bowerbank, as Burton (1924b) correctly pointed and tylasters, commonly mixed. Width of these out. This also explains Dendy's following comment lines equals that of 1-3 spherasters. Some columns under Donatio seychellensis (1916a: 265): "The of oxea surrounded by scattered asters. Spherasters large red Donatio so common in the neighbourhood rare elsewhere in inner cortex, but tylasters scat- of Port Phillip, Victoria [the true Tethya ingalli], tered throughout. Chones indistinct, probably coin- possesses large oxyasters very similar to those of ciding with radial lines of crowded asters. One pillar D. seychellensis, but I am inclined to think, on of spherasters contains an excurrent canal. Inner account of other characters, that it is specifically cortex also contains subradial styles and styloids, distinct both from D. seychellensis and D. ingalli." irregularly flexuous, hastate to slightly fusiform, Indeed the whole spiculation of Tethya ingalli is most common below outer cortex, single or in very similar to that of T. seychellensis, as Hechtel's groups of 2-3, pointed ends outwards. Ends blunt, detailed description (1965: 65) makes clear. The mucronate, or conical to variable length, frequently cortical structure of the new specimen is sufficiently embedded in outer cortex. Some with bases in similar to that represented in Sollas (1888, pi. 44 peripheral choanosome. fig. 16) to justify the specific separation of T. ingalli Choanosome packed (periphery) to crowded with and T. seychellensis in the sense of Sollas and asters of three categories: two like in cortex, less Hechtel. inwards, frequent inward; third, more abundant As to Tethya diploderma, I concur with Hechtel oxyasters with thin, distally microspined rays. (1965: 66) that the species must be considered as

Compact radial columns of oxea, 1 80-200 urn wide, unrecognisable at present. There is, however, a equally distant, issue from small focal area of type-slide extant in the British Museum (Natural packed megascleres. One half of columns expand, History), BMNH 1870.5.3.93, cabinet 2, tray 52. 32 F. WIFDliNMAYBR

Burton (1924b: 1039) re-examined it, but gave no Burton, 1930c: 670 (definition, discussion). — Bergquist, details. Sollas' brief and insufficient redescription 1968: 21, 24 (key to New Zealand species). (1888: 439) is probably based on this slide. Top- Diagnosis. Encrusting to massive, sessile Poly- sent (1918: 574f.) was unaware of its existence. No mastiidae; papillae variable in number and length; complete survey of Schmidt's type material has ever without marginal fringe of radial spicules. been published. This applies particularly to the Tylostyles, subtylostyles, styles and styloids com- Atlantic material of 1870 (Museum of Compara- monly in three sizes. Pronounced cortex with tan- tive Zoology; unpublished survey by W.D. Hart- gential and radially bundled spicules, smallest man, Peabody Museum); to his material in the megascleres in palisade at surface. Chiefly littoral Copenhagen Museum, and his collections of types, and sublittoral. schizotypes and slides in the Landcsmuseum Joanneum, Graz, Austria. The specimens of the Polymastia crassa Carter latter collection are now temporarily in Geneva, Plate 2 figures plate 21 figures 3-5, curated Dy R. Desqueyroux-Faundez, and partly 7,8, being revised by her and by J.N.A. Hooper (per- text-figure 20 sonal communication). Polymastia bicolor . Unfortunately, many published records and war crassa Carter, 1886b: 120 (Port Phillip Heads, Vie.). short descriptions of Indo-Pacific sponges, assigned Polymastia crassa. - Dendy, 1 897: 250 (2 syntypes re- to Tethya ingalli, T. seychellensis, and T. examined, new records from Port Phillip Heads). diploderma, are dubious. Material examined. Station KG 7, two specimens (F51945, Polymastiidae Gray, 1867 F51946); station WB, one specimen (F51947). Type specimens Polymastia Bowerbank of Polymastia bicolor var. crassa (BMNH). Diagnosis. Base thin, encrusting; numerous papil- Polymastia Bowerbank, 1863c: 1 104. -Ridley and lae variable in width, low to moderately high Dcndy, 1887: 210.-lX-ndy, 1922: 148 (definition). - (mostly 5-20 mm), often contorted, irregular, wrin-

n n

k I «

i V

Figure 20. Polymastia crassa. spicules, a-b. Ranges of laroc stvUs

kled, some with apical oscules. Firm, little com- larly and in cortex expand to form confluent to pressible. Surface otherwise smooth. Colour overlapping umbels. Pattern may be obscured or variable, papillae commonly orange, base greyish replaced by large spicules in confusion. Outer orange, to reddish or dark purple, the latter occa- cortex formed by palisade of small spicules. sionally on whole sponge. Interior pale orange to Matrix of cortex finely fibrous, mostly without yellow. Cortex 0.7-1 .5 mm thick, with palisade of orientation, much lighter, in transmitted light, than small styles at surface. Deeper cortex may be choanosome. Contrast accentuated by interstitial vaguely stratified by layers of paratangential spic- confused spicules in choanosome, and by abun- ules. Choanosome dense, with spicules in confu- dance therein, of minute, vesicular to elongate and sion and in ascending bundles; these branch in contorted (sausage-shaped) bodies, presumably periphery, and often traverse cortex in umbels. algal symbionts. They also occur in cortex, but Walls of papillae reinforced by longitudinal spic- much less profusely. Between spicular columns, ular bundles. boundary with cortex usually bulges outward, with sheaths of cortical matrix penetrating deeply along columns. Description. Largest specimen (F51945, pi. 2 fig. Skeletal structure in intermediate specimen some- 7) pear-shaped in outline, 9 x 12 cm wide, 3 cm what different: spicular columns more distinct, high. Papillae crowded, stubby, with interstices more widely spaced, their branching in peripheral equal to their width or narrower. Papillae smaller choanosome and expansion in umbels in cortex on sides, 1-5 mm wide and high, larger (5-20 mm more regular. Interstitial confused spicules paratan- wide and high) in median zone. Many papillae con- gential in periphery. Second layer of paratangen- fluent, compressed and vermiculate in section; tial spicules below outer 3/5 of cortex. Cortex in larger ones bear longitudinal folds and wrinkles, papillae only 330 /mi thick, spicular columns more and are mostly inclined to recumbent. Apical crowded than in base. Boundary with cortex rarely oscules rare on small papillae, more frequent on bulges between spicular columns. larger ones. Spiculation (individual measurements): F51945: In life salmon to orange on papillae (10 R - 2.5 (1) Large styles, strongyles, strongyloxea (styloids), YR7/8-10), greyish red around base (5 R6/2-3), 492-677-699 x 7.1-70.7-16.2 jim. (2) Medium with pale orange to yellow (10 YR - 2.5 Y8-9/6-10) styles, subtylostyles, strongyloxea (distinction from in cortex and choanosome. Firm, little the large spicules is somewhat arbitrary, but inter- compressible. mediate sizes are rare), 311-407-462 x 5.6-7.4-9.1 Specimen F51946 (pi. 2, fig. 8) growing on sand, /an. (3) Small styles and tylostyles, 124-755-188 x encrusting base of indented outline, 2-3 mm thick, 2.4-4.2-5.3 fim. F51946 : (1) Large styles, stron- wide. 21 tapering papillae more 7 X 5 cm mostly gyles, strongyloxea (styloids), 477-750-964 x widely spaced than in larger specimen, often com- 7.6-/7.4-15.3 pan. (2) Medium styles, tylo- and sub- pressed, 2-8 wide at the base, up to 15 mm mm tylostyles (with small heads), oxea, 179-252-340 X long. Some weakly rugose longitudinally, mostly 4.7-7-9.1 urn. (3) Small tylostyles, 100-729-177 x recumbent in different directions, contorted. Some 1.9-2.9-4.4 /on. bear minute apical oscules. Base incrusted with sand. Papillae bright orange (2.5 YR7/12), base greyish orange (2.5 YR7/3). Rubbery. Remarks. Carter (1886b: 119, 121) also described Specimen F5I947 similar in shape to F51946, but Polymastia bicolor (as typical variety) and P. mas- smaller: base 3 x 4.5 cm wide, 1-2 mm thick. salis from Port Phillip Heads. Dendy (1897: Papillae crowded, not above 3 mm wide at the base, 249-250), who re-examined the types of both, latter variety of P. bicolor 5 mm long. Very dark purple, almost black, regarded the as a and that this species belongs to Polymastia, thoughout (5 RP3/1). Now (in alcohol) dull yel- doubted lowish brown (10 YR5-6/3-4), other two specimens stating: "It appears to be intermediate in structure Suberites Polymastia." I now cream-coloured. Cortex in largest specimen between the genera and examined these types (BMNH), particu- 1-1.5 mm thick in base, 700-870 /mi in intermedi- have not type slides, and cannot comment on this ate specimen. Choanosomal skeleton, in former, larly the the four species described by Bergquist of closely spaced subparallel columns, 40-180 urn issue. Of three are known from New Zealand only, thick, of large megascleres. They frequently branch (1968), Bowerbank also from Britain and from and anastomose obliquely; some break up into dis- P. conigera Ocean. The latter is spherical with a orderly bundles. Interstitially crowded intermedi- the Indian single osculiferous papilla. P. hirsuta Bergquist has ate spicules, some larger spicules in confusion. surface pile and involute papillae. P.fusca Berg- Below cortex, spicular columns branch more regu- a 34 f. wilde ;;nmayer

quist and P. granulosa Brndsted are closer to P. S. areolata Dendy (1897: 255) may well be a good crassa in shape and surface, but have a distinct, species, characterised by a regular polygonal- uniform colour: dark chocolate brown externally areolate surface (Vosmaer, 191 1, pi. 3 fig. 4). This in P.fusca, mostly conserved after fixation, deep pattern is reminiscent of Raphyrus hixoni Lenden- orange throughout in life in P. granulosa. P.fusca feld (1885) described from Sydney. also differs from P. crassa in cortical structure S. digitata Hentschel (1909) seems to be suffi- (packed with spicules, small subtylostyles often ciently distinct by its shape, colour (orange-brown exotylote), while in P. granulosa the large sub- surface, whitish choanosome) and ecology. S. tylostyles are commonly polytyloie. massa Ridley and Dendy (1886) (holotype from Bass Spiraslrellidae Ridley and Dendy, 1886 Strait, BMNH 1887.5.2.33, not examined by me) is described as massive, pale yellow, with very Spirastrella Schmidt diffuse skeleton, and commonly slender spirasters with up to six bends. Vosmaer (191 1: 17), who re- Spirastrella Schmidt, 1868: 17. -Wilson, 1925: 345 (dis- examined the holotype, regards it as very close to cussion). - Bergquist, 1965: 185 (discussion). - .S'. wilsoni Wiedcnmayer, (correctly .S". purpurea sensu Topsent), 1977a: 162 (synonymy in pari, not Spiras- except for colour. trona, = Diplastrella; references). It may therefore be conspecific with Suberites wilsoni var. albidus Carter (holo- Diagnosis. Incrusting to massive Spiraslrellidae of type: BMNH 1886.12.15.256; spirasters figured by variable, often irregular shape. Tylostyles and sub- Vosmaer, 191 1, pi. 12 fig. 3). This species, if future tylostyles, subordinate^ styles, in more or less iso- revision based on primary types and topotypes lated ascending bundles. Spirasters mostly coarse, should confirm its validity, would have to be called scattered to packed in choanosome, often concen- Spirastrella utbida (Carter). Suberites spirastrel- trated in superficial crust, absent in some speci- loides Dendy (1897) is probably another mens. Boring svnonym. assumed in two species. Colour in Vosmaer ( 1 91 1 : 23) observed "traces of red color" life commonly yellow, brown, grey or deep purple, on the holotype of Suberites wilsoni var. albidus. never blackish. The possibility should not be excluded that purple and yellowish or brownish specimens occur Remarks. within This genus is notoriously difficult and one species (S. purpurea), with transitions, as this controversial in taxonomy, especially in delimita- seems to be the case in the Japanese Spirastrella tion of species. Vosmaer's (191 1) extreme lumping pant's Thiele, as redescribed by Hoshino was criticised (198P by several authors (notably 224).

Hallmann, • 1914a: 291!'.; Topsent, 1918- 542 f S. spinispintlifera (Carter), first described from Wilson, 1925: 345). South Africa, then from Victoria, Of the Indonesia and species recorded from southern Austra- New Zealand (Bergquist, 1968: 16) is sufficiently lia, S. purpurea (Lamarck), with synonyms S. distinct wil- by its weakly spined spirasters soni (Carter) and (spinispirae). S. bonneti Topsent, and A number of species from the Indo-West Pacific redescribed by Topsent (1918: 546; 1933: 43), is dis- reg.on have been described up to 1905 (S punc- tinct chiefly by its deep, stable purple colour (spon- tata R.dley, S. vagabunda Ridley, S. solida gioporphyrine; see Bergquist Ridley and Skinner, 1982, pi. and Dendy, S. inconstans (Dendy), S. transttoria 7 fig. 3). Its thick cortex is differentiated into an Ridley, S. cyltndrica Kieschnick, S. inner zone of felted megascleres, aurivillii Lind- and an outer zone gren, S. semilunaris of packed Lindgren, S. carnosa Topsent spirasters and erect megascleres. S lacunosa Kieschnick, S. spiculifera Kieschnick' Spirasters arc mostly small, feebly spiral, often S. tentortoides Dendy) which approaching amphiasters. might be good spe- In some specimens, cies. This applies particularly to those microscleres may be rare or lacking, redescribed and most more recently: S. megascleres vagabunda and S. aurivillii by may be modified or reduced, often Bergquist (1965: 182f.); with subterminal S. inconstans by Vacelet tyles or without tyles (with styles et al. (1976: 33); S. carnosa and even strongyles and 5. solida by resulting). Sand is commonly Uey UX FaUndeZ (1981: incorporated throughout 734 >- Vaeelet et al. the sponge body. The (1J76:no^ L 1 : 34) have shown that presence of S. cunctatrix species in the in southern Australia mconstans-vagabunda group, with similar needs to be confirmed, but is not and vari- unlikely from its able morphology, can be distinguished in known distribution (Western Atlantic, Mediterra- the field C al affinitieS ° th°l, h attribution nean, Indo-West Pacific [see Desqueyroux- 7,u ' S to any1 ot the oldTspecies remains Faundez, 1981: problematical Their 733], North Australia [as Spiras- spcc.es 1 prefers back-reef trella decumbens Ridley], habitats (sand Hats with Hawaii, Gulf of sea-grass, mangroves); their California). species 2 occurs in shaded and sheltered recesses in shallow portions :

SPONGES FROM BASS STRAIT 35

of the coral reefs; their species 3 inhabits somewhat Material examined. Station KG 10a, one specimen deeper portions of the fore-reef. 5. auhvillii, as (F51948); station BSS 187, two specimens (F51949, redescribed by Bergquist (1965), besides contain- F5 1 950). Holotypes of Spirastrella papillosa and of Spiras- ing characteristic spirasters, has a predominantly trella cunctatrix var. porcata (BMNH); type specimens of Spirastrella boring habit, largely hidden in fairly large cavities papillosa var. porosa (NMV) and of Pupil- lina panis in coral rock, with papillae protruding through (AM). Hypotype of Spirastrella cunctatrix: Carter straight channels. (BMNH 1886.12.15.250).

Diagnosis. Shape variable, often massive, like turtle-shell, also thickly flabellate, radially folded, Spirastrella papillosa Ridley and Dendy with wide, truncate margin containing regular depressions. Firmly spongy. Oscules marginal. Sur- Plate 2 figures 10, 11, plate 21 figure 6, face except margins commonly nodulose or papil- text-figure 21 late, but this relief sometimes subdued; otherwise smooth. Deep yellow, olivaceous or Spirastrella papillosa Ridley and Dendy, 1886: 491. - purplish grey, internally light yellowish brown or Ridley and Dendy, 1887: 232, pi. 41 fig. 5, pi. 45 fig. I] dark olive-grey. Cortex 0.7-2.5 (Port Jackson). - Dendy, 1 897: 253 (synonymy; Port Phil- mm thick, full of felted megascleres, lip Bay).- Whitelegge, 1897: 331 (Funafuti). -Vosmaer, more croweded (packed) at surface, as for 1911: 17, 33, 41, 56, 60, 61 (references, resume, type data, spirasters. Choanosome finely and regularly caver- description, discussion). -Hallmann, 1912: 126 (Shoal- nous, with felted megascleres mostly in thin con- haven Bight, Port Jackson, NSW, Port Phillip Bay). torted walls. Spirasters intermediate in size (for Cliona papillosa. -de Laubenfels, 1936a: 142 (trans- genus), rather robust, shafts often straight, terete. fer only). Spirastrella papillosa var. porosa Dendy, 1897: 253 Description. Shape and size of first specimen (pi. (Port Phillip Heads).- Vosmaer, 1911: 17 (discussion). 2 fig. 10) like figure in Ridley and Dendy (1887, Spirastrella cunctatrix. -Carter (nee Schmidt), 1886b: pi. 41 fig. 5). Roughly pear-shaped, 11 x 15 x 114 (pan, only BMNH: 1886.12.15.250, Port Phillip 8 cm; flat base of attachment almost circular in out- Heads, figured in Vosmaer, 191 pi. 3 fig. 3, pi. 10 fig. 2). 1, line, 9 cm in diameter. Most of free side covered Spirastrella cunctatrix var. porcata Carter, 1886bl 1 15 with crowded round papillae, 5-8 mm in diameter, (Port Phillip Heads). -Vosmaer, 1911: 10,27,33,39,61 2-3 mm high. Two blunt conical elevations (references, resume, type data, decriplion, discussion), on top, pi. 10, fig. 3 (spicules of holotype). and on one side near base, respectively. Papillae Pupillina in upper half tend to panis Lendenfcld, 1888: 58 (part, not pi. 1 be aligned longitudinally, and figs 1, 2, fide Hallmann, 1914a; Port Jackson). - coalesce in apices of osculiferous cones, passing Whitelegge, 181. 1889: -Hallmann, 1914a: 294 (synlypes into longitudinal folds separated by narrow fur- redescribed). rows. Apical oscules, now closed, have deeply Spirastrella - purpurea. Vosmaer (nee I.amarck), 191 1 incised, irregularly stellate margins. Deep yellow 17 (part, specimens S.E. 426d, 92a-l only), pi. 3 fig 5, in life (10 YR to 2.5 Y8/20) on outer surface, light pi. 4 fig. 3, pi. 9 fig. 6, pi. 10 fig. 1, pi. 14 fig. 1. (Selat yellowish brown (7.5 YR7/4-6 and 7.5 YR8/4-8) Sape, Flores Sea, Indonesia). around base and in choanosome. Other two specimens thickly flabellate. F 51949

(pi. 2 fig. 1 1) 2.5 cm thick near base, 1.5 cm distally. Width and height 14-15 cm. Free margin three-quarters-circular in outline, truncate with several contiguous squarish depressions. Small oscules situated partly in these •^ depressions, partly irregularly scattered on one side. Papillae on both sides smaller, more numerous, lower than in F51948, often vague and coalescent. One side

coarsely corrugated longitudiually. Olive in life. F51950 a section (about one-third) of irregularly crescent-shaped lamella, 18 cm wide, 10-11 cm high, up to 5 cm thick at base. Both surfaces more vaguely and irregularly nodulose than in previous specimen. Surface purplish slate-grey, dark olive- grey in choanosome. All specimens firmly spongy, Figure 21. Spirastrella papillosa, spicules from F51948. internally finely cavernous. a. Range of megascleres, x68. b-c. Enlarged ends of Cortex variable in thickness, 700 /xm in F51948, megaselercs, x263. 1. Spirasiers, x 790. 2.5 mm in F5 1949, 1 mm in F 5 1950. Full of felted 3

36 F. WIEDENMAYER

megascleres, with rnicroscleres scattered in lower was less often used since its revival by de Lauben- half to three-quarters, crowded further out (zone fels (1936a). commonly pigmented), packed at surface. Surface Chondrosia Nardo layer, 80-170 pim thick in F51949, not always distinct, often with still felted megascleres, either flush Chondrosia Nardo, 1847b: 267. — Wiedenmayer, 1977a:

with surface or protruding from it. In F51950 187 (synonymy, references). spirasters gradually decrease in abundance from Diagnosis. Chondrillidae without spicules. surface to middle of cortex. Foreign debris widely scattered throughout choanosome and cortex in Chondrosia reticulata (Carter) two flabellate specimens. In massive specimen Plate 3 figures 1-6, plate 22 figure 1 choanosome often packed with sand, while cortex contains very few grains. Halisarca reticulata Carter, 18861": 274 (Port Phillip

Spiculation (in F51949) : (1) Tylostyles, sub- Heads and Westernport Bay). tylostyles and styles, straight, curved or bent; with Chondrosia spurca Carter, 1887a: 286 (Port Phillip tyles frequently subterminal, faint, or composite, Heads). — Carter, 1887c: 356 (synonymised with Halisarca occasionally tilted, or with second, faint subtermi- reticulata). -Topsenl, 1896b: 513 (synonymised with Chondrosia collectrix Lendenfeld). — Vosmaer, nal tyle; points may be telescoped, mucronate, 1932-1935: 289 (resume).- Wiedenmayer, 1977a: 188 strongyloxeate or strongylate; 184-554-510 x (resume). 2.6-6-8.4 fim. (2) Spirasters mostly straight to Chondrosia reniformis forma spurca. — Burton and gently twisted, occasionally curved or bent, irregu- Rao, 1932: 324 (discussion, in part). larly or spirally spined, with distinct terete shafts, Chondrosia collectrix Lendenfeld, 1888: 74 (net- total length 32. 1-40. 1-49 .4 fim, width of shaft Schmidt, 1870; Port Jackson). -Hallmann, 1914a: 307 2.8-5.7-4.5 /an, width with spines 10.2-/5. /-19. (redcscription and discussion of holotype). — nee Bnj>ndsted, 1926: 319 (a Psamniopemma fide Bergquist, 1968: 63). Remarks. In morphology, the spicules in the new Reniera collectrix Lendenfeld, 1888: 78 (Port Jack- specimens, particularly the rnicroscleres, compare son).- Whitelegge, 1902b: 277, 280 (redescription of 2 well with pi. 10 figs 2, 3 in Vosmaer (1911), syntypes).-Hallmann, 1914a, b: 308, 327 (redescription representing those in Carter's two specimens from of 2 syntypes, synonymised with Chondrosia collectrix Port Phillip Heads, quoted in synonymy above, Lendenfeld). Chondrosia chucalla de Laubenfels, and with pi. 45 fig. 1 1 in Ridley and Dendy (1887). 1936a: 184 (nomen novum for Chondrosia collectrix Lendenfeld as secon- Of the measurements tabled in Vosmaer (1911: 61), dary junior homonym). -?de Laubenfels, 1951: 267 and based on re-examination of the type specimens, (Hawaii). -?de Laubenfels, 1954a: 154, text-fig. 178 those for the Victorian specimens (B.M.I, B.M.10) (Marshall Islands). -?de Laubenfels, 1955a: 142 (Gilbert again agree with mine (Dendy, 1897, gives 200-460 Islands). x 4-8.3 nm for tylostyles, 50 x 12 /un for Material examined. Station rnicroscleres). In the holotype of Spirastrella papu- BSS 187, one specimen (F51951). Type specimens of Halisarca losa, Vosmaer found maximum thickness of reticulata and Chondrosia spurca (BMNH). Type specimens of Chon- tylostyles to be 17 /un (Ridley and Dendy, 1887, drosia collectrix and Reniera collectrix (AM and BMNH). give 15.7 /urn), i.e. twice the size in my measurements. Vosmaer's own specimens from Indonesia Diagnosis. Encrusting to irregularly massive, firm, rubbery. Surface (quoted in synonymy above) are comparable to the smooth, glabrous, to finely rugose or verrucose, holotype from Port Jackson in dimensions of deep purple in life, light beige or greyish when preserved. megascleres (range: 183-600 x 15-20 /on) and in Oscules mostly minute, scattered. Cortex the segregation of spirasters into two size-categories variably thick, 0.5-1.5 mm in massive larger (22.5/55 /tm in the holotype, 12.5-14/40-85/im in specimens, little differentiated except for absence of Vosmaer's material). This may reflect a bio- foreign detritus, which is otherwise scattered geographical gradient; but more material from Port to packed through most of the choanomsome. Peripheral Jackson and other localities will have to be choanosome sometimes free of detritus, examined. with irregularly meandering canals, paratangential below cortex.

Description. Chondrillidae Gray, 1872 Massive, bluntly ridge-shaped, with asymmetrically triangular cross-section, flat base' Remarks. Gray's name has priority over Chon- of attachment. 4 x 8 cm at base, 3.5 cm in height drosiidae Schulze, 1877 (in the modern sense, with Partially enveloped clump of coralline algae and Chonclrilla generally included) though the former bryozoans on one side. Deep purple in life, firm, SPONGES FROM BASS STRAIT .17

little compressible, rubbery. Now faded to light in Reniera colleclrix. The syntype (syntypes?) in beige, with light brownish to greyish areas. Surface Sydney (AM G9014, wet, two halves or two cut on sides rugose to verrucose without orientation, specimens, pi. 3 figs 4, 5) is light yellowish brown otherwise smooth, glabrous. Elevations 2-4 mm internally, speckled dark greenish grey. wide, depressions similar or narrower. Apical ridge The West-Indian Chondrosia colleclrix (as 1-1.5 cm wide and glabrous. Most oscules minute redescribed in Wiedenmayer, 1977a: 189) differs

(100-800 /iiti wide), scattered on ridge, few from C. reticulata by its darker, almost black clustered down one narrow side. Single larger colour, softer consistency, and by having a thin, oscule, 2.5 mm wide, on one of elevations below detachable ectosome. ridge. Cortex 0.5 to 1.5 mm thick, marked by absence of foreign detritus. Matrix of cortex Latrunculiidae Topsent, 1922 vaguely and finely fibrous, without orientation Remarks. The sigmatose early ontogenetic stages except along subradial, meandering chones. Excur- of discorhabds in Sigmosceptrella and the frequent rent canals subradial in cortex, equal in width to occurrence of polytylote megascleres in the family smaller oscules. Subcortical crypts or excurrent (the latter stated to be common only in various canals (connections not seen) meandering in poecilosclerids) prompted Topsent (1922) to include paratangcntial plane, 0.5 mm wide, with smooth the Latrunculiidae in the Poeciloselerida. He reiter- walls. Similar canals meandering deeper in choano- ated this view (1928: 47), granting a common, some, surrounded by fibrous matrix, without though distant origin of Spirastrellidae and Latrun- preferred orientation. culiidae. His view was upheld by Levi (in Brien et Foreign debris scattered to crowded throughout al., 1973: 61 1). Bergquist (1978: 166) included the choanosome, packed in some places, ill-sorted, up family in the Hadromerida, based on earlier to 600 ftm in diameter. Choanocyle chambers biochemical work, followed by Boury-Esnault and mostly spherical, 25-36 urn in diameter. Cortex and van Beveren (1982: 42). Van Soest (1984: 146) left choanosome contain large quantities of translucent the ordinal assignment of the family open, provi- microsymbionts, of one, commonly two to three sionally following Levi (1973). aggregated globular cells, diameter 7-12 /un. Most Polytylote monactinal megascleres also occur in colourless in transmitted light, scattered to Polymastia (P. granulosa, see Bergquist, 1968: 23; crowded, those lining canals commonly packed, P. polytylota, see Pulitzer-Finali, 1983: 490). Early single, smaller (not exceeding 8 jtm), light yellow- ontogenetic stages of discorhabds in Sigmoscep- ish. Another form of symbiotic alga, mostly trclla are not like typical sigmata and chelae, in attached to sand grains, irregular, amoeboid to having a straight shaft, and spines always appear- digitate-frondose, light yellowish green. ing simultaneously on distal recurved ends and Remarks. The syntypes of Cliondrosia spurca and opposite on the shaft. The homology with those of Haliscarca reticulata relabelled (by poecilosclerid sigmatose microseleres is weakened Carter?) "Chondrosia spurca", in London by the rhabdose erliest stage in diseorhbds of loveni (see Remarks on Latrunculia (BMNH), which 1 have examined only macroscop- Podospongia of ically, compare well with the new specimen. The below). Another type "pseudosigmatose" phylogenetically related to spirasters, largest one (the leclotype, BMNH 1886.12.15.128, microsclere, in anthosigma. Similarity by wet, with slide G2756 in the Australian Museum) might be seen the con- has deep portions of the peripheral choanosome vergence in these "sigmatose" microseleres seems to that in dentate chelae and sig- almost devoid of sand (pi. 3 figs 2-3). The mean- lo be analogous 1921: fig. the dering excurrent canals in these areas are up to 3 maspires (Dendy, 119, 31). On other hand, the presence of true chelae (palmate mm in diameter. In the syntype of Chondrosia col- wet), the anisochelae) in liarhozia primitiva Dendy (1922: lectrix Lendenfeld in Sydney (AM G9061 , 1311.) might be adduced as a stronger argument for cortex is thinner than in the Victorian specimens, peocilosclerid affinity of the Latrunculiidae (Dendy so that individual foreign inclusions or their clusters as included the genus in the Spirastrellidae). However, in the underlying choansome are reproduced skeleton, and Dendy's statement, that conules or fine tubercules (pi. 3 figs 6a-e). The its reticulate present speci- its "oxydiscorhabds" are not arranged in a special colour is somewhat darker than the like in Lalrunculiu, make it certain men, greyish. The microsymbionts, described by surface layer 15-20 is an aberrant poecilosclerid (as de Hallman as "cystencytes", are slightly larger, that Barbozia kind of Laubenfels, 1936a: 53, and Topsent, 1928: 48, pm in diameter. Possibly a different Its "oxydiscorhabds" (Dendy, microsymbionts was described by Flallmann maintained). 1921, 1922, pi. 18 fig. Id) are remarkably simi- (1914a: 309) as being confined to the smaller canals fig. 34C; 38 F. WIEDENMAYER

lar to some amphiasters in the spongillid Dosilia tirotalis Topsent (1928, pi. 7 fig. 19b). Three types plumosa (see Dendy, 1921, fig. 26M). Topsent (figs 22, 23, 45) have their upper half decorated by (1928: 48) speaks of "isancres amphiasteriformes." microspined or roughened globules, several in one Levi and Levi (1984: 956) argued convincingly that whorl and one apical, similar to one-half of a mas- the microsclere-complement is unstable in this sive amphiaster of the clionids Thuvsa and Alec- genus (sigmata or ioiui (figs "oxydiscorhabds" may be 41, 42 of the same plate). Two types (figs. absent), and that Barbozia is a junior synonym of 32, 44) are close to discorhabds in the modern Phlyctaenopora Topsent (1904), which they assign Didiscus. That in figure 32 is uniformly to the Coelosphaeridae. microspined throughout and its discs are marginally The fossil record has not been sufficiently inflated. Similar, isolated examples are known from appraised in this context. Though it does not Recent sediment of the Indian Ocean (Carter, resolve the issue, it is apt to add a new perspec- 1879b, pi. 29 fig. 20) and from the Upper Jurassic tive. The oldest fossil discorhabds are those figured of Poland (Wisniowski, 1888, pi. 12 fig. 10). Such by Rust (1885, pi. 45 figs 35, 36), now known to types particularly in connection with the occasional be of Lower Cretaceous (Albian) age. That of fig- slight swelling of blunt apices of the axis, as in ure 36 is remarkably similar to the modern type in Didiscus Jiavus van Soest (1984), suggest transitions Carter (1879b, pi. 29 fig. 16). From the same phos- to, and homology with short criccorhabds with few phorite nodules from Northern Germany, Rust annul! (Wiedenmayer in Hartman et al., 1980: 72), figured spirasters (pi. 45 figs 34, 38) and what looks with the microsclere type in Carter (1979b, pi. 29 like an anthosigma (pi. 45 fig. 37). Normal sigmata, fig. 21), and microspined, nodular amphiasters in forceps and chelae (some of otherwise unknown Alectona and Thuosa, which may be asymmetri- types) are also reprented (pi. 45 Tigs 9-11, 13-15, cal (Topsent, 1904, pi. 12 fig. 3b; Annandale, 17, 20, 21) and so are tylote sigmaspires (pi. 45 figs 1915a: 18, text-fig. 3). Those in Topsent (1904) are 12, 16). Normal sigmata, however, are known also suggestive of the type attributed to Alectona already from sediments of Triassic age (Mostler, by Hinde and Holmes (1892, pi. II fig. 44). Its 1976, text-fig. 12). shaft is smooth, strongyloxeote, and bears two Some modern discorhabds are exceptionally large symmetrically arranged discs with serrated margins, for microscleres (up to 217 /mi long in Latrun- like in Didiscus, but, in addition, a few irregularly culia multirotalis Topsent, 1928: 222; up to 377 pm distributed tubercles between discs. Other parallels long in L. biformis, fide Boury-Esnault and van are the microspined sanidaster of Latrunculia Beveren, 1982: 44). Even larger are the giant "dis- (Negombata) acanthosanidustcra (Hoshino, 198 1 : casters" in SceptrintusTopsent (1904: 1 17), a genus 221, fig. lib; pi. 4 figs 3, 4), with its blunt'clads; now virtually extinct, known only Tram the type and the amphiasters of Thoosa laeviaster Annan- locality in the Azores. These spicules, more aptly- dale (1915a: 22) resembling early stages of, or described as hypertrophied verticillate sanidasters, atrophied discorhabds of Latrunculia. have sizes of 150 x 7 /mi to 530 x 40 /mi. They The microsclere attributed to Thoosa by Hinde were regarded as megascleres by de Laubenfels and Holmes (1892, pi. 12 fig. 3, 30 pm long) com- (1936a: 79), who was undecided whether to include bines two structural elements: the massive the genus in his "Acarniidae" or in the Agelasidae. amphiaster of Thoosa mollis Volz (Pulitzer-Finali Topsent (1904), however, has clearly stated that 1983: 502, fig. 30 A-C) and the denticulate axial these spicules are arranged like spirasters in Spiras- apices like in many discorhabds of Latrunculia Of Irella and thus should be ranged as microscleres. the microscleres ascribed to Spirastrella bv Hinde He therefore included Sceptr'mtus in the Spirastrel- and Holmes (1892), those on plate 12, figures 4 and lidae rather than in the Latrunculiidae. Sceptrintus 6, can be regarded as transitional between spirasters must have been very common in the Oligocene and discorhabds, with some around spines still in verticils Tasmania (Kennett, Houtz et al., 1975: 333, or clustering around the bulbous apices. figs Analogous 10-12), and possibly occurs today at great transitions were pointed out by Dendy (1921-123 depth in the Gulf of Mexico (Spongolithis hystrix fig.. 39) in Latrunculia (Negombata) cor'ticata Ehrenberg, 1873, pi. 5 fig. 22, isolated spicule in Carter. dredged sediment, 2720 m). The tendency, common to some spirasters Among the fossil and microscleres from the Oamaru irregular discorhabds, to approach amphiasters Diatomite of New Zealand figured by Hinde and (compare e.g. Diplastrella biste/lata, Suberites wil- Holmes pi. 1 (1892, 1 figs 15-45) are many typical SOmm Vosmaer, 1911, pi. n figs discorhabds of 4, 9, with Latrun- Latrunculia. The largest (fig. 34) culiix hallmanni, below, and Latrunculia is 206 ixm long and resembles those of L. mul- (Negombata) tarentina Pulitzer-Finali, 1983, fig, SPONOES FROM BASS STRAIT .19

37) has been noted by Topsent (1928: 47). Com- in Didiscus, some are known in Latnmculiu or Sig- parison of the fusiform vcrticillate saniciasters and mosceplrella (Hinde and Holmes, 1892, pi. II, fig. 7-whorled discorhabd in Hinde (1910, pi. 1 figs 15; Dendy, 1922, pi. 18 figs 4b, 4b'; Bergquist, 11-13; Late Eocene) with the sanidasters from the 1978, pi. 7b, here with dichotomy at right angles, Oligocene of the continental rise off Tasmania i.e. in axial planes, a feature also apparent in some (Kennett, Houtzet al., 1975: 333, figs 10-12) sug- fossil examples in Hinde and Holmes (1892), in gests a connection between Sccptrinlus and Latrun- some extremes resulting in erect apical blades). The culia, closer than Topsent imagined. I wo types of "Ditriaenella" differ only in the The spicules ascribed to a new (hypothetical) dichotomy being proximal in figure 35, with six genus Ditriaenella by Hinde and Holmes (1892, pi. equal clads in one whorl (in analogy to the 12 figs 34, 35) merit consideration in this context. amphiaster of Thoosa). Amphiasters in Samus The one in figure 34 was reproduced by Rauff (possibly derived from "Ditriaenella" amphiasters (1893, p. 155, fig. 11) and named by reduction of the rhabd ends) are often heter- "amphimesodiehotriaene." The closest modern opolar (Carter, 1879b, pi. 29 fig. 3, here equivalents in my oponion are the "amphitriaene" microspined; Topsent, 1928: 28; Pulitzer-Finali, of Samus and the normal amphiaster of Thoosa 1983, fig. 17, right side), and in this trait they are (Wiedenmayer, 1977a: 33, figs 29, 30). With regard comparable to some fossil anisodiscorhabds, par- to the homology of the latter two, 1 am merely fol- ticularly the curious stubby types in Hinde and lowing Topsent's opinion (1928: 28), countering Holmes (1892, pi. II figs 29-31) with only two Sollas' influential and misleading view (1888: 57), whorls and no apical extension of the axis (their who qualified the amphiasters of Samus as similarity to candelabra is here regarded as due to amphitriaenes. Topsent was cautious with regard convergence). to the other, smaller microseleres of Samus, quali- Amphiasters of Samus, with maximum dimen- fied as sigmaspires by Sollas, a view followed by sion (diagonally) up to 200 /mi, are considerably most subsequent authors (e.g. Pulitzer-Finali, 1983: larger than amphiasters of Thoosa (up to 37 jan 481 ). These microseleres are C-or S-shaped, in one long in normal types, Wiedenmayer, 1977a, fig. 30; plane or twisted, microspined (not smooth as up to 70 nm in the large amphiaster of Thoosa claimed by Sollas, 1888, and Lendenfeld, 1903). mollis: Pulitzer-Finali, 1983: 582). The fossil spic- They are thus indeed morphologically similar to the ules of "Ditriaenella", which could be regarded common microseleres of Spirophorida. Topsent, equally well as amphiasters and as special oxydis- rejecting the homology with spirasters, ultimately corhabds, are 185 pan long. They, like those of accepted them as sigmaspires, and admitted a dis- Samus, can well be qualified as microseleres, con- tant relationship with the Spirophorida (in which sidering the size ranges in discorhabds and sigmaspires would be the only ancestral trait shared sanidasters of Latnmculia and Sceptrintus cited with Samus). Considering the presence of above. spinispirae (microspined spirasters) in Spirastrella The special minute microseleres localised in the (S. spinispirulifera, see Bergquist, 1968, pi. 11 G) surface of papillae of Cliona levispira and Dolona and the similarity with the spiroscleres in Trachy- pulchella (Topsent, 1904, pi. 12 figs Id, 2c) can be cladus (Bergquist, 1970, pi. 10C/2), convergence regarded as reduced discorhabds, with lengths (6-10 of such microseleres cannot be excluded. Reduc- jim) less than one-third of those in the smallest tion of spination and of twisting of spirasters in modern lalrunculiid discorhabds, and about half ancestors of Samus might well have occurred. as long as the smallest fossil one (Hinde and

The fossil spicule of "Ditriaenella" connects I he Holmes, 1892, pi. 11 fig. 27, 18 /t m long). Cliona amphiasters of Thoosa and Samus with certain acuslella Annandale (1915a: 14) contains similar types of discorhabds, in which verticils are more microseleres exclusively. or less regularly lobed, with lobes bifid, trifid, or The analogy with the Chondrillidae in the sup- multifid (the latter in Hinde and Holmes, 1 892, pi. pression of megascleres in Samus (frequently also

1 1 fig. 40). Individual verticils of such types com- in adults of Aleetona and Thoosa: Topsent, 1928: monly comprise four lobes (Sigmosceptrella quad- 28) should be pointed out in this context. The ques- rilobala) or three lobes (S. fibrosa). Adjoining tion whether the microseleres of" Ditriaenella" and verticils are often twisted regularly around the axis, Sceptrintus are ancestral or derived with respect to so that lobes alternate between whorls when viewed discorhabds in Latrunculiidae has to remain open. axially. In such view some examples show tendency The much earlier record of the latter (Early Cretace- to dichotomy (Topsent, 1904, pi. 12 fig. 5; 1928, ous) is probably fortuitous. Furthermore, all fossil pi. 7 fig. 18b). Oxydiscorhabds are not exclusive types are isolated: we therefore do not know what 40 F. WIEDENMAYER

complements occurred in species of the geological second species is Sigmosceptrella quadrilobata past. However, the balance of these considerations Dendy (1921, 1922), from the Indian Ocean. Fully favours a common ancestry of Latrunculiidae, grown microscleres of this genus are similar to those Spirastrellidae and Clionidae, and their differen- in Latrunculia conulosa, described below, but their tiation in Early Cretaceous time or earlier. early ontogentic stages are always "sigmatose". The diagnostic value of this trait is Latrunculia Bocage lessened by the ontogeny of discorhabds in Poclospongia loveni du Latrunculia (Latrunculia) Bocage Bocage (as redescribed by Topsent, 1928: 219, pi. 7 fig. 17) being intermediate: earliest stages Latrunculia Bocage, 1869: 161. -Ridley and Dendy, as 233. slightly wavy rhabds, young stages similar 1887: -Dendy, 1921: 121.-Topsent, 1922: 7. to those Sceptre/la Schmidt, 1870: 58. -Ridley and Dendy, in Dendy's drawings, but more irregular. 1887: lxii, 234 (synonymised with Latrunculia). -van Negombata de Laubenfels (1936a: 159) has as Soest, 1984- 149 (synonymised with Latrunculia). type species (by original designation) Latrunculia Negombo Dendy, 1905: 127. -de Laubenfels corticata Carter (1879b: 298) from the Red Sea. The 1936a: 132. "genus" is distinct only by having diactinal instead Diacarnus Burton, 1934a: 549. -de Laubenfels, 1936a- of monactinal megascleres, and is 133. therefore better treated as a subgenus. Oxylatrunculia Hoshino Diagnosis. Sanidasters transitional to discorhabds; (1981: 221, type species by monotypy O. acan- whorls of spines more than two, but not always dis- thosanidastera Hoshino) is a junior synonym. A tinct (some types transitional to amphiasters). third species is Latrunculia purpurea Carter (1881a: Microscleres may be symmetrical or asymmetrical, 380) from Bass Strait, not recorded again. Two with one end pointed (anisodiscorhabd). more species occur in the Red Sea and a sixth spe- Amphiclads (reduced discorhabds with spines only cies was described from the Mediterranean as in distal whorls) may be present with transitional Latrunculia tarentina by Pulitzer- Finali (1983: types (Topsent, 513, 1892: 128; 1928: 220). Basic rhab- with remarks on Red Sea species). It should be dose character of microscleres evident in earliest noted that diactinal megascleres also occur in ontogentic stages. Megascleres styles or sub- Didiscus and Podospongia, but together with tylostyles, often polytylote; not differentiated mor- monactines and transitional types (styloids). phologically in parts of the body. Shape of sponge encrusting to massive, then commonly irregular; Latrunculia (Latrunculia) conulosa Hallmann not stalked with globular head (as in Poclospongia). Plate figure 3 7, plate 22 figures 2, 3, Remarks. Other genus-group aggregates of the text-figure 22 Latrunculiidae recorded from southern Australia are Sigmosceptrella and Negombata. Sigmoscep- Latrunculia conulosa Hallmann, 1912: 126,' text-fig. trella pi. Dendy (1921: 122; see also Dendy, 1922: 136) 24, 22 fig. 1 (off Devonport, Tasmania). is a good genus and has as type species (subsequent Material examined. Station KG 6, two specimens (F51952, designation by Dendy, 1922: 138) Spirastre/la F5 1 953); station KG 7, one specimen (F5 . fibrosa Dendy 1 954) Holotype (1897), from Port Phillip Heads. A of Latrunculia conulosa (AM). a b n n p\ ^

^ ^

Figure 22. Latrunculia conulosa, spicules from F5 1 952. a. Range of megascleres, x 85. b. Enlarged ends of nLascleres X329. c. Large samdasters with range of lengths on right, x980. d. Small choanosomal sanidas.er, x980 SPONGES FROM BASS STRAIT 41

Diagnosis. Irregularly lobose, compressible, elas- larger umbels, with bases deep in choanosome, in tic. Surface mostly conulose, vermilion on top in shape of trees or candelabra, with outermost life, dull orange below. Few clustered oscules. branches often curved, some recurved like festoons. Cortex indistinct, 65-435 ftm thick. Sanidasters Central trunks of umbels up to 500 urn thick; away with large spines in 4 whorls (inner ones more from cortex, some break up again inwards into widely spaced), packed to crowded through most curved branches, mangrove-like roots, or festoon- of cortex, scattered with smaller ones in choano- like anastomoses between neighbouring trunks. In some. Styles in compact thick columns, branching choanosome between spicular columns, irregularly in umbels and like candelabra below and within scattered, disoriented megascleres and sanidasters cortex. Few scattered styles. Additional lax skele- of various sizes. ton of light collagenous fibres. Additional lax skeleton of slightly translucent collagenous fibres of irregular cross-section, mostly Description. Mostly irregularly lobose. In F51952 crooked, twisted, lined by crowded microscleres. and F51953 (the latter for two specimens contigu- Width 0.3-0.5 mm, branching irregularly, with ous in situ), lobes are 0.5-3 cm wide, twisted and sparse anastomoses, intervals 1-2 mm. Tapering angular, partly free, partly coalescent, separated slightly to surface forming axes of conules, pro- by lacunae or narrow grooves. F51954 (pi. 3 fig. truding occasionally. Many collagenous 7) more regular, single compressed lobe, 3 x 5.5 anastomoses in choanosome much finer, com- cm wide, expanding from narrow base of attach- monly below 50 nm wide, straight to bent, terete. ment (1.5 cm wide) to height 4 cm. Vermilion in Spiculation: (1) Styles, occasionally with blunt life (10 R6-8/12) on top (deeper on conules, lighter points, thinner ones with telescoped points; a few in depressions), grading to dull orange below. Some modified as strongyles, oxea, and anisostron- areas become brownish to greenish grey and dull gyloxea (styloids), 203-297-353 x 3.5-7.2-9.7 urn. purple when drying. Recessed areas and base pale Large sanidasters of cortex and choanosome, orange-beige. Moderately firm, compressible, (2) total length 37.9-4J.7-47.6 /tm, width with spines elastic. 20.3-24. 7-28.8 /mi, width of shaft 4.2-5.6-7.4 /un. In F51952 and F51953, surface irregularly conu- (3) Small sanidasters of choanosome (developmen- lose (conules frequently coalescent in short ridges) tal stages), 24.4-36.9 x 8.9-20 x 1.5-4.1 /un. in exposed areas; faintly verrucose to rugose in Typical, most frequent form of large sanidasters other areas, otherwise smooth. In F51954, top and between forms illustrated in Hallmann (1912: 127, three sides regularly conulose, conules 1 -2 mm wide text-fig. 24b, c). Four distinct, symmetrical whorls and high, 1-3 mm apart, one wide side weakly of spines: distal ones like crowns, with spines verrucose-rugose. Ostia regularly scattered in inclined at 45°; inner whorls with spines slightly depressions between conules, barely visible. Few inclined, their outside in one plane. Shaft between oscules, 0.5-2 mm wide, single or clustered in small inner whorls distinctly terete. Distance, at spine subpetaloid groups. Many conules have terete tips tips, between inner whorls twice that between each of variable length, being protruding ends of col- contiguous inner and outer whorl. Spines of inner lagenous fibres. whorls frequently, those of outer whorls occasion- Cortex indistinct, 65-435 /on thick, weakly ally, bifid. Frequent departures: whorls may be collagenous-fibrous matrix, commonly packed with asymmetrical with respect to size, placement and sanidasters, less frequent inward, leaving some inclination of spines; contiguous whorls may vir- inner portions free. Peripheral choanosome com- tually merge; in more massive forms, like in monly packed with segmented blue-green algae. Hallmann's text-figure 24b, centre of shaft no Two specimens (F51953 and F5I954) contain scat- longer terete. Even smallest sanidasters, with spines tered, radially fibrous, fuzzy balls in deeper hardly developed, always have straight shafts. choanosome, diameters 48-64 ^m; possibly another type of microsymbionts, analogous to that in Anco- Remarks. The closest relation is with Latrunculia rinasuina. F51952 contains embryos, orange in life. (Latrunculia) hallmann i, see below. Main skeleton of compact columns of Latrunculia (Latrunculia) hallmanni sp. nov. megascleres with variable diameters and little orien- surface. columns straight tation except below Some Plate 3 figures 8-10, plate 22 figures 4, 5, to surface, but most branch repeatedly from text-figures 23-29 peripheral choanosome outwards, forming complex within and irregular umbels. In final subdivision Hololype: F51955, Station KG 3 (Tasmania, East Cove, cortex, strands dissolve into parallel tracts of 2-3 Deal Island, Kent Group; VV of jetty of settlement, some spicules abreast, approaching palisades. Some distance from shore; outcrops and boulders with algal 42 F. WIHDI'NMAYKR

pa " r p,cal "epre!ii (s- r-sarss Sir " °° " °< ssstsss sssjs: forest to 10 m). Five 1-5 paratypes: 1956, station KG 3; Diagnosis. Shape variable, mostly F51957, station KG 4 (Tasmania, West Cove, Hrilh Island! irregularly massive, large. Rubbery, fairly Kent Group; boulders covered with algae, much kelp); compressible. Surface variably vermcose F51958, Station KG 5 (Garden Cove, N side of Deal to rugose, brain-like, or almost Island, Group; smooth. Irregular Kent boulders with algae); 15 1959 and depressions frequent on top and F51960 (selii/oparalypes). Station KG 8 (l.asl sides. Colour Cove, Deal variable, dull red-purple, red-orange- Island, Kent Group, hallway between jelly and North brown, and off-white, often in different patches Point; rocky level bottom). on same sponge. Oscules scattered or clustered SPONGES FROM BASS STRAIT 43

AJ >r *.** Wa# *T* *J\#

x4.8. Figure24. F51957, depression with stellate Figures 24 25 l.uirunculia haUmanni sp. nov. Details of surface, area of apical depression will, two types ol oscules. I lie area is clusters Of ostia. Figure 25. Holotype, marginal bordered above by the tabulate margin, below by a narrow fold.

outer crust in cortex, but numerous in choanosome. tending to segregation by 2 sizes. Cortex tough, col- styles thinner, branching mostly within lagenous, 0.6-2 mm thick, detachable, tends to Columns ol with branches more closely spaced than in shrink and crack after preservation. Sanidasters cortex, previous species. Scattered styles and sanidasters massive, more irregular than in L. (L.) comilosu, choanosome more numerous, reticulation of col- with outer and next-inner whorls often merging in lagenous fibres tighter than in /.. (/,.) conulosa. (approaching amphiasters), mostly confined to thin 44 7 I . WIEDENMAYCR

Description. Shape and size variable (text-fig. 23). Paratype F51956 (pi. 3 fig. 9) depressed-pear- More massive and larger than Latrunculia shaped, 6x9x5 cm; surface as type 2 on one conulosa. Three types of macroscopic surface con- broad side, type 3 on the other, type 1 on narrow formation: (1) finely and regularly verrucose to sides near base. rugose, more or less like in Latrunculia conulosa; Paratype F51957 (pi. 3 fig. 10) irregularly bul- convoluted, with (2) meandering and branching bous, expanding to inverted pear-shape from con- furrows, i.e. cerebroid, commonly also coarsely stricted base, which is 3 x 4 cm wide. Height 7.5 verrucose; (3) almost smooth or faintly and irregu- cm, upper width 5 to 6 cm. Sides furrowed by con- larly verrucose-rugose. The three types may be cave depressions, elongate upwards, with sharp associated with transitional zones and overlaps. rims and regular to sinuous outlines. Top has two Some specimens have a truncate lop, with one or shallow depressions containing one conspicuous more concave depressions bearing oscules. Concave osculc (3 and 4 mm) each. depressions may occur on sides. Paratype F51958 like a stump truncated obli- Holotype (pi. 3 fig. 8), growing on a steep rock quely, with top regularly hollow. Width 4 cm, face, a wide inverted cone, 8 high, cm 9.5 x 1 1.5 height 4-6 cm. One side has surface mostly of type cm wide at top. Sides convolute, with deep furrows. 2, other side and concave depression on top with Truncate top bears a central concavity, 1.5 cm surface type 1. Rim of apical depression and its deep, 4^6 cm wide, folded lengthwise, containing inside distinctly bristly. Five conspicuous oscules oscules of two sizes. Marginal area of top flat, on sides, one at bottom of pit on top. mostly smooth. Upper portion of freee sides, to a F51959 and F51960 cut from large specimens. level 3 cm below top, shows surface type 1 superim- The former originally cake-shaped below, with posed on type 2, which prevails deeper down. undulating, horizontal ledges at mid-height, with Verrucose-rugose pattern, with furrows spaced 3 volcano-shaped, slightly constricted upper portion, mm, perpendicular to rim of top. flattish smooth top containing central pit. Greatest

i«'""'~ ~ i

Figure 26. Latrunculia hallmanni sp. nov. F51956, sagittal section in reflected liebt y 8 )Ti su.e on top and ,s J0 ,ned by a thicker and a thinner collagenous fibre. A eoniea, '^S^SSSS^Si SPONGES FROM BASS STRAIT 45

rp^mMk.

iUi ? Vv, hVi /

^3&k

1 28

reflected light. F.gure 27. Sagittal section, x4.1, Figures 27, 28. Lalrunculia hallmanni sp. nov. F51957, sections in parallel to the base, below, shows longitudinal collagenous fibres with cortex on left The oblique view of the section Section parallel to base, from below, with cortex brought out by shrinkage of the interstitial choanosome. Figure 28. above, x5.5. 46 F. WIEDENMAYER

width 22 cm, height 20 cm. F51960 like a thick depressions. Some grooves with ostia form isolated angular club or menhir (megalith), 20 cm high, 6 stellate groups, particularly in lateral depressions cm wide at constricted base and top, 12 cm in bulg- of F51957 (text-fig. 24). ing middle. Cortex tough, collagenous, easily detachable, 0.6 Colour never uniform, basically three hues, shar- to 2 mm thick. Surface commonly microhispid with ply set off, or intergrading and mottled: (1) dull protruding megascleres. Collagenous matrix dis- red-purple, chiefly on prominent portions of sides tinctly fibrous, mostly without orientation. Single

with surface type 1; (2) red-orange-brown, darker terete chones disposed radially or obliquely, rarely in depressions, lighter on portions with surface paratangentially, lined with microscleres. One con-

types 2 and 3; (3) off-white with tinges of hues 1 ical cribriporal group seen in section (text-fig. 26). and 2, mostly confined to narrow area around base, Extensive narrow, slitlike subcortical crypts. Incur- but in F51958 in irregular patch on portions of one rent canals dendritic in choanosome. Excurrent side and of apical depression (text-fig. 23d). Hue canals straight, mostly radial, up to 4 mm wide, 1: 2.5 R5-6/6 (F51958), 5 R5/4 (F51959). Hue 2: surreunded by thinner extensions of cortical 10 R6-7/6-8 (F51959), 10 R~ 2.5 YR6-7/6 matrix. Lumina terete, except irregular constric- (F51955). Hue 3: N 9- 10 (YR, 1) (F51955). Fresh tions. Other excurrent canals compressed, twisted, choanosome beige to orange-brown, darker for the with curving folds on insides, like in human ear canal linings: 5 YR8/8 and 5 YR9/4, respectively, (text-fig. 27). Their collagenous walls frequently for F51959. separated from surrounding chaonosome by thin Rubbery, fairly compressible. Cortex on top voids, possibly accentuated by shrinkage in alcohol. tends to shrink and tear in alcohol (holotype). Little Skeletal structure much like in Lalrunculia (L.) consistence in size and distribution of oscules with conulosa, but collagenous fibres better developed, regard to sides, top, depressions and surface types. more closely spaced (text-figs 27, 28). No secon- They may be scattered or clustered. Oscules in holo- dary reticulation of finer fibres in choanosome. type virtually confined to apical depression, of two Tapering main fibres at surface of type 1 form axis categories: (I) regularly spaced, numerous, small; of conules and protrude. Around rim of F51958 (2) irregularly scattered, larger, often elongate and bristles are protruding fibres. twisted, 0.5-1 mm wide (text-fig. 25). Larger Differences in structure of spicular skeleton most oscules round, up to 4 mm wide, commonly single, pronounced in cortex and periphery. Microscleres some paired, confined to top (F51957), or more crowded in outer quarter to half of cortex in numerous on sides (F51958). Larger oscules with Latrunculia (L.) hallmanni, scarce further in, slightly raised lips on top of F51960. unlike in L. (L.) conulosa, where microscleres are Ostia inconspicuous, in very fine grooves mean- packed at surface and crowded below, throughout dering, anastomosing and branching, mostly in cortex. Repeated branching of spicular columns

a b n n

Figure 29. Latrunculia hallmanni sp. nov .spicules t mm F5 1958 a. Range of styles, x 85. b. Enlarged ends of styles, x329. c. Massive sanidaster, x98(). d. e. Small and intermediate sanidaster, x 980. SPONGES FROM BASS STRAIT 47

begins well below cortex in L. (L.) conulosa, with spines, distally often all, described as being branches in inner cortex still thick and irregularly recurved. spaced. In L. (L.) hallmanni, profuse branching There are several similarities, of both species of begins just below cortex, with cortical branches thin Lalrunculia here described, with the descriptions and regularly spaced, and final, palisade-like sub- of Sigmosceplrella fibrosa (Dendy, 1897: 254; division within zone of crowded microscleres. lnter- Dendy and Frederick, 1924: 507). However, the stitially scattered megasclercs and larger protorhabds (early stages of sanidasters, beginning microscleres in choanosome more numerous in L. as tylotes) in all preparations examined are always (£.) hallmanni. straight, and no single "sigmatose" one was found. Spiculation (holotype): (1) Styles to faint sub- Though bifid spines are common in the inner tylostyles, straight to slightly bent, terete or thick- whorls of sanidasters of Lalrunculia (L.) conulosa

est nearer to point, points short, conical to (rare in L. (L.) hallmanni), I could not detect the mucronate or blunt, 237-J//-404 x 3.5-6.5-9.1 regularly "trilobate" arrangement described for Sig- jim. (2) Massive sanidasters transitional to mosceplrella fibrosa, nor the quadrilobate one of amphiasters, in cortex and in cortical matrix lining S. quadrilobata. excurrent canals, collagenous fibres and columns Order Axinellida Levi, 1955 of megascleres in choanosome, total length 44.4-55. 7-60.6 nm; total width 28.8-54.4-42.6 ,mt; Axinellidae Ridley and Dendy, 1887 width of shaft 7.1-9.5-13.2 /un. (3) Small to Pseudaxinella Schmidt medium sanidasters, in choanosome and inner cortex, total length 15. 5-34. 7-44.7 /an; total width Pseudaxinella Schmidt, 1875: 120.-Thielc, 1903b: 378 10.1-77.5-25.6 nm; width of shaft 2.3-4.4-6.4 Mm. (type species redescribed). — Wiedenmayer, 1977a: 155 (revised diagnosis). Etymology. The name honours E.F. Hallmann, Diagnosis. Mostly massive, unbranched Axinelli- who, during his activity at the Australian Museum dae without axial or mesial condensation of skele- in Sydney, did much to promote the knowledge of ton. Spiculation as in Axinella. More or less parallel the Australian sponge fauna, particularly by revis- and plumose, anastomosing ascending columns of ing Lendenfeld's types. spicules may be so crowded that the structure Remarks. Besides the differences from Lalruncu- appears confused. (L.) lia conulosa in skeletal structure, mentioned Pseudaxinella decipiens sp. nov. above, Lalrunculia (L.) hallmanni is distinct in

Plate 3 figure 1 plate 22 figure colour, in a more collagenous, detachable cortex, 1 , 6, and in having larger, more massive sanidasters in text-figure 30 the cortex. While there is some overlap in size and shape of smaller sanidasters between the two spe- Holotype: F51961, Bass Strait, 17 km S ol'Warrnambool (38°32.0'S, 142°28.6'E); depth 52 m, coarse sand and cies, those in /.. (/..) hallmanni frequently have shells (sin BSS 187). their shaft spined or tuberculate, and spines at both ends rarely in distinct whorls. The large sanidasters Diagnosis. Small, ridged-crescent-shaped, very in L. (L.) hallmanni commonly look like a pair of firm; in life, buff with greenish grey areas. Surface oxyspherasters joined by a short, stocky shaft. smooth to verrucose, microhispid. Oscules scat-

Judging from its description, Axos spinipocidum tered, minute. Spicular skeleton of intergrading Carter (1879b: 286, pi. 25 figs 1-9; type species of Diacarnus), from Port Jackson, Sydney, is also a

Lalrunculia (Lalrunculia). Its shape is that of a waisted tall cup, and its surface is extraordinarily convoluted and laciniate. It agrees with both species above in having cortex, canal linings and fibrous skeleton of similar collagenous, fibrillate material. Its sanidasters are distinctive: long and C slender (68 x 3 /on), symmetrical, with four whorls r of small spines (2 median, 2 distal) about equally spaced along the terete shaft. The one figured resembles the young stage in L. (/..) hiannulata Figure 30. Pseudaxinella decipiens sp. nov., spicules, a. lopsent, 1892 (see Topsent, 1904, pi. 12 fig. 6b' Range of oxea and styles, X 67. b. Enlarged ends of oxea, right), but the whorls are irregular, with some x260. c. llcxnous strongyles, with range, x67. 48 F. W1EDENMAYER

oxea and styles, mostly confused, with rare long of such spicules in the Axinellida, as expressed by and flcxuous slrongyles. Vacelet (1969: 177) and Pansini (1983: 79). Vace- lei's discussion also included the definition and dis- Description. Asymmetrical wedge, bound by three tinction, from Axinella, of Phakellia. Phakellia was crescent-shaped areas separated by sharp edges. redefined and discussed, chiefly with regard to Base of attachment and narrower side flat, but with In do- Pacific species, by Bergquist (1970: 17, 18). angular folds and depressions. Other side, sepa- A further question that needs to be introduced in rated by sharp curved ridge along lop, regularly this context, pending a comprehensive revision of convex, smoother than opposite side. 3 X 4 cm the Axinellida, is the position and suitable diagnosis wide at base, 2 cm high. In life, buff with greenish of Pseudaxinella and "Teichaxine/la" (recte Axino- grey areas. Very firm, little compressible. sia) for which I have given revised diagnoses Surface on larger, convex side almost glabrous (Wiedenrnayer, 1977a: 154, 155). These definitions along base, verrucose in one corner. Areas near are admittedly based on species of the central other corner and apical edge rough, microhispid, Atlantic, and are probably too restrictive. If like opposite side. Oscules minute, indistinct, elon- allowance is made for accepting decipiens and con- gate, vermiculate. Sand grains scattered on surface. vexa in Pseudaxinella, with their confused skele- Choanosome dense, with few small meandering ton; and if the excurrent canals. diagnosis of "Teichaxinella" is slightly widened, so that Phakellia (see Kollun, Skeletal structure mostly confused, except tracts 1959: 200) and Axinosia Hallmann (1914b: and bundles of 2-4 spicules intersecting at variable 349) become synonyms, skeletal structure angles. Much light-yellow spongin, with small would no longer be viable as diagnostic criterion round meshes, 20-125 /*m wide, in interstices of in this group. Nor would relative proportions of spicular meshwork, occasionally traversed by single oxea and styles, including the extremes of their spicules. Spicular meshwork of intergrading styles mutual suppression (which would sink and oxea with some longer flexuous slrongyles Homaxinella). Lack of interlaced. a special axial (or mesial) skeleton, and external shape (thinly flabellate to Spiculation: (1) Oxea and styles, both infrequent, caliculate in Axinosia, massive to thickly flabellata connected by numerous strongyloxea and anisos- in Pseudaxinella, surface smooth to aculeate or hispid in trongyles (one end mucronate); points most fre- both) could alone be used. quently blunt or telescoped; all commonly curved or bent; Distinctions could be arbitrary with 278-JJ0-483 x 4,1-7.5-1 1.2 /mi. (2) Flex- some thickly flabellate species such as Phakellia uous strongyles, about 1-2% of all spicules (recte Pseudax- inella"!) 542-770 X 4.7-9.1 /mi. lumida Dendy (1897: 236).

Remarks. Comparison should be made with Reniochalina Lendenfeld Axinella auranliaca Lendenfeld (1888: 235, pi. 5 fig. Reniochalina Lendenfeld, 1; redescribed by Hallmann 1914c: 424, text- 1888: 82. - Hallmann, pi. 1914b: 346f (discussion, revision: genus fig, 19, 21 fig. 1). The spiculation is very simi- dubiurn). Axiamon Hallmann, 1914c: 440 (objective lar, but otherwise only the structure of the axial synonym). -de L.aubenfels, skeleton in 1936a: 130. A. auranliaca is comparable to that in Pseudaxinella decipiens. Diagnosis. Mostly frondose or anaslomosins- The ramose, external shape and the differentiation of the occasionally lobose Axinellidae without peripheral axial or skeleton in the branches of Lendenfeld's mesial condensation of skeleton. Spicula- species are characteristic tion as in of Axinella as generally Axinella. Surface thrown into fine accepted. Burton anastomosing (1928: 129) transferred A. auran- grooves and ridges, the latter tuberculate liaca to Bubaris on account of the flexuous stron- or (commonly) lacinulate. gyles. This cannot be upheld if the definitions of Remarks. Lendenfeld (1888: Buhahs by Dendy (1922: 82f.) described two 62) and Hechtel (1969: species in Reniochalina, R. stalagmitis, 25) are accepted. There is the type a better agreement of species (subsequent designation bv Hallmann, Pseudaxinella decipiens with Axinella (correctly 1914b: 347), and R. Pseudaxinella) convexa lamella. Thev are here svn- Hoshino (1981: 207, text- onymised. In contrast to Whiteleege (1902b: 277 fig. 1; pi. 1 fig. 1). The main spicules in the 283), who accepted the types, Japanese species are chiefly oxea, Hanmann (1914b: about twice the 3461 .) rejected the types of Lendenfeld's size of those in P. decipiens, and the species in flexuous diacts the Australian Museum (2 syntypes of are oxeote, also larger than in our species. R. slalae- mitiS, AM C',9004 and B5478, wet, and one of The presence of long flexuous diacts in Pseudax- the syntypes of R. lamella inella known to him) as such, confirms the doubt as to the dignostic value because of inconsistency with Lendenfeld's descrip- SPONGES FROM BASS STRAIT 49

tions. Admitting only the latter, he pronounced R. R. stalagmitis, here figured in pi. 4 figs 4-5) over- stalagmitis unrecognisable and R. lamella as pos- laps with that of Ptilocaulis. Pending a more sibly belonging to Axinosia Hallmann. The three thorough re-examination of Lendenfeld's specimens rejected types were redescribed, and those in the in London, Reniochalina is here retained as a valid Australian Museum illustrated (one for each spe- genus, having oxea in addition to styles, and lack- cies) by Hallmann (1914c: 441, text-fig. 23, pi. 18 ing axial and mesial condensation of the skeleton. figs 2, 3, the latter quoted in reverse in text and But the variability in spiculation noted by captions) under the name Axiamon folium gen. et Hallmann, and the rather confused skeletal struc- sp. nov., with no other types. The three type speci- ture in the following new species raise the same sus- mens in question, plus the second syntype of R. picion expressed above, concerning Axinosia and lamella (BMNH: 1925.12.1.18, unknown to Pseudaxinella, that presence or absence of certain

Whitelegge and Hallmann), all of which I re- types of megascleres and of axial/mesial conden- examined (see pi. 4 figs 2-3) are sufficiently docu- sation may be synapomorphies with little systematic mented as such by respective museum-register importance. entries and autograph labels. Despite the incon- Reniochalina sectilis sp. nov. sistencies in Lendenfeld's descriptions, it is clear that Lendenfeld based Reniochalina on these speci- Plate 4 figure 1, plate 23 figure 1, mens. Hallmann's action is indefensible (1CZN text-figure 31 Article 18a), and both Axiamon and A. folium are objective synonyms. Holotype: F51962, Bass Strait, 17 km S of Warrnambool D (38°32.0'S, 142 28.6'E, depth 52 m, Reniochalina was hitherto only known from the coarse sand and shells), stn BSS 187. specimens discussed above. It is closest to Axino- sia in skeletal structure, spiculation and general Diagnosis. Small, massive-lobose, rubbery, dull shape, but the laciniate surface pattern is distinc- orange in life. Surface with regular grooves and tive. In this respect, it is similar to Ptilocaulis (see tabulate ridges branching and anastomosing. Wiedenmayer, 1977a: 152). Morphological varia- Oscules inconspicuous, in grooves. Spicular skele- bility of specimens assigned to Reniochalina by ton variably dense, confused to subisodictyal, felted Lendenfeld (including two unnamed species of or densely umbellar in ectosome. Spongin encas- Hallmann, 1914b: 443, probably synonymous with ing bundles of styles only.

styles, X84. b. Main style, x328. c. Range Figure 31. Reniochalina sectilis sp. nov., spicules, a. Range of main styles, x 84. f, Dermal oxeon and style, of main oxea, x 84. d. Main oxeon, x 328. e. Range of dermal oxea and g. i. Choanosomal filiform diactines, x328. X328. h Choanosomal styles (with example of mean length), 87. 50 F. WIEDENMAYER

Description. Massive, with two basally coalescent but rare, ectosomal spicules; there is imperfect diverging lobes: one little individualised, 2 cm long, segregation, in measurements, of oxea and free end 5 mm high, other one tapering, twisted, styles/styloids (styles being rare); interstitially scat- 3.5 cm long (free end 2.5 cm long). Width at base tered spicules are lacking. The strong development 2.5 x 3 cm. In life dull orange, now, in alcohol, of spicular umbels in the ectosome of R. sectilis

dark orange-brown (5 YR3-4/2-4). Rubbery, is somewhat reminiscent of the Raspailiidae. But moderately compressible. Surface with narrow non-specialisation and absence of echinators in the ridges and grooves, both 1-2 mm wide, straight to main skeleton do not favour such an affinity. The crooked. Ridges tabulate, microhispid with pro- lack of surface-lacinules in R. sectilis is a further truding spicules; often merging, branching and distinction. In the smaller specimen figured by anastomosing, as for grooves. Grooves semiterete, Hallmann, and in some areas of the larger speci- smooth except frequent fine transverse striation of men on the same plate, the lacinules are short and collagenous nervures. Few small oxcules in grooves. distinctly aligned on ridges. This is particularly evi- Collagenous dermis of grooves occasionally want- dent in the second syntype (fragment) of Reniocha- ing, especially near base, with choanosome lina lamella (pi. 4 fig. 2a), where lacinules are unprotected. scarce. Main skeleton in two elements: (1) Irregular subisodictyal reticulation of bundles of styles and Rhaphoxya Hallmann styloids, frequently encased in clear spongin. Some Rhaphoxya Hallmann, 1917a: 641 (definition, discus- spongin fibres up to 185 /mi thick. (2) Oxea scat- sion), 674 (systematic position).- Bergquist, 1970: 18 tered interstitially. Main skeleton generally con- (discussion). fused, lax to locally dense. [Acanthellina] Carter, 1885e: 365 (nomen oblitum).- Ectosome contains felt of oxea, styles and Bergquist, 1970: 18 (in discussion, = A can the/la semu styloids, little different from those of choanosome, Carter, nee Schmidt). but smaller, thinner, often in overlapping umbels. Diagnosis. Massive to thickly flabellate, fleshy Spiculation: (1) Styles and styloids of main skele- Axinellidae with lax, not condensed spicular ton (anisostrongyles, anisostrongyloxea), mostly skeleton. Surface strongly conulose, curved to strongly bent, 388-425-487 x rugose to lacinulate. Choanosome cavernous. Spiculation as in 7.7-72.2-16.5 /mi. (2) Oxea of main skeleton, Axine/la, but frequently with straight or, more often, gently curved, 298-345-390 filiform spicules. Dermis may be reinforced by tangential X 4.7-5.5-7.6 /mi. (3) Dermal oxea, styles and megascleres. styloids, 250-306-367 x 4.7-5.5-7.5 /mi. (4) Long styles, rare, in choanosome, 762-974 x 9.1-11.5 Rhaphoxya caetiformis (Carter) /mi. (5) Straight to irregularly flexuous filiform diactines, rare in choanosome, around 190 X 1.2 Plate 4 figures 6-12, plate 5 figures 1-3, /mi. plate 23 figure 2, text-figure 32 Remarks. The spiculation is similar to that of Acanthella caetiformis Carter, 1885a: 114, pi. 4 fig. Reniochalina stalagmitis, as described by Hallmann (part? Port 6 Phillip Heads). -Carter, 1885e: 364 (old (1914b: 442, text-fig. 23). R. stalagmitis has special, record).

Figure 12. Rhaphoxya caetiformis, spicules, a. Range of styles and styloids, x84. b Enlargedc-margea * Range x styloids,stvloids x 328.*M c. ( 84) and example ( x 328) of fine oxea. SPONGES FROM BASS STRAIT 51

Rhaphoxya cactiformis. -hurton, 1934a: 565 (trans- Choanosome cavernous; wide, annulate excur- fer only). — Bergquist, 1970: 18, pi. 4A (discussion; lec- rent canals commonly parallel to plane of lamella totype designated (as "holotype"), re-examined and below depressions of surface, perpendicular in figured). prominent parts of larger specimens. Subdermal Acanthella hirciniopsis Carter, 1885e: 364 (Port Phil- apices of regularly spaced perpendicular incurrent lip Heads). -Burton, 1938: 20 (Tasmania, new record only). canals 150-700 fim wide, 0.7-1.5 mm apart, con- Acanthellina parviconulatu Carter, 1885e: 365 (Port spicuous in smaller specimens with wide radial Phillip Heads). sulci. Mesial portion of frond contains lax, random Acanthellina rugolineata Carter, 1885e: 365 (part? Port arrangement of spicules, some grouped in whispy Phillip Heads). columns (pi. 23 fig. 2). Interstitial and peripheral — Acanthella slipitata. Dendy, 1897: 237 (part: syno- choanosome collagenous, densely granular, light nyms of Carter, 1885, only, with type data; not Acan- brown in transmitted light, mostly devoid of spic- thella stipitata Carter, 1881a (unrecognisable, holotype ules except plumose to chaotic zones, 350-1000 /xm destroyed) nor Dendy's specimens ( = Phakellia dendyi wide, oblique to surface, ending in conules, some fide Bergquist, 1970: 18). piercing surface. Dermis around 175 fim thick, Material examined. Station BSS 187, two specimens fibrinous, clearer than choanosome, with scarce (F51963). Type specimens of Acanthella cactiformis, foreign spicules and debris; occasionally pierced by Acanthella hirciniopsis, Acanthellina parviconulatu, and groups of spicules. Acanthellina rugolineata (BMNH). Spiculation: (1) Styles and styloids (strongyles, strongyloxea, anisoxea, the latter mostly with tel- Diagnosis. Flabellate, commonly pedunculate. escoped ends; some styles with a mucronate, curved Rubbery; little compressible in peduncle, softer, or bent, occasionally flexuous, rarely straight apex); in elastic frond. Buff to flesh-coloured in life. Both 248-545-397 x 1.9-5.6-9.7 //m. (2) Fine fusiform faces of frond with radial folds sparsely branch- oxea and filiform diactines, often broken; possi- ing and anastomosing, bearing complex conules bly not a distinct category, 192-281 x 1.3-3.1 /urn. and lacinules, separated by equally wide, mostly With respect to Carter's synonyms smooth grooves. Oscules marginal. Dermis gla- Remarks. four their type specimens, most of which are illus- brous, diaphanous in grooves, roofing regularly and following variables in punctiform vestibules. Cartilagenous collagen of trated here, the external shape can be outlined: more or less symmetrical peduncle continues as mesial ribs in frilly folds, fan-shape with rounded outline can be maintained with lateral branchlets in axis of lacinules. Spicu- with growth to a medium size. Particularly in larger lar skeleton in frond lax, confused to vaguely columnar. specimens, the lamella may be asymmetrical, lacunose, with an irregularly sinuous margin. A

Description. Intermediate, in size and shape, peduncle is not always developed: in some speci- between complete syntype of Acanthella hirciniop- mens, the base of attachment occupies the whole lamella. peduncle also sis (pi. 4 fig. 12), and lectotype of A. cactiformis width of the The may be figured by Bergquist. Like in former, peduncle dis- covered by conules, though small ones. In smaller the conules be scattered at tinct, almost smooth, slightly compressed; conules specimens may random, low, mostly aligned radially; dermis diaphanous or, more often, are connected by ridges arranged over numerous, scattered small subdermal cavities. radially. These may be more widely spaced and or equal to the ridges in width, both show- As in lectotype, fan wide, thin, symmetrical, concave, cross-section. Oscules in smaller without marginal oscules. Buff-to flesh-coloured ing v-shaped specimens are mostly irregularly scattered, and seem to in life, paler in alcohol. Firmly rubbery to tough, favour positions on the lower part of the lamella, little compressible, in peduncle and mesial near the margin, along parts or all of the margin, branches; softly rubbery in conules and ridges. but rarely with consistence. They are up to 5 Soft, limp interstitially and in dermis. Dermis with mm wide, often elongate or sinuous. fine reticulum of collagenous nervures, radial In larger specimens, the lateral ridges grow out- around conules, transverse in furrows. Oslia regu- ward, thickening and coalescent in irregular larly scattered between nervures. Few sand grains. with conules proliferating in various Hard peduncle with dense collagenous matrix sur- fashion, directions. The result is a disorderly and complex rounding spicules, extending into mesial region of arrangement of frilly combs and tubercules and lamella, repeatedly branching to margin. Branches intervening deep lacunae of sinuous outlines. Many anastomosed. All collagenous elements end in per- oscules are hidden inside the lacunae, but others pendicular branchlets within conules and ridges at are visible in prominent parts. surface. 52 F. WIEDENMAYER

Rhaphuxya felina sp. nov. walls about half as wide. Labyrinthine pattern of walls with compound lacinules and conules at sur- Plate 5 figure 4, plate 23 figures 3, 4, face. Ostia scattered. Dermis with tangential spic- text-figures 33-35 ules scattered, matted or in vague bundles. Oscules elongate to sinous and vermiculate, on top of apical Holotype: F51964, Station KG I (Tasmania, West Cove, cones and on sides. Inhalant canals parallel in Erith Island, Kent Group; wreck of S.S. "Bullseye"). peripheral choanosome, perpendicular to oblique Diagnosis. Fist-or paw-shaped, firmly spongy, to surface; meandering and sinuous further in viscid. Dull orange to yellow in life. Surface choanosome thus thoroughly cavernous (text-fig. verrucose-rugose in life. After preservation, thin 34). Megascleres of choanosome in branching and dermis with tangential spicules largely collapses, obliquely anostomosing plumose columns, and surface then conulose to lacinulate and contorted- scattered in confusion interstitially, together with alveolate (deep pits formerly vestibules). Oscules filiform spicules. Distinction between thinner on apical cones. Choanosome thoroughly caver- branches, anastomoses and scattered megascleres nous. Spicular skeleton indistinctly reticulate- frequently vague. Distinct principal spicular plumose, with many megascleres scattered columns straight or vaguely sinuous, about 50 /zm interstitially. thick and 170 to 350 /*m apart. Arrangement at surface, in edges, lacinules and conules of partitions, Description. Like a mitten or feline paw: massive, denser, matted or penicillate, plush-like. erect, compressed, regularly expanding from Spiculation: (1) Oxea, straight, gently curved narrow base to 4-5 apical osculiferous cones. Base or bent, some slightly flexuous; hastate not attached; side to mid-height with imprint and to slightly fusiform, points often conical; rare remnant of algal stalk used for attachment. Height styles and styloids (anisoxea with one end blunt); 214-25J-308 x 11 cm, width 4x9 cm. In life dull orange to 2.2-5.0-8.2 jxm. (2) Filiform diactines, straight chrome-yellow (7.5 YR7/10 to 2.5 YR8/10). Firmly to slightly flexuous, 138-767-220 1-1.2 spongy, viscid. Surface regularly verrucose with X fim. dendritic and meandering sulci. Semidiaphanous Re/narks. Rhaphoxya felina is easily distinguisha- weak dermis now mostly collapsed, except base and ble from R. eactifonnis by surface characteristics, some areas around top. Now exposed subdermal by its delicate dermis, different skeletal structure, cavities circular to elongate-sinuous to vermiculate almost exclusive occurrence of regular oxea, and (text-fig. 1-2 wide, 33), mm separated by contorted a distinct category of filiform diactines. Rhaphoxya iM

figures 33, 34. Rhaphoxya felina sp nov., details in reflected light, x 4. Figure 33. Surface with irreeular subdermal spaces. Figure 34. Block sectioned from surface (above) showing the cavernous interior, with chiefly erect disnosi (ion ol canals below surface. ' v -

SPONGES FROM BASS STRAIT 53

Figure 35. Rhaphoxya felina sp. nov., range and examples of spicules, x327. a-d. Oxea. e. Filiform diactines.

pallida (Dendy, 1897) and R. typica Hallmann Spirophorella Lendenfeld, If 236 (probably as (1917a) have a more substantial dermis than R. nomen novum for Spirophore/). felina, and their choansome is denser, not as Diagnosis. The only genus in the family. Axinel- as in this species. spic- thoroughly cavernous The lida with spiroscleres and commonly microrhabds. ular reticulum reinforcing the dermis of R. typica Trachycladus laevispirulifer is coarse, visible without magnification. Carter skeletal structure of/?, pallida and R. The main Plate 5 figure 5, text-figure 36 typica resembles that of R. felina, but the principal columns are more sinuous, and the surface plush Trachycladus laevispirulifer Carter, 1879b: 343, pi. 28 figs 1-5 (South Australia). -Carter, 1885e: 357 (Port Phil- of the latter is wanting. The chief difference resides lip Heads). -Dendy, 1897: 245 (Port Phillip Heads). in the average and maximum sizes of oxea and fili- Dendy, 1921: 107 (discussion of spirula). — Dendy, 1924: form diactines, which in R. pallida and R. typica 378 (discussion). — Dendy and Frederick, 1924: 506 are about twice those in R. felina. (Abrolhos Islands, WA; discussion). Spirophora digitata Lendenfeld, 794 Traehycladidae Hallmann, 1917 1887a: (Port Jackson). Synonyms: Spirophorellinae Lendenfeld, 1889 (nomen Spirophorella digitata.- Lendenfeld, 1888: 236 (old oblitum); Rhaphidistiinae de Laubenfels, 1936a. record). Trachycladus digitatus. — Hallmann, 1914a: 268, 429 Trachycladus Carter (discussion). — Hallmann, 1916a: 453, 466, text-figs 3, 4;

pi. 22 figs 1, pi. 23 fig. pi. 26 fig. pi. fig. I Trachycladus Carter, 1879b: 343. -Hallmann, 1916a: 2, 1, 2, 27 (redescription from syntypes and hypotyes). — Levi and 453 (definition, discussion). -Topsent, 1928: 37.- Levi, 1984: 942, text-fig. pi. 1 fig. 6. Bergquist, 1970: 21 (definition of family), 22 (discussion). 8, Spirophora bacterium Lendenfeld, 1887a: 795 (Western [Spirophore/] Lendenfeld, 1887a: 794 (nee Milne- Port Bay, Vic). Edwards, 1836).

a r

] fb tfW L C

Figure 36. Trachycladus laevispirulifer, spicules, a, b. Range and example of megascleres, x326. c. Spirulae and sigmaspire, x970. d. Microrhabds, x 326. 54 F. WIEDENMAYLK

Trachycladus bacterium.— Kallmann, 1916a: 453-457, Surface smooth with some vermiculate subdermall 486 (discussion). canals. No oscules. Trachycladus scabrosus Hallmann, 1916a: 459, text- Spiculation of branches: (1) Oxea, strongyles. fig. pi. 21 fig. pi. fig. pi. fig. (off 1, 4, 23 9, 28 6 Port styloids and styles, indistinguishable in size, Jackson). 265-277-314 x 5-6.2-7.1 /*m, curved, bent or flex- Trachycladus fastigatus Hallmann, 1916a: 462, text- uous, rarely slraighl. Oxea most frequent, hastate fig. 2, pi. 21 fig. 1, pi. 23 fig. 10 (Great Australian Bight). to slightly fusiform, commonly with telescoped Trachycladus digitatus var. gracilis Hallmann, 1916a: points; styloids (strong> loxea, anisoxea with 472, text-fig. 5, pi. 22 fig. 3, pi. 23 fig. 2, pi. 27 fig. 2 une- (Port Jackson). qually mucronate points) moderately frequent;

Trachycladus digitatus var . clavatus Hallmann 1916a: styles 1-2% of all megascleres, strongyles about 474, text-fig. 6. pi. 22 fig. 4, pi. 23 fig. 3, pi. 25 fig. 2, equally scarce. (2) Spirulae, toxaspires and sig-

27 fig. 1 p. 3, pi. 28 fig. 5, pi. 29 fig. (Port Phillip Bay, maspires, mostly smooth, length 6.6-/0.0-14.6 /*m, including hypotypes of Dendy, 1897). width 5.1-5.9-7.2 /tm, thickness 0.8-1.5 Mm. C- Ttachycladus digitatus var. strongylatus Hallmann. shaped sigmaspires, especially little-curved ones, 1916a: 477, text-fig. 7, pi. 22 fig. 5, pi. 23 fig. 4, pi. 26 commonly thicker and finely tuberculate. figs 3 6, pi. 27 fig. 4 (Port Phillip Bay). (3) Trachycladus reteporosus Hallmann 1916a: 479 (defi- Microrhabds, smooth, occasionally faintly cen-

nition, discussion), 480, text-fig. 8, pi. 21 fig. 2, pi. 23 trotylote, 6.4-/2.4-15.3 x 0.7-2.6-3.8 ,/.m.

fig. 5, pi. 24 fig. 3, pi. 26 figs 1, 4, 7, pi. 27 fig. 5 (typical Remarks. Trachycladus laevispirulifer is here conform, Port Phillip Bay); p. 482, pi. 21 fig. 3, pi. 23 figs ceived as a highly polymorphic species. such it 6-8, pi. 24 figs 1, 2, pi. 25 fig. 1, pi. 28 figs 1-4, pi. 29 As fig. 2 (var. or varr., Port Phillip Heads, based on 10 hypo- is the most thoroughly described and illustrated of types of Dendy, 1897, and one specimen in AM). all Australian sponge species (see synonymy). The Trachycladus pustulosus Hallmann, 1916a: 486, text- new specimen is externally most similar to that fig. 9, pi. 21 fig. 5, pi. 26 figs 5, 8, pi. 27 fig. 6, pi. 39 described by Hallmann as Trachycladus pustulo- figs 7 (Port Phillip Bay). 6, sus. Dendy (1924: 378) and Dendy and Frederick Trachycladus stylifer Dendy , 1924: 377, pi. 12 fig. 7, (1924: 506) doubted the justification of Hallmann's pi. 15 figs 39-42 (Three Kings Island, New Zealand).— species and varieties and thought that all Australian Bergquist, 1970: 21, pis. 5a, 10c (E of North Cape and specimens Poor Knights Island, New Zealand). of Trachycladus should probably be included in T. laevispirulifer. Trachycladus stylifer Material examined. Station KG 7, one specimen (F51965). is based on megascleres being almost exclusively Type specimens of Trachycladus laevispirulifer, styles. Dendy (1924) noted only a few stongyles, Spirophora digitata, S. bacterium (BMNH). Type speci- but no oxea. Bergquist (1970) recorded a propor- mens of Trachycladus scabrosus, T. fastigatus; T. digita- tion of in tus varr. gracilis, clavatus, strongylatus; T, reteporosus, 2% oxea her material. In most of T. pustulosus (AM). Hypotypes of Trachycladus Hallmann's (1916a) descriptions, mention is made laevispirulifer. Carter, 1885e (BMNH). Dendv, 1897 of scarce styles, which, however, seem to be want- (NMV). ing in the specimens assigned by him to T. digitatus var. strongylatus and T. reteporosus. In T. Diagnosis. Prolifically branched, often bushy. digitatus (typical form), Hallmann estimated the Stalk and branches commonly terete, of variable proportion of styles as being between 3 and 7%. length. Branches sometimes nodose, rarely In T. digitatus var. strongylatus and 7". reteporo- anastomosed, often curved, commonly pointed, sus, true oxea are rare, while strongyles and stron- but very short ones may be blunt, club-shaped, gyloxea predominate. Trachycladus stylifer is laterally coalescent. Tough, flexible, elastic. Ver- therefore synonymised with f. laevispirulifer. milion to bright orange in life. Surface smooth to rugo-reticulate, rugo-vermiculate or verrucose, Raspailiidae Hentschel, 1923 occasionally hispid; lipostomous. Condensed axis Synonym: Euryponidae Topsent, 1928. of longitudinally packed oxea and/or styles with lateral thin, plumose columns oblique to surface. Remarks. In merging the Euryponidae with the Raspailiidae, Little spongin in stalk. Interstitial spiroscleres 1 have followed Bergquist's view (1970: crowded, packed at surface. Microrhabds less 26-32; 1978: 167). In the earlier instance, Bergquist abundant, often clustered. has pointed out various insecurities, points of contention, and the need for revision, par- Description. Small, thickly fan-shaped, with short ticularly within Raspailia. This report is not the stalk, expanded base, and short, repeatedly divided place to discuss this matter any further, particu- clavate branches, laterally coalescent, with ends larly since I am describing only one species in the flush, mostly few shorter ones jutting foreward family, in Clathriodendron, one of the less con- from one face. Firmly rubbery, vermilion in life. troversial genera. SPONGES FROM BASS STRAIT 55

Clathrindcndron Lendenfeld cored by styles/subtylostyles, echinated by acanthostyles. Clathriodendron Lendenfeld, 1888: 215.-Hentsehel, 11911: 383 (synonymised with Raspailia).— Hallmann, Description. From narrow pedicel, 2 cm long, a I 1912: 295 (discussion, retained). -de Laubenfels, 1936a: 1102 (definition in Raspailiidae). — Bergquist, 1970: 30 series of compressed, mostly flabellate branches (definition in Raspailiidae). diverge chiefly in one plane. These coalesce partly along edges, in fronds more or less deeply incised Diagnosis. Raspailiidae without axial or mesial con- radially, with radial folds of variable width and densation of skeleton, with strong development of depth. Two flabellate branches with parallel planes reticulate spongin; fleshy dermis without special partly coalescent along radial portion of their spicules. Outermost erect spicules of periphery pro- inside. Total height 9 cm, total width when out- i truding only from most prominent parts of surface. stretched 13 cm. Thickness of fronds around 1 cm. Clathriodendron cacticutis (Carter) Surface thrown into prominent thin ridges, chiefly radial, branching and anastomosing here and Plate 5 figures 6, 7, plate 23 figure 5, there, with uniform lacinules at equal intervals. text-figure 37 Lacinules perpendicular, more frequently oblique upwards. Radial ridges 0.5-1 mm wide, about 3 Dictyocylindrus cacticutis Carter, 1885e: 354 (Port Phil- mm apart. Length and spacing of lacinules 1-3 lip Heads Vie.). mm. Raspailia cacticutis. — Dendy, 1896: 48 (Port Phillip Heads). — Pick, 1905: 35 (excluded from Raspailia).— Dark brown (10 R3/2) in life, slightly faded in Shaw, 1927a: 427 (Maria Island, Tasmania). alcohol (2.5 YR4/2). Elastic, rubbery. Small oscules irregularly scattered, scarce, on one face Aulospongus? cacticutis. - Dcndy, 1905: 1 76 (tentative of fronds, in furrows between ridges. Conspicu- transfer only). ous fleshy, glabrous dermis. Tips of lacinules Clathriodendron cacticutis. —Hallmann, 1912: 297 microhispid with protruding spicules. In furrows (redescription of spiculation from type slide). between ridges, dermis reinformed by faint reticu- Material examined. Station KG 9, one specimen (F51966). lum of collagenous nervures, with transverse lines Holotype, BMNH 1886.12.15.120 (wet, from Port Phil- generally more prominent. Ostia within these lip Heads, ex J.B. Wilson collection). meshes barely visible. Diagnosis. Flabellate, with thin, often contorted Choanosome regularly and finely cavernous.

pedicel. Fronds around 1 cm thick, may be com- Mesial areas devoid of condensation, contain regu- plex, radially folded, more or less deeply incised lar reticulation of spongin fibres with small poly- from upper margin. Both faces traversed by gonal meshes. Styles and acanthostyles less sparsely branching and anastomosing radial ridges, numerous here than further out, with little relation to fibres in placement and orientation. Mesial fibres 1 mm wide, beset with numerous conules or lacinules up to 3 mm high. Intervening grooves 3 50-70 fim thick, with mesh sizes 135-175 /an. At mm wide, mostly terete, transversely striate, bear- periphery, almost continuous sheet or dense retic- ing few scattered, small oscules. Rubbery, dark ulum of spongin bears palisade of acanthostyles on

brown in life. Choanosome finely cavernous. Main outside. Inside ridges and lacinules, long primary skeleton reticulate, denser, richer in spicules at fibres at intervals of 175-260 /*m, cored by styles periphery, here with palisade of acanthostyles. and subtylostyles, echinated by acanthostyles. The Ridges and lacinules reinforced by longer fibres latter perpendicular or oblique outward. Dermis 65-700 nm thick, thinner on ridges. Outermost spicules not protruding except apices of lacinules. a r Spiculation: (1) Styles and subtylostyles, straight, r curved or bent, 395-5 /J-591 x 6.1-/2.J-16.5/tm; b « (2) acanthostyles with pointed to blunt ends, 90.9-/00-123.5 x 8.5-73-15 /mi.

Remarks. Clathriodendron arbuscula Lendenfeld (1888, pi. 5 fig. 2, misidentified in the caption, fide Hallmann, 1912: 147, 297; 1914a: 267; redescribed by Hallmann, 1912: 296f.) is a distinct species, with proliferous thin branches, thinner conules, Figure 37. Clathriodendron cacticutis. spicules, a, b. in C. cacticutis, and with a Range and example of styles, x67. c, d. Enlarged ends tylostyles larger than of style and subtylostyle, x261. e. Acanthosiyles, x261 complement of oxea. -

56 I . WIEDENMAYER

Clathriodendron ruhrum Kirk (1911), (revision, monotypic restriction). -Topsent, 1928: 62.- Topscnt, 1930a: 24, 46 (2 Australian species added). redescribed by Bergquist ( 1 970: 3 1 ) has compressed- (synonymised with Clalhria). digitate branches, a surface without conules, styles Van Soest, 1984: 7, 129 Aiuhcrochalina Lendenfeld, 1887a: 741, 786 (part, for and tylostyles of different sizes (the former very exceptions sec Echinoclathria leporina. below). — Burton, large) and oxea about twice the size of those in C. 1934a: 588 (revision).- Wiedenmayer, 1977a: 140 arhuscula. (references), hololype of cacticutis, here figured for The C. Clathriopsamma Lendenfeld, 1888: 227. -Hallmann, comparison (pi. 5 fig. 7), is much smaller than the 1920: 771 (definition, discussion, review). -Vacelel, Vas- present specimen, obviously immature, with con- seur and Levi, 1976: 75 (discussion, Aulenella syn- ules little developed. onymised). -Van Soest, 1984: 129 (synonymised with Clalhria). Subclass Ceractinomorpha Levi, 1953 Dlciyociona Topsent, 1913a: 61 8. -Levi, 1960: 60 (synonymised with Clalhria). — Van Soest, 1984: 7, 129. Order Poecilosclerida Topsent, 1928 IDendrocia Hallmann, 1920: 767. -Van Soest, 1984: Microcionidae Carter, 1875 129 (synonymised with Clalhria).

Leploclalhria Topsent, 1928: 61 , 62, 298 (part, fide van Synonym: Clathriidac Lendenfeld, 18X4. Soest; type species vacant). — Levi, 1960: 54 (synonymised wilh Microciona}.— VIM Soest, 1984: 129. Remarks. I have already established the priority of Pseudanchinoe Burton, 1929a: 433 (monotypic). — de Carter's name (Wiedenmayer, 1977a: 139, with Laubenfels, 1936a: 109 (part). - Van Soest, 1984: 129. resume on usage of both names), noting that the Ilymanlho Burton, 1930a: 502. -Van Soest, 1984: 90, priority of Clalhria, if accepted as a synonym of 130. the priority Microciona, has no bearing on of the Aulenella Burton and Rao, 1932: 345. family name (1CZN, Article 40). Thalyseurypon de Laubenfels. 1936a: 107. — Levi (1960: 50; 1973: 513) regarded the two Wiedenmayer, 1977a: 143, 144f. (as synonym of Pan- family names (but not the respective type genera) daros). -Van Soest, 1984: 108, 128 (synonymised wilh as synonyms and chose Clathriidac (wrongly Clalhria). attributed to Hentschel, 1923), as valid, without Cionanchora de Laubenfels, 1936a; 108. — Van Soest, giving reasons. Clathriidae sensu Levi was used by 1984: 7. Quizciona de Laubenfels, 1936a: 111. — Van Soest, most authors after I960, but was never justified, 1984: 130. with correct reference to the rules of nomenclature. Wetmoreus de Laubenfels, 1936a: 1 12. — Van Soest, Microcionidae was also used by Hechtel (1965: 41) 1984: 130. and Hoshino (1981: 153), both in the sense of de Laubenfels. Van Soest (1984: 89) treated Clalhria Diagnosis. Encrusting to erect and bushy Micro- and Microciona as congeneric, but as distinct sub- cionidae with simple, dendritic or anastomosing genera, and in this context treated Clalhria as senior spongin fibres cored by (acantho-)styles and echi- generic synonym, which is here accepted as correct. nated by distinct (acantho-)styles. Other (single) However, his choice of Clathriidae Lendenfeld, categories of special styles or tylostyles at surface, 1884 (wrongly attributed to Hentschel, 1923) as arranged paratangentially or piercing ectosome. Yet valid, in preference to Microcionidae Carter, 1875, another category of styles or tylostyles may occur breaches ICZN Article 40a and has to be rejected. interslitially in choanosome, but may intergrade Clalhria Schmidt wilh ectosomal megascleres.

Clalhria Schmidt, 1862: 57. - Ridley and Dendy, 1887: 1912: Remarks. The above I46.-Uendy, 1905: 170. -Kallmann, " 205.- synonymy follows chiefly that Topscut, 1928: 62. -Burton, 1934a: 558 (excessive syn- suggested and implied by van Soest (1984), who onymy, discussion). — Levi, I960: 50 (resume, defini- based his revision of the Poecilosclerida, and par- tion).- Bergquist, 1965: 168 (discussion). — Wiedenmayer, ticularly of the Microcionidae, on taxonomic prin- 1977a: 140 (references, resume). -Van Soest, 1984: 7, 90 ciples expressed in Table 1 (p. 7) and in the (synonymy, definition, discussion). discussions of families and genera. In essence, he Microciona Bowerbank, 1863c: 1 109. - Levi, 1960: 51 emphasises skeletal structure, and degrades habi- (resumtS definition). - Bergquist, 1965: 168 (discus- tus and spicular characters (chela-types (particu- sion).- Wiedenmayer, 1977a: 140 (references, larly discussed on resume). — Van Soest, 1984: 7, 90 (discussion, subgenus p. 84), absence of spicule of Clalhria). categories (p. 108), presence or absence of acan- Ophlitaspongia Bowerbank, 1866: 14.— Simpson, those condition of megascleres, growth form). 1968a: 95 (synonymised with Microciona). Simpson (1968a: 119, cited in Wiedenmayer, 1977a: Witsonella Carter, 1885c: 320. -Hallmann, 1912: 237 141) had devalued similar characteristics used as (definition, excessive revision). — Hallmann, 1920: 768 diagnostic in the past. Regarding the inclusion of SPONGES FROM BASS STRAIT 57

Ophlitaspongia, see Remarks on Echinoclathria, Thalysius transiens. -deLaubenfels, 1936a: 105 (trans- below. fer only).

Clathria transiens Hallmann Material examined. Station KG 7, one specimen (F51980). Type specimens of Clathria transiens (AM). Plate 5 figure 8, plate 23 figure 6, Diagnosis. Erect, much branched, irregular, often text-figure 38 bushy. Branches variable in length, rather thin, some anastomosing, mostly complex nodular and Clathria transiens Hallmann, 1912: 266; with four lacunose, frilly, with many lateral branchlets as unnamed forms; form a ( = typical form), p. 229, text- long as thickness of figs. 47, 47a, pi. 33 fig. 1 (off Devonport, Tas.); form main branches. Firm, elastic. b, p. 231, text-figs. 48, 48a, pi. 33 fig. 2 (40 mi. W of Bright vermilion in life. Oscules inconspicuous. Kingston, SA); form c, p. 232, pi. 33 fig. 3 (same local- Spiculation (megascleres) variable, toxa often ity); form d, p. 233, pi. 34 fig. 2 (Port Phillip Bay, scarce, chelae may be very rare. Vic). -Burton, 1934a: 559 (discussion), 599 (Great Bar- rier Reef, record only). —Shaw, 1927a: 426 (Maria Island, Description. Short stalk. Branches roughly in same Tas., record only). plane, asymmetrical. Secondary branches short. Contorted rugae and nodules 0.5-2.8 mm wide, abed interstitial lacunae 0.4-1.6 mm wide. Main n branches chiefly 5-6 mm thick, distally blunt. Spiculation: (1) Choanosomal coring styles, rarely straight, commonly gently curved or bent, 126-/49-180 x 2.9-7.5-10.9 /an. (2) Peripheral styles, coring and projecting through ectosome, 220-469 x 7.6-8.5 ftm. (3) Interstitially scattered choanosomal styles, strongyles, tylostrongyles and tylostyles, mostly straight, some gently curved or bent, 156-254-327 x 2-5-3.9 /mi. (4) Smooth or slightly roughened echinating styles and subtylostyles, 56-64-72 x 2.1-2.7-3.6 /an. (5) Toxa with straight or recurved arms, scarce, 96-/45-157 x 0.9-/. 8-2. 5 /an. (6) Palmate chela (only one seen), 12.8 x 4.7 /an.

Remarks. The new specimen agrees in growth form with Hallmann's form c. Its colour in life was vermilion (5R5-6/12). The spiculation does not closely agree with any of Hallman's forms and attests to the variability of this species.

Clathria wilsoni sp. nov.

Plate 5 figure 9, plate 24 figure 1,

text-figure 39

Hoiotype: F51967, Station KG 5 (Garden Cove, N side of Deal Island, Kent Group, Tasmania; boulders with algae, much kelp).

Diagnosis. Small, with many crowded, short, nodu-

lar, angular, blunt branches from flat common base. Dull vermilion in life, spongy. Surface verrucose to hispid. Oscules small, sinuous, elongate, scattered. Reticulate skeleton of angular spongin fibres, typical megascleres. Microscleres rhaphidotoxa and rare atrophied chelae.

Figure 38. Clathria transiens, spicules, x263. a. Description. Small; thickly encrusting base with Choanosomal coring slylc. b. Peripheral coring style, c. irregular sinuous outline, 1 x 2 cm wide, 5 mm Interstitial choanosomal style, d. Echinating style, e. Toxa. high, topped by 10-12 clustering, partly coalescent, 58 F. WIEDENMAYER

(3) Echinating acanthostyles, 31.6-57.2-67.2 x 2,6-6.8-10 ixm (width with spines). (4) Thin strongyles and rhaphidotoxa, with ends occasionally roughened, 71-137 x 1-1.9 ftm. (5) One weak, atrophied chela (palmate?), 6.4 ^m chord.

Etymology. The species is dedicated to Dr Barry Wilson, formerly Director of the Division of Natural History and Anthropology, Museum of Victoria, whose initiative made this study possible.

Remarks. Hallmann (1912: 215) segregated several species, mostly Australian, as a distinct group within Clathria, which he called the "spicata- group". As characteristics of the group, he noted (1) imperfect differentiation of principal (coring) and accessory (echinating) megascleres, (2) participation of the former in the echination of the fibres,

(3) semi-plumose or spicate arrangement of coring megascleres in the main fibres of the peripheral main skeleton, (4) absence of coring spicules from the connecting fibres, and (5) relatively dense echination on transverse fibres below the surface, chiefly on their outer side. Clathria wilsoni shows some Figure 39. Clathria wilsoni sp. nov., spicules, a. traits outlined above, but the absence of others Choanosomal coring styles, x263. b. Interstitial and shows that the "spicata-group" is really an artifi- ectosomal styles, x263. c. Rhaphidotoxa and thin stron- cial one. In skeletal structure and spiculation, gyles, x 263. d. Range of echinating acanthostyles, x263. Clathria wilsoni shows most affinity with Clathria e. Enlarged acanthostyle, x788. costifera Hallmann (1912: 215), particularly in the stubby branches, 3-5 mm high, 2-3 mm wide. In ectosomal skeleton without special spicules, life dull vermilion, yellowish around base. Con- presence of rhaphidotoxa, and lack of chelae in C. sistency spongy. costifera. The external shape of the latter, however, Surface finely verrucose to hispid, the latter chie- is distinctive. Rhaphidotoxa also occur in Thaly- fly on branch-tips. Oscules small, commonly elon- sias juniperina (Lamarck, 1814). gate, sinuous to vermiculate, scattered on base and sides of branches. Echiaoclathria Carter skeleton Choanosomal of spongin fibres, 18-35 Echinoclathria Carter, 1885e: 355. -Hallmann, 1912: /im thick, of angular disposition, with little orien- 275f. (discussion, revision). — Wiedenmayer, 1977a: !43f. tation; irregular meshes of variable size, smaller (references, discussion; not Pandaros). ones round, larger ones rhombic to polygonal. Holopsamma Carter, 1885b: 211 (part). -nee sensu Fibres always cored by styles and subtylostyles, one Loeblich and Tappan, 1964: C792 (under to eight abreast, echinated by short acanthostyles. Xenophyophorida, wrong type species). [Halme] Lendenfeld, 1885f: 285 Interstitially scattered thinner styles and sub- (nee Halme Pascoe. 1869). tylostyles, filiform diactines and rhaphidotoxa. Aulena sensu Lendenfeld, 1888: 228. -Lendenfeld, Chelae extremely rare, atrophied. Interestitial styles 1889b: 90 (nee Aulena Lendenfeld, 1885f.). form special dermal skeleton, in paratangential Ophlitaspongia sensu auctorum (nee Bowerbank, 1866): bundles and wide, regularly radiating penicils. Hallmann, 1912: 253 (discussion, revision). -Burton, Paratangential fibres just below surface commonly 1959: 246 (key, list of" 5 Australian species). - have most echinators on outside. Ascending fibres Wiedenmayer, 1977a: 140 (references). below surface occasionally with plumose arrange- Litaspongia de Laubenfels, 1954a: 162 (type species: Ophlitaspongia ment of coring styles. arbusculu Row, 1911). Spiculation: Choanosomal coring styles (1) and Diagnosis. Microcionidae of variable shape, often subtylostyles, commonly slightly curved or bent, thinly frondose or honeycombed, with generally mostly smooth, some roughened to faintly spinu- reduced spiculation. Principal megascleres smooth late, 1 77-257-287 X 2.4-4.5-7.4 /tm. Interstitial (2) styles or subtylostyles, coring or irregularly plumose and ectosomal styles, more frequently straight than to semi-echinating; auxiliary (interstitial) gently curved or bent, 83-777-264 x 1-2.2-3.3 jun. megascleres, if present, often filiform (strongyles SPONGES FROM BASS STRAIT 59 or subtylostrongyles). Microscleres, if present, toxa Echinoclathria carteri Ridley and Dendy and/or isochelae. Reticulate spongin fibres well Echinoclathria carteri Ridley and Dendy, 1886: 476. — developed, but may be obscured in some Ridley and Dendy, 1887: 162, pi. 29 fig. 12, pi. 31 fig. thoroughly sandy species, where proper spicules 3 (Moncoeur Island, Bass Strait; Twofold Bay and off may be rare or lacking. Port Jackson, NSW). -Hallmann, 1912: 284, text-fig. 65. Echinoclathria macropora. — Whitlegge, 1907: 504 (off Bolany Bay, off Coogee, off Wollongong, Shoalhaven Remarks. Simpson (1968a: 95, 105) merged Oph- Bight, NSW; nee Plectispa macropora Lendenfeld, 1888; litaspongia in Microciona, on the basis of a re- nee Echinoclathria macropora: Whitelegge, 1901). examination of topotypes of the type species, Oph- Axociella carteri.— de Laubenfels, 1936a: ^(transfer only). litaspongia papilla Bowerbank (junior synonym of Spongia seriala Grant). This view is upheld here, Material examined. Station KG 9 (record and colour-slide but a new genus for Ophlilaspongia sensu aucto- only). rum is not necessary. Hallmann, in his discussion Diagnosis. Long branches mostly simple, terete, of Echinoclathria, admits that merging this genus around 1 cm thick, straight to bent or crooked, and Aulena with Ophlitaspongia is equally feasible rarely concrescent. Surface regulalry alveolate, with as keeping the latter distinct: "Echinoclathria is ulti- tabulate partitions around 1 mm, deep, rounded- mately separable from Ophlilaspongia only by polygonal pits 2-3 mm wide. Rubbery, dull orange- virtue of its characteristic honeycomb-like struc- yellow when fresh, turning purplish brown. Sty- ture". The definition of Ophlitaspongia by Levi lote principals, 132 X 9 //m; subtylostylote aux- (1960: 58, 64) emphasises spongin fibres containg iliaries, 160 x 2 jtm; palmate isochelae 15 /ira long. quasi-echinating magascleres, without distinction of principal and accessory types, and absence of Remarks. The single specimen was accidentally lost palmate isochelae. This obviously conforms largely soon after collecting. But the field notes, colour with the diagnosis of Ophlitaspongia seriata. This slide, published descriptions, and plate 31, figure 3 in little virtually monotypic diagnosis is even more appar- Ridley and Dendy, 1887, leave doubt as ent though only implied, in van Soest (1984: 7, to the proper identification of this specimen. The after collecting noted as dull bottom), where incrusting habit is linked with Oph- colour soon was litaspongia. In contrast, Burton (1959: 246), like orange-yellow (7.5 YR7-8/6-8) with some areas Hallmann, included, in Ophlitaspongia, species becoming purplish brown after a while. The con- without microscleres, species with isochelae and sistency was rubbery. For comparison, see Remarks toxa, and species with toxa only. Species of on following species. Echinoclathria revised by Hallmann are mostly devoid of microscleres, but two contain isochelae. Echinoclathria favus Carter Given the variability in shape within Oph- litaspongia sensu Hallmann and Burton, there Plate 5 figure 10, plate 24 figure 2, seems to be little value in the "honeycomb-like mass text-figure 40 of anastomosing flattened trabeculae" (Hallmann's diagnosis of Echinoclathria) as a diagnostic trait. Spongia cellulosa. — Lamarck, 1814: 373 (nee Spongia A similar structure occurs in Pandaros (type spe- cellulosa Esper, 1797; King Island, Bass Strait). Echinoclathria Carter, 292 cies), but in that genus, as redefined by van Soest favus 1885d: (South coast). -Ridley and Dendy, 1887: 160, pi. 31, figs 4, 5 (1984: 127), spongin fibres may also be woven into (Moncoeur Island, Bass Strait). -Dendy, 1896: 40 (old continuous sheets. The latter structure also charac- records). -Hallmann 1912: 276, text-fig. 61 (off Devon- terises Echinoclathria tenuis Carter (junior syno- port, Tasmania).— Topsent, 1930a: 20, pi. 1, fig. 3 the type species nym of Spongia leporina Lamarck), (redescription of Spongia cellulosa: Lamarck). of Echinoclathria subsequently designated by Axociella favus. —de Laubenfels, 1954a: 164 (transfer Burton (1934a: 562, where the genus is merged in only). Clathria). Contradicting himself, and the rules of Material examined. Station WB, one specimen (F51972). (1934a: 599) transferred E. nomenclature, Burton Type specimens of Echinoclathria favus (BMNH). tenuis to Ophlitaspongia, maintaining this combi- Diagnosis. Massive-lobate, small to medium-sized, nation later (Burton, 1959: 246), in accordance with commonly on Pecten or other large shells. Softly Dendy (1896: 37), Hallmann (1912: 261) and Top- spongy to rubbery, light red to light orange in life. sent (1932a: 101). But his earlier subsequent desig- plenary Surface contorted-honeycombed, with alveoles nation is of course irreversible, except by commonly 2-3 mm wide, partitions 1 mm and power of the International Commission on Zoo- below. Some consecutive partitions lacking here logical Nomenclature (ICZN Article 79). 60 F. WIEDENMAYER

minal constrictions and terminal swellings elongate and indistinct, 132-746-163 X 1.4-/.6-2.1 /an.

Remarks. This Echinoclathria is distinct from other honeycombed species without sand inhabiting the same area (£". carteri Ridley and Dendy, E. glabra Ridley and Dendy) by its light red-orange colour; from the former by its massive habit, from both by the tendency of some neighbouring pits to merge in meandering trenches, and its common associa- G= tion with large shells, especially Pecten. Megascleres are generally larger in the other two species. The Figure 40. Echinoclathria favus, spicules, x259. auxiliaries of E. glabra were originally described a. Range and examples of principal megascleres. b. Raage and example of auxiliary megascleres. as "smooth bicapitate cylindricals" (presumbably subtylotes). and there, pits then elongate, meandering. Prin- Echinoclathria laminaefavosa (Carter) cipal megascleres mostly in 2 sizes. Plate 5 figure 11, text-figure 41 Description. Contorted-lobate, dimensions 6x7 X 10 cm. Honeycombed surface comparable to Holopsamma laminaefavosa Carter, 1885b: 212 (Port both specimens figured in Ridley and Dendy, 1887. Phillip Heads, Victoria). Cellular spaces more regular, with slightly thinner Haime nidus vesparum Lendenfeld, 1885f: 288, pi. 26 partitions; characteristic lack of anastomoses and figs 1, 2; pi. 27 figs 4, 5, 7; pi. 28 figs 8, 9, 11; pi. 29 resulting meandering lacunae infrequent. Common figs 12, 13 (Port Phillip Bay, Vic.; Port Jackson, NSW). -Lendenfeld, 1888: 157 (Port Stephens, association with Pecten not noted. In life light red NSW). -Lendenfeld, 1889b: 457, pi. 11 figs 1, 7, 9, 10, (7.5 YR7/10-12) above, light orange (2.5 YR8/8) 15-18. -Whitelegge, 1889; 184 (Maroubra Bay, NSW) below. Softly spongy to rubbery. Skeleton a mesh- lEchinoclathriafavus van arenifera Carter, 1885c: 350 work of clear to faintly stratified spongin fibres, (south coast?). with principal fibres in plane of lamellae. Principal lEchinoclathria arenifera. — Dendy, 1896: 40 (Port fibres terete, straight to crooked, sparely branched, Phillip Heads, Queenscliff jetty, Vic). -Shaw, 1927a: 426 35-60 fjm in diameter. Transverse (connecting) (Maria Island, Tas.). fibres frequently oblique and bent, around 20 ^m Hatme laxa Lendenfeld, 1886a: 845 (collective name thick. Meshes small, round (width in the order of for 2 varieties, no nominotypical subspecies). Halme laxa var. minima fibre-diameters) to irregularly polygonal, up to 100 Lendenfeld, 1886a: 847 (Port Jackson). (im wide. Principal megascleres indistinctly Halme laxa var. digitata Lendenfeld, 1886a: 847 (Port segregated in two categories. Distribution of larger Jackson). ones inconsistent, rare in some portions of Autena too. -Lendenfeld, 1889b: 95 (collective meshwork. name). -Whitelegge, 1889: 187 (off Green Point and off Where frequent, larger styles are both/either Ball's Head, Port Jackson). -Dragnewitsch, 1906: 442 coring, 1-5 abreast, with angular course, and/or (Pulu Brani, Singapore). -Hallmann, 1912: 287 (2 syn- oblique, in plumose to semi-echinating arrange- types re-examined). -Hallmann 1914a: 268 (partial syn- ment, the whole irregular. Outermost transverse onymy). -Burton, 1938: 20 (Maria Island, Tas.). fibres mostly free, tapering to surface, often echi- Aulena laxa var. minima.- Lendenfeld, 1888: 228.- Lendenfeld, 1889b: nated at tips with protruding styles. Thinner prin- 96, pi. 8 figs 10, 11, 21. Aulena laxa var. digitata. — Lendenfeld, 1888: 229.— cipal megascleres everywhere, in fibres and Lendenfeld, 1889b: 96, pi. 8 figs 12, 15, 20; pi. 9 fig. 1. interstitially. Auxiliary megascleres strongyles, Halme gigantea Lendenfeld, 1886a: 848 (collective instertitial only. Chelae (described and figured by name for 3 varieties). Ridley and Dendy, and Hallman) not found. Halme gigantea var. micropora Lendenfeld, 1886a: 849 Spiculation: Principal (1) megascleres, larger (nee Halme micropora Lendenfeld, 1885f; Illawarra, ones as fusiform subtylostyles, slightly constricted NSW). below head, thinner ones with narrow heads, Halme gigantea var, intermedia Lendenfeld, 1886a: 849 approaching anisoxea, overall dimensions 69-55-99 (Broughton Island, NSW). Halme gigantea x 2-5.5-4.8 fim. Disregarding rare transitional var. macropora Lendenfeld, 1886a: 850 (east coast). types, two categories: 80-55-99 x 3.3-5.5-4.8 /an, Aulena gigantea.- Lendenfeld, 1889b: 97 69-75-79 x 2-2.5-3.1 /an, (collective and respectively. (2) name). -Whitelegge, 1889: 187 (off Green Point, Port as strongyles Auxiliary megascleres, and subtylos- Jackson). -Hallmann, 1914a: 268 (partial synonymv).- trongyles, the latter faintly fusiform, with subter- Burton 1938: 20 (Maria Island, Tas.). SPONGES FROM BASS STRAIT 61

gin, foreign debris and proper spicules; presence or absence of orientation of fibres and of their differentiation into primaries and secondaries, specialisation in the periphery and at the surface. Where differentiation and orientation of fibres prevails in the main skeleton, debris may be agglutinated in columns, or may be encased in knotty fibres. Where specialisation below and at the surface is developed, principal specules (styles) seem to be frequent. They echinate the outermost Figure 41. Echinoclathria laminaefavosa, spicules from paratangenlial fibres on the outside and may

1 395442. 51969, x a. Range and example of principal echinate free fibres ascending to the surface, or may styles, b. Range and example of auxiliary megasclcres. be arranged in plumose columns without spongin. Aulena gigantea var. micropore/. - Lendenfeld, 1888: A sandy dermis may or may not be developed. 232. — l.endenfeld, 1889b: 100, pi. 9 fig. 2. - Whitelegge, These traits are based on l.endenfeld's observations

1901: 93, 1 18 (Tuggerah Beach, Port Jackson, Maroubra and figures (1889b, pi. 8 in particular; the tangen- Bay, NSW). -Whitelegge, 1907: 504 (off Barranjoey, tial dermal spicules in fig. 7 are more likely NSW).-Guiler, 1950: 10 (Blackmans Bay, Tas.). foreign). Aulena gigantea var. intermedia. - Lendenfeld, 1888: Either principal or auxiliary megascleres, or 232. -Lendenfeld, 1889b: 100, pi. 8 figs 3, 4, 7, 8, 18, 19; pi. 9 figs 3, 4. both, may be rare or wanting. Aulena and Halme sensu latter Aulena gigantea var. macropora. — Lendenfeld, 1 888: Lendenfeld (1889b), the without Halme 231 (Fremantle, \VA: Port Jackson, NSW). -Lendenfeld, (formerly Aulena) villosa (a Dysiclea), are in prac- 1889b: 99. - Whitelegge, 1889: 187 (off Green Point, Port tice indistinguishable except for virtual or total lack Jackson). of proper spicules in ''Halme". Halme irregularis Lendenfeld, 1889a: 49 (in key).— Eehinoelathria arenifera Carter is here treated Lendenfeld, 1889b: 453 (collective name for two varie- as a doubtful synonym because of its principal ties, no nominal subspecies). megascleres having been described as fusiform sub-

Halme irregularis var . lamellosa Lendenfeld, 1889a: 49 tylostyles (like those in E. favus), while those in (in key). -Lendenfeld, 1889b: 454, pi. 1 1 fig. 3 (Port Phil- are hastate. Dendy (1896) noted lip Bay, Vic). E. laminaefavosa Halme irregularis var. [micropora] Lendenfeld, 1889a: proper spicules in three of his specimens but prob- - ones. 49 (in key, nee Halme micropora Lendenfeld, 1 8851")- ably none in two other Lendenfeld, 1889b: 455, pi. 9 fig. 8 (Port Phillip Bay, The present specimens are palm-to hand-sized, Port Jackson). cake-shaped. F51969and E51971 are of the "laxa- type", having wide, rather irregular compartments, Material examined. Station KG 1, one specimen (F5I969); partitions thin in the former, intermediate in station KG 5, two specimens (F51970, F51971). Type with specimens of Holopsamma laminaefavosa (BMNH); of the latter. F5 1970 is of the "irregularis-type", with Halme nidusvesparum, II. laxa varr. minima and digitate; very wide compartments and contorted partitions Aulena laxa varr. minima and digitata; Halme gigantea extending into partly free lamellae. The colour in varr. micropora, intermedia and macropora; Halme life was noted as dull orange (5 YR8/6) in F51970, irregularis varr. lamellosa micropora (AM and and a deeper orange (5 YR7-8/10-12) in F51971. BMNH). Spiculation: Only dissociated preparations. Diagnosis. Massive, cake-shaped to lobose, or F51969 has frequent styles among foreign debris, thickly digitate, light to dull orange-brown, easily 52.6-60.5-68.1 X 2-2.9-3.1 /an. Auxiliary torn, thoroughly honeycombed; always full of megascleres: bent to flexuous strongyles, with wide sand. Proper spicules may be rare or lacking. Very axial canal, 128-/45-160 x 1.7-2-2.2 /an, about common in Bass Strait. 10% of all proper megascleres. In other two specimens only auxiliaries as proper megascleres, mostly Remarks. As with Trachycladus laevispirulifer, the subtylostrongyles, weakly mineralized and flexu- extensive synonymy reflects the polymorphism of ous, 148-0.9-1.8 iim in F51970, 155-160 x 1-1.5 this species. The variability relates to general shape ,xm in F51971. (massive, cake-shaped to globular or lobose, coarsely digitate), regularity or irregularity of the honeycombed, lacunose or lamellate surface pat- Kchinoclathria leporina (Lamarck) tern, prevailing width of compartments, relative Plate 6 figure 1, plate 24 figures 4, 5, width of partitions. With regard to skeletal structure, variability affects relative abundance of spon- text-figure 42 62 F. WIEDENMAYER

Spongia leporine/ Lamarck, 1814 (1813-1814): 444 microscleres. frond (Southern Seas). Description. F51973 small, 6 cm high; main Echinoelathria tenuis Carter, 1885e: 355 (Port Phillip 2.8 cm wide, bears two small secondary fronds on Heads, Vic.). upper half. Growth lines and radial striation barely Ophlitaspongia tenuis. — Dendy, 1896: 37 (revision; perceptible. Other specimens like Lamarck's holo- Port Phillip Heads). -Hallmann, 1912: 261, text-tig. 56, type, as figured by Topsent, but both comprise pi. 35 fig. 1 (Port Phillip Bay; W of Kingston, SA).- more of the distally incised palmate branches. Topsent 1932a: 101, pi. 6 fig. 1 (redescription of holo- Height of F51974, 33 cm, that of F5 1968, 20 cm. type of Spongia leporina; discussion). — Burton, 1934a: irregular pattern of longitudi- 599 (Great Barrier Reef). Fronds marked by transverse nee Ctathria tenuis. - Hentschel, 1911: 377, text-fig. 49 nal ridges, truncated or alternating along (fide Hallmann, 1912). growth lines in form of swellings curved parallel Phakellia papyracea Carter, 1886g: 379 (Western Port to frond margins. Pattern more pronounced than Bay, Vic; fide Dendy, 1896; nee Phakellia papyracea on Topsent's and Hallman's illustrations. On lar- = Phakettia wellsi de Lauben- Ridley and Dendy, 1886, gest specimen, some ridges branch and/or fels, 1936a, nomen novum, here transferred to Axinosia). anastomose, and may extend into small secondary Antherochalina perforata Lendenfeld, 1887a: 788 fronds, more often longitudinal, occasionally trans- (part?), pi. 22 fig. 44 (Broughton Island, NSW).- verse. surface-structure modified into a rugo- Lendenfeld, 1888: 89.- Whitelegge, 1902b: 279, 287 (part: Same schizoholotype and two doubtful "syntypes" reticulate pattern in one area. Fronds, where almost redescribed). — Burton, 1934a: 558 (holotype revised: syn- smooth, 1-2 mm thick. In largest specimen, fronds onymised with Ophlitaspongia tenuis). pierced by few round lacunae, 1-3 mm wide. Some Antherochalina tenuispina Lendenfeld, 1887a: 789 are submarginal and partly open, like in (Western Port Bay, Vic). -Hallmann 1912: 265 (schizo- Antherochalina perforata as figured in Lendenfeld holotype re-examined). — Burton, 1934a: 558 (holotype (1887a). revised: synonymised with Ophlilaspongia tenuis). In life vermilion (7.5 R6/12), with thin paler coat in some places (7.5 R7/10). Rubbery to leathery, Material examined. Station KG 6, one specimen (F51973); hard in stem. Oscules scanty and surrounded by station KG 7, two specimens (F51974, F51968). Holotype of Echinoelathria tenuis (BMNH); type specimens of radiating, dendritic-confluent subdermal canals in Phakellia papyracea (BMNH); holotypes of Antherocha- life, pattern invisible in alcohol. Ostia conspicu- lina perforata and A. tenuispina (BMNH). ous, regularly scattered in depressions on both sides of fronds. In small specimen (pi. 24 fig. 5), mesial Diagnosis. Complex-frondose, with terete, basally region comprises a tight subisodictyal reticulation expanded stalk. Fronds 2 mm thick, distally lobate, of thick yellow spongin fibres, with rounded incised to variable depths, occasionally lacunose; meshes equal in width or smaller than fibre- their surface in large specimens with radial and con- diameters, encasing principal megascleres of centric ribs. Rubbery in fronds, firm in stalk. Ver- narrow size-range (73-97 x 3.2-5.7 /an), with rare milion, with thin pale coat in places. Oscules larger principal and auxiliary megascleres. Few scanty, commonly stellate with branching, radial fibres somewhat longer and multispicular. canals. Fibro-reticulate subdermal and spicular Peripheral skeleton with distinct ascending and skeleton tending to radial-penicillate differentiation paratangential fibres, larger meshes, pale spongin, periphery and surface. Megascleres typical, in no principal megascleres of wide size-range, and

b r O:

Figure 42. Echinoelathria leporina, spicules from F51974, x 402414. a. Range and examples of prineiml stvles- h ' ' Range and example of auxiliary subtylostyles. ' SPONGES FROM BASS STRAIT 63

numerous auxiliary megascleres, of which outer- Holopsamma crassa (BMNH, sec table 1), Halme globosa most tend to be arranged perpendicular to surface. (BMNH), Halme micropora (AM and BMNH). For com- In largest specimen, spongin meshwork of mesial parison: holotype of Phortosopongia reticulum Marshall, 1880 region wider, multispicular fibres common; inter- (ZMB, with sehizotype and slide). mediate and larger principal, and auxiliary Diagnosis. Small, round to irregularly lobate, megascleres abound. In periphery, auxiliary thoroughly sandy, more or less deeply pitted to megascleres scattered at random below dermis, lacunose. Pits and lacunae round to elongate, some chiefly paratangentially in dermis, between regu- contorted, mostly 1-3 mm wide; tabulate to convex larly hispidating outermost principal styles (pi. 24 interstices of similar width. Pits may be covered fig. 4). by recessed sandy dermis bearing central oscule. Spiculation (F51973): (1) Principal styles and Internally cavernous. Firm, easily broken. Greyish subtylostyles, rarely straight, mostly curved or flex- buff to yellow, finely mottled by coarse sand. uous, 69-724-281 x 3.2-5.5-9.4 urn. (2) Auxiliary Small, thin strongyles only (?) subtylostyles, mostly straight, occasionally flexu-

ous, 179-249-312 x 2-2.4-2.9 ,uti. Description. Specimens contiguous in life, small, largest (F51975) 5.5 cm in greatest dimension. This Remarks. The skeletal structure agrees in general specimen (pi. 6 fig. 2) the most regular, cocoon- with Hallmann's detailed description. The varia- shaped, growing around several branches of brown tions ascribed by this author to age and growth alga. Other three specimens irregularly lobate. stages within the same specimen could not be Lacunose pattern more regular in largest specimen, observed, mostly because time did not permit mul- with round to elongate openings, sharp, flush to tiple sectioning. Variations of a similar kind were slightly recessed rims; irregularly cavernous observed in transverse sections of specimens F51973 interior. Some elongate openings contorted, often (small) and F51974 (largest). The differences could like footprints. Tabulate to slightly convex inter- equally be individual, or the result of unequal orien- stices about equal in width to lacunae, in F51977 tation with respect to longitudinal ribs (not and F52077, interstices discontinuous in some observed in sectioning). areas, lacunae larger, winding or sinuous. F51976 Exhinoclathria globosa (Lendenfeld) (pi. 6 fig. 3) has most lacunae closed by deeply concave fleshy dermis, reinforced by scattered sand Plate 6 figures 2-5, 7, text-figure 43 grains, occasionally pierced by a small oscule. Some Holopsamma crassa Carter, 1885b: 211 (part: at least of these pits are composite, petaloid. 6 of 31 symypes; Port Phillip Heads, Vic, below 20 I'm). In alcohol greyish buff to yellow, finely mottled /1i//cTO7awsa.-Lendcnfelds, 1889b: 101, pi. 8 figs 1, by sand grains, difficult to grade (about 10 YR - 2, 5, 6, 22, 23; pi. 9, figs 5, 9 (description and figures 2.5 Y7-8/6). In alcohol firm to hard, moderately from unspecified Carter symypes). to barely compressible, easily broken. Sand always nee — Psammopemma crassum. Lendenfeld, 1 889b: 638 coarse, packed at surface, irregularly scattered to (partly based on other unspecified Carter symypes of crowded internally. detailed observations Holopsamma crassa). No on skeletal structure. Echmoclathria crassa. - Hallmann, 1912: 287 (discussion, wrong conclusions). Spiculation: Only dissociated preparation Ha/me globosa Lendenfeld, 1885f: 303 (Port Phillip (F51977): thin strongyles, straight to slightly bent Bay, Vic; St. Vincent Gulf, SA, shallow water).- or flexuous, 127-/57-148 x 1-1.3 /mi. Lendenfeld, 1888: 157 (old records).- Lendenfeld, 1889b: 456, pi. 9 fig. 11. Remarks. The polymorphic concept of this species, Halme micropora Lendenfeld. 1885f: 304 (Illawarra, as reflected in the synonymy given above, needs to near Wollongong, NSW, on beach). - Lendenfeld, 1888: be justified and clarified, albeit provisionally, with 159 (old record). -Lendenfeld, 1889b: 461, pi. 9 fig. 12, reference to the confused history of past descrip- pi. 11 fig. 4 (not synonyms, old record). tions, and to what I know of the type material

Material examined. Station KG 4, four specimens involved. (F51975, F51976, F5I977, F52077). Type specimens of Carter's description of Holopsamma crassa is quite ambiguous and unsatisfactory, considering the notorious taxonomic difficulties affecting the Australian sandy sponges. The extant type series of Holopsamma crassa is certainly composite; though only macroscopic examination was poss-

Figure 43. Echinoclathria globosa, spicules from F51977. ible during my survey at the British Museum (May, Interstitial strongyle, the only type found, x 390. 1983), at least five species are obviously 64 F. WIEDENMAYER

represented. In Carter's manuscript catalogue of regular central oscules, smooth or reinforced by sponges from Port Phillip Heads and Western Port sand grains; (2) degree and regularity of interior Bay (J.B. Wilson collection), identified and cavernosity; (3) presumed absence of either prin- described by him, and deposited with the type speci- cipal styles or auxiliary strongyles, or both (in anal- mens, 24 syntypes (of which one wet, 23 dry) are ogy to Echinoclathria iaminaefavosa); and (4) listed for Holopsamma crassa. According to the variability of spicule dimensions (Lendenfeld indi- BMNH register and specimen labels, 31 specimens cated 200 x 3(im for strongyles). are now catalogued (not all of which were found A final word of caution is needed with regard during my survey). Though some mistakes are to Phoriospongia reticulum Marshall, which is known to occur in Carter's catalogue, it seems more externally indistinguishable from Echinoclathria likely that some specimens were broken (acciden- globosa. The holotype (pi. 6 fig. 6), from Tasma- tally or for study) and fragments subsequently nia, could be re-examined from a schizotype, registered separately in the British Museum. The through the kindness of Dr and Mrs D. Kiihlmann, results of my survey of this type series are Curators in charge, Museum fur Naturkunde der represented in table 1. Humboldt-Universitat East-Berlin. Its spiculation

Lendenfeld (1889b), who had examined most, if is distinctive: (1) Strongyles, mostly modified to not all syntypes of Holopsamma crassa, described subtylostrongyles, subtylotes, and (rarely) tylos- two different species based on largely unspecified trongyles, variably curved, bent or flexuous, syntypes of Carter. He compounded the confusion 204-222-235 x 1 .9-2.4 pm; (2) sigmata, often con- by using Carter's specific name for both species. torted, very abundant intcrstitially (pi. 24 fig. 3), Aulena crassa sensu Lendenfeld is obviously iden- 22.3-50.5-36.3 x 1.2-2.4 /tm. tical with Echinoclathria globosa, as here under- Echinoclathria egena sp. nov. stood, as particularly some of his figures (pi. 8 figs

1, 2; pi. 9 figs 5, 9) suggest. The figures of plate Plate 6 figure 8, plate 24 figure 6, 8 are diagrammatic, and those on plate 9 could not plate 25 figures 1,2, text-figure 44 be matched with any specimens I have examined, hence are probably of specimens not found in the Holotype: F51978, Station KG 6 (Tasmania, Winter Cove, dry systematic collection. The other species, E side of Deal Island, Kent Group; boulders with algae, described by Lendenfeld as Psammopemma eras- depth 3-6 m). sum, without figures, is more puzzling. It is partly Diagnosis. Encrusting, lobate and bullate base with based on unidentified Carter syntypes (more likely short, knotty, compressed, irregular branches. among those not found), partly on specimens of Oscules conspicuous, numerous, raised, scattered. his own, none of which I could trace during my Spongy to limp. Dull orange when fresh, turning surveys in Sydney, London, and East Berlin. The purplish brown. Surface smooth. Regular fibre net, description is suggestive of a Phohospongia mostly condensed in periphery. Megascleres typi- without, or with overlooked proper spicules, or, cal but small, principals coring primary fibres only. perhaps, of a Psammoclema as understood below. No microscleres. But its status as a good species, and hence the need for a replacement name, are doubtful at present, Description. From incrusting base of irregular, contrary to the allegation in Hallmann (1912: 287). sinuous outline, 1-5 mm thick, several knotty and It could be argued that a lectotype of Holop- compressed branches arise, mostly oblique, con- samma crassa should be designated from among torted, up to 4 cm long, 0.3 to 2.5 cm wide. Oscules the two specimens illustrated by Lendenfeld (1889b, conspicuous and numerous on base and branches, pi. 9), in which case crassa (published in March) mostly raised on volcano-shaped or bullate eleva- would have priority over globosa (published after July). That specimen, however, has to be identified first as extant, and re-examined, including spiculation. In the meantime, it seems safer to use globosa, the holotype of which, previously known only from Lendenfeld's inadequate figure, is here figured anew (pi. 6 fig. 4). The polymorphism of Echinoclathria globosa, as here conceived, is apparent in (1) the conforma'- tion of the surface: regularity in distribution and contours of lacunae, relative width and convexity Figure 44. Echinoclathria egena of interestices, presence or absence of a dermis sp. nov., spicules, x 342. tym- a. Range and example of coring styles and panising or recessed in the lacunae, subtylostvles. with or without b. Range and example of interstitial strongyles. A

SPONGES FROM BASS STRAIT 65

Table 1. Type series of Holopsamma crassa Carter in the British Museum (Natural History).

BMNH register Preservation Location in type Additional MS. entries Present interpretation, number collections on label' remarks

86.12.15.58 wet 4. IV. " ." = Ancorina. . Ancorina sp., has (R.K. 21/7/08) sandy cortex

86.12.15.313, dry K. 63 "92, 12, 39" Echinoclathria -314,-316 (H.J.C.), "Holo- globosa (Ldf); psamma tuberculata" 86.12.15. 313 here and figured, pi. 6, "Psammopemma fig. 7 tuberculata var." (Ldf.)

86.12.15.315 dry not found

86.12.15.317 dry K. 63 "54" and "PW11" Psammoclema (H.J.C.), "Holo- densum (Marshall) psamma tuberculata" and Psammopemma tuberculata var." (Ldf.)

86.12.15.318 dry K. 63 "103" and remark Psammoclema on colour in life fuliginosum (Carter) (H.J.C.) "Holo- psamma fuliginosa" (Ldf). "Psammo- pemma fuliginosum"

86.12.15.409 dry not found

86.12.15.410 dry K. 37 "21" (H.J.C), "Dys. Echinoclathria rugosa" (Ldf.) globosa

86.12.15.411 dry not found

86.12.15.412 dry K. 62 "52" and "conulated" Phoriospongia (H.J.C.) wilsoni (Dendy)

86.12.15.426-430 dry K. 62 "10" (H.J.C.) Phoriospongia wilsoni (Dendy); in box, possibly 5 frag-

ments of 1 specimen

86.12.15.431 dry not found

86.12.15.474-487 dry not found 14 syntypes

1 H.J.C. = Carter; Ldf. = Lendenfeld; R.K. = Kirkpatrick.

tions, 1-2 mm wide, some smaller. Mostly res- ermost tangential meshes of peripheral skeleton. tricted to edges of branches, some scattered on one Meshwork of strong spongin fibres regular, par- face of wider branches. In life dull orange (5 ticularly in outer portion. Primary fibres 27-52 fim YR7-8/12), turning dark purplish brown (5 R3/8) in diameter, thicker at surface; secondaries 16-40 in elevated areas when drying. Spongy, moderately /an in diameter. Meshsize larger in deeper choan- firm in branches, limp in base. some, up to 350 x 500 /an, typically around Choanosome dense, but base contains conspic- 120-175 ixra. Common peripheral condensation, uous meandering canals, 3 mm wide. Surface opti- 300-350 fim deep, of closely spaced secondaries and cally smooth, but slightly rough, micropapillate adventitious primaries, with mesh sizes mostly with protruding ends of primary fibres. Ostia equal to greater fibre-diameters. Peripheral fibres mostly conspicuous, regularly scattered inside out- commonly stratified, with 1-2 layers of granular 66 F. W1EDENMAYER

Secondaries aspiculous. Air matter. Primary fibres cored by styles and sub- plumosely arranged. scattered. No microsclere* tylostyles, parallel, single or up to 4 abreast, with iliary styles irregularly points out, occasionally askew, frequently broken. Description, Four stout tubes, concrescent basally mostly aspiculous. Thin slrongyles Secondary fibres and by pairs along contiguous sides. Width 6 scattered at random interstitially, frequent in 3 cm, maximum height 5 cm. Individual tubes 1. scarce below. Some adventitious periphery, cm wide, with apical oscules and atria 5-6 mr peripheral skeleton free, bud-like, primaries in wide. rarely protrud- encasing styles singly or in brushes, - In life dull vermilion above ( 10 R 2.5 YR7/10) ing; more frequent at surface, mostly blunt or trun- dull yellow-orange below (2.5Y9/10). Firml; cate, like tips of main primaries. spongy, slimy in life. Ostia now conspicuous- Spiculation: (I) Coring styles and subtylostyles, 0.1-1.5 mm wide, mostly collapsed into subderma fusiform, thickest in the middle, length spaces. Surface microhispid. Atria terete, taperini 38 45.8-52.9 fim, median/basal width near base. Choanosome cavernous, riddled b; 2.3-3.1/1.2-2.1 /mi; (2) interstitial slrongyles, canals as wide as subdermal spaces.

1 24-/46- 178 x 1-2.4 /mi. fibres irregular I ax meshwork of spongin Remarks. The specimen contains many embryos, primaries 34-46 /un thick, often crooked, secon

C, 175 /

is some resemblenee to Slylinos (probably compo- fibre-meshwork, particularly crowded at surface. site), particularly Slylinos scariola (Lamarck), Microscleres absent. which also has fusiform, subbasally constricted Spiculation: (1) Principal styles, hastate, subtylostyles (fide Topsent, 1932a: 95). 102-/25-132 x 3.5-4.9-7 /

specimen Material examined. Station KG 6, one (F51979). Desmacididae Schmidt, 1870 Two syntypes of Ophlitaspongia tubulosa (AM). Synonym: Esperiopsidae Henlschel. 1923. Diagnosis. Short, stout tubes, concrescent basally and partly above. Single apical oscules. Vermilion Remarks. Van Soest (1984: 35) repeated Levi's error

to dull yellow-orange, firmly spongy, slimy in life. (1973: 611) of using Esperiopsidae Hentsehel, 1923, Surface smooth; thin dermis veiling regular pune- and of disregarding the priority of Schmidt's family tiform verstibules. Internally cavernous. Fibre net name. Both authors cited Dcsmacidonidae (pars) irregular, lax. Primaries with hastate styles often as synonym, and include Desmacidon in the family.

As I have already stated (Wiedenmayer, 1977a: 79, footnote 4), Schmidt's original spelling (as Des- macidinae) has to be retained in accordance with ICZN Article 29bii (see also Appendix DVII, Greek entry no. 25, excluding nos. 32, 33), making Des- macidonidae and Desmacidontidae incorrect subsequent spellings. This provision does not at feci the gender of Desmacidon, which was established as feminine by Bowerbank (1861), in accordance with ICZN Article 30d (30bii in previous editions, Figure 45. Echinoclathria tubulosa. not 30ai2 as spicules, x 197. a slated in my footnote). Though all Principal styles, b. Auxiliary styles. names published in Bowerbank (1861, merely a list) SPONGES FROM BASS STRAIT 67

are nomina nuda, his intention therein is clear and conclusions that Balzella is a junior synonym of consistent, all adjectival species names associated Stylotella, and that Stylotella is related to Phori- with Desmacidon and Hymen iacidon being femi- ospongia. further considerations, arising from the nine. Later, Bowerbank was still consistent in 1864, study of our diverse material of Phoriospongia and but inconsistent in 1863 and 1866 (feminine and Psammoclema, prompted the inclusion of this masculine species names in both genera). If there group in the Desmacididae. This view was con- are any lingering doubts as to proper fixation of firmed at this time by van Soest's (1984: 47) plac- the gender of these genera, reference should be ing of Balzella in the same family. made to the footnote in Thiele (1905: 421), which Apparent discrepancies and difficulties, in for- I had overlooked in 1977a. Though Thiele's etymo- mulating a definition of the Stylotellinae, are logical argument is open to doubt, his acceptance resolved by regarding them as extreme differences of the feminine gender is sufficient for the purpose of degree. Loss of spicules in sandy sponges is not of stability in nomenclature. uncommon, and also occurs in Echinoclathha. The family is here subdivided into two subfami- Absence of spongin (e.g. in Stylotella and in Psam- lies. The nominotypical subfamily corresponds to moclema densum) versus a well developed reticu- the family as previously understood Levi by (1973: lation of spongin fibres (e.g. Phoriospongia 61 If., part) and restricted van Soest by (1984: 48). guettardi Topsent, 1933) are linked by intermedi- The name Stylotellinae Lendenfeld is revived for ate stages. The spicular vicariance (ambivalence) a rather confused and controversial group of of monactinal versus diactinal megascleres, in itself genera, of which, beside the type genus, the most typical of the Desmacididae, is occasionally linked important are Phoriospongia and Psammoclema by intermediates within the same species, or even (as redefined below). While Styloielta was recently within one specimen (Phoriospongia reticulum, included either in the Suberitidae or in the described above; Stylotella inaequalis, described Hymeniacidonidae, Phoriospongia was included in below). the Myxillidae by Dendy (1924: 362), Topsent Of particular importance, in the definition of the (1928: 54, as Phoriospongiinae Lendenfeld, 1889), Stylotellinae, is the dissociation, if both are present, and Burton (1934a: 552). Phoriospongia, Psam- of the skeleton formed by spongin and/or sand mascus, Psammoclema, and their synonyms, were from the spicular skeleton. included by de Laubenfels (1936a: 96) in a new Desmaeidinae family Psammascidae, placing most emphasis, in the diagnosis, on the high content of sand in these Synonym: Lspcriopsidae sensu van Soest, 1984. sponges. The type genus Psammuscus was misin- Diagnosis. "[Desmacididae] with a reticulate or terpreted by de Laubenfels, by ascribing to it proper plumo-reticulatc choanosomal skeleton of monac- megascleres and microscleres, and by in merging tinal or diactinal megascleres. Ectosomal skeleton it Phoriospongia. The original definition of Psam- absent (but organic dermis often strongly developed mascus and the description of its type species (P. or charged with foreign material). Microscleres [if decipiens, still alone in the genus) by Marshall present] include palmate, anehorate or unguifer- (1880: 92f.) contained no mention of proper spicate isochelae and sometimes sigmata." (Van Soest, ules. Vacelet, Vasseur and Levi (1976: 63) followed 1984: 35, for Lsperiopsidae.) Special types of de Laubenfels in erroneously placing Phoriospon- microscleres (birotulates, placochelae, bipocilli, gia lamella Lendenfeld in Psammuscus. The genus canonochelae) occur in particular genera. was still assigned to the Myxillidae by these authors, Strongylaeidon Lenden feld in agreement with Topsent and Burton. In the Stylotellinae sensu novo, the spiculation, as a rule, Slrongylacidon Lendenfeld, 1897b: 1 10. -Topsent, is scanty, if present, both in density and in weak- 1928: 45. -Burton, 1934a: 555 (diagnosis in Myxillidae, ness of individual spicules. The arrangement of included species). -de Laubenfels, 1936a: 54 (in Des- macididae). -Levi, 1973: 611 (definition in Lsperiopsi- megascleres, if present, is poorly organised and dae). — Van Soest, 1984: 42 (definition in Esperiopsidae), microscleres, if present, are commonly sigmata 46 (discussion). only. The exceptions are Psammochela and Psum- motoxa, probable synonyms of Phoriospongia. The Diagnosis. "| Desmaeidinae) with plumosely reticulate spicular skeleton, well developed and arranged bundles of strongyles for megascleres and often rather complex microsclcre complement, and unguiferate or anehorate isochelae and [or] sigmata the special ectosomal skeleton of the Tedaniidae for microscleres." (van Soest, 1984: 42). (Myxillidae auctorum) are not or only exception- Microscleres may be absent. Reticulate skeleton ally developed in the Stylotellinae. only in thickly encrusting, massive and branching

The study of the new material of Stylotella itiae- specimens, plumose ascending columns with only quail's (Hentschel) (described below) has led to the basal anastomoses in thinly encrusting specimens. 68 F. WIEDENMAYER

scarce, irregularly scattered in depressions. Ostia inconspicuous, in rows along fine meandering and dendritic subdermal canals. Skeletal structure (pi. 25 figs 5, 6) irregular reticulation of fibres filled with megascleres; these also scattered interstitially in choanosome and dermis. Reticulation irregular in orientation and thickness of fibres (35-150 /mi), and in extremely variable shape and size of meshes. Individual fibres straight, angular or sinuous; compactness varies, some break up, intergrading with surrounding interstitial megascleres. Clear spongin barely visible in more r compact fibres. Thicker fibres usually more continuous. Short ones (anastomoses) frequently expand at junctions. Dermal skeleton of thinner

Figure 46. Strongylacidon stellidernw, spicules, a, b. spiculo-fibres; arrangement lacey, vaguely umbel- Strongyles of two size ranges, x 322. c, d. Enlarged ends, lar, or, most frequently, festoon-like (closely x948. e. chelae, x948. spaced parallel fibres arching inwards between principal fibres forming axes of conules). Dermis 350-430 /mi thick, dense, light brown in Strongylacidon stelliderma (Carter) sections, with granular collagen, few interstitial Plate 6 figures 10, 11, plate 25 figures 5, 6 megascleres. Choanosome distinctly lighter, less collagenous, riddled by incurrent canals, around text-figure 46 50 /im wide. Excurrent canals around 120 /im wide, Halichondria? stelliderma Carter, 1886h:451 (Western- commonly surrounded by denser matrix with port Bay, Vic). apochetes and some megascleres both radial. Small, - 1896: 20 (holotype Desmacidon stelliderma. Dendy, frail chelae numerous everywhere, crowded in lin- re-examined, new records from Port Phillip Heads).— ings of most canals. Br^ndsted, 1927: 3 (Port Chalmers, New Zealand). Spiculation: (1) Strongyles, often with one or Strongylacidon stelliderma. — Burton, 1934a: 554, 555 both ends slightly swollen to subtylote, of two size (transfer and synonymy only). Burlonispongia stelliderma. — de Laubenfels, 1936a: 53 ranges: (a) 108-/77-126 x 0.8-7.6-2.2 /mi and (b) (transfer to gen. nov. only). 176-755-197 x 1.2-/. 9-2. 5 /tm; both sizes occur Desmacidon plicaluin Hentschel, 1911: 321, text-fig. interstitially, distinction in fibres not determined. 18 (WA). (2) Frail unguiferous anchorate chelae, C-shaped, teeth atrophied, little recurved if at all, Material examined. Station BSS 181, one specimen chord (F51981). Holotype, BMNH 1886.12.15.148, wet, from 9.4-70-11.1 /mi, depth 3.4-J.5-4.4 /mi. Westernport Bay, ex J.B. Wilson collection.

Diagnosis. Small, massive-lobate, or irregularly Remarks. The holotype of this species (pi. 6 fig. compressed, frondose to ramose, with encrusting 11) is massive, bulbous, with conspicuous apical base. Spongy to limp, easily torn. Surface regularly oscules. The conules are blunt in the apical region, conulose to lacinulate, with axial spicular column. but pointed below, with tips of spiculo-fibres regu- Oscules scarce, irregularly scattered, apical in larly protruding. Ostia, distinctly visible at low massive-lobate specimens. Irregular, angular net of magnification, are regularly scattered in depressions compact to disheveled fibres packed with stron between conules. Strongylacidon sansibarensis Len- gyles, which are also scattered interstitially. Struc- denfeld (1897b, the type species) is little known ture denser, lacy or festooned below surface. Frail It was described as thickly encrusting, finely conu- chelae numerous throughout. lose, permeated by a dendritic alga, and possess- Description. Attached to chaetopterid worm tube. ing, as only microscleres, unguiferous anisochelae

Base encrusting, about 1 mm thick, bearing few 6.7-9 /im long, and strongyles measuring 160-260 short irregular fronds and branches. Thinner x 3-4 pm. Strongylacidon intermedia Burton branches angular, with polygonal cross-section, (1934a) and S./ratex(Pulitzer-Finali, 1982b), from larger ones much compressed, almost paper-thin. the Great Barrier Reef, both lack microscleres (see Colour in life not noted, now off-white. Limp, below, Remarks on Stylotella). The genus is well easily torn. Surface finely conulose, long conule represented in the West Indian region (van Soest, tips often recurved and ending in whiplike exten- 1984: 42f., with descriptions of three new species, tion of primary spicular fibre. Density of conules all thinly incrusting, and discussion of other spe- variable, highest on edges of branches. Oscules cies, some as species inquirendae). -

SPONGES FROM BASS STRAIT 69

Slylotellinae Lendenfeld, 1888, sensu novo loss of chelae (1984: 7, table l,b) and his greater emphasis on skeletal structure. Synonym: Phoriospongiinae Lendenfeld, 1889.

Diagnosis. Desmacididae with reduced spiculation. Slylotella inaequalis (Hentschel) Megascleres monactines or diactines or both, with Plate 7 figures 1, 2, plate 26 figures 1-4, intraspecific and individual transitions. text-figure 47 Microscleres, if present, usually sigmata only, but Batzella inaequalis I chelae, toxa, microtylotes and rhaphides may Hentschel, 1911: 325, text-fig. 20 occur. Microscleres or the whole spiculation may (Shark Bay, Denham, Albany, WA).- Pulitzer-Finali, 1982b: 98 (in discussion). be lost, in some species always, in others in some nee Strongylacidon inaequalis. — Burton, 1934a: 550 specimens. Spongin skeleton, when present, retic- (Great Barrier Reef). - Burton, 1959: 242 (Arabian coast). ulate or dendritic. All spicules usually interstitial nee Cacochalina truncatella var. mollissima Lenden- except in type genus, where spicules also occur in feld, 1887a: 763 (cited as synonym of Strongylacidon inae- whispy, vaguely plumose but not reticulate fibres. qualis by Burton, 1927a: 292, and 1959: 242). Most species incorporate much foreign detritus, Collosclerophora I urenacea Dendy, 1917: 321, pi. II commonly aggregated or encased by spongin, (Port Phillip Heads, Vie.). -Dendy, 1921: 133, text-fig. [ and/or scattered to packed interstitially. Com- 49 (discussion). -Dendy, 1922: 74, 76 (discussion). monly a cortex packed with debris, but missing in Material examined. Station BSS 181, seven specimens some species or individuals. Cortical | concentration (F51982 through F51988). of erect or felted proper megascleres may occur. Diagnosis. Small, variably encrusting, irregularly Stylotella Lendenfeld lobulate, or with short, crooked, frilly branches more or less concrescent. Frequently attached to Slylotella Lendenfeld, 1888: I85.-Hallmann, 1914b: or enveloping tall objects (worm tubes, algae). 348 (discussion, revision: to Suberitidae).— Burton, Limp to rubbery. in 1934b: 41 (discussion, synonymised with llymenia- Brownish orange life. Surface cidon). — 6e Laubenfels, 1936a: 152 (definition in Suberit- smooth, semi-diaphanous, occasionally finely ver- idae). — Bergquist, 1968: 24 (in Suberiiidae). rucose. Oscules small, round or slit-shaped. Spic- — Pulitzer-Finali, 1978: 72 (in Hymcniacidonidae, ules partly scattered, partly in loose, often Mediterranean records excluded). indistinct, whispy fibres of variable thickness and nee Stylotella (as applied to records) sensu European spacing. Peripheral umbels and felt of spicules may Topsent, 1925a: 637f. (Mediterranean).- Vacelel, I960: occur. Spongin scanty. Variable amounts of sand 264 (Mediterranean). — Borojevic, Cabioeh and Levi, and scattered foreign spicule debris internally and 1968: 17 (English Channel). -Boury-Esnaull, 1971: 3271'. at surface. Macro-and microsymbionts common. (Mediterranean with further references). Spicules variably strongyles or Bulzetlu Topsent, 1894b: xxxiv (English Channel). styles/subtylostyles, Thiele, 1905: 438 (discussion, not the two species often combined. included). — Burton, 1934a: 5541". (disseussion: monotypic; Description. Six specimens small, around 1 cm in definition in Myxillidae). — de Laubenfels, 1936a: 138 greatest dimension, attached to chaetopterid worm (definition in Hymeniacidonidae). — Pulitzer-Finali, 1978: tubes or to algae, together with other 68 (part: not Batzellafriabilis). - Pulitzer-Finali, 1982b: sponges. Thinly or thickly encrusting; 98 (in Halichondriidae: part; not B. frutex, not B. or irregular, with con- intermedia). — V an Soest, 1984: 47 (definition in Esperi- crescent lobes separted by thin invagination; also opsidae, discussion). with angular or compressed, crooked and folded Collosclerophora Dendy, 1917: 320.-Dendy, 1922: 74 branches, some crowded, others free. Seventh (discussion). — de Laubenfels, 1936a: 53 (definition in specimen (F51988, pi. 7 fig. 2) up to 4.5 cm high, Desmacididae). a group of compressed, frilly and aculeate branches from basal disc and pedicel, largely pseudo- Diagnosis. "[Slylotellinae] with a reduced, loosely morphic, from growing around agal plumose skeleton of strongyles (tornotcs); no branches. Compressible, limp in branches, firmer, rubbery ectosomal skeleton; no microscleres." (Definition in lobes and thick encrustations. In life brownish of Batzella [in Esperiopsidae] in van Soest, 1984: 47).

Remarks. Batzellafriabilis and B. frutex Pulitzer- Finali possess a reticulate skeleton, and so does Strongylacidon intermedia Burton (1934a), which was transferred to Batzella by Pulitzer-Finali (1982b). These species are best included in Stron- gylacidon, despite the lack of chelae. This is con-

sistent with van Soest 's recent revision of West Figure 47. Stylotella inaequalis, spicules, a. Strongyles (subtylostrongyles), 394. b, Indian Poeeilosclerida, particularly with regard to x c. Enlarged ends, x 1 169. 15

70 F. WIEDENMAYER

subdermal canals; finely, orange (5 YR5/12 in F51987, 7.5 YR5/8 in dritic "and anastomosing larger specimen. Surface F5 1988); beige to cream in alcohol. Surface gener- irregularly conulose in protruding spicules. Some ally smooth in small specimens, with narrow, den- partly microhispid with

Table 2. Skeletal features and symbionts in Stylotella inaequalis

Specimen Skeletal Dominant Spicular dimen- Foreign debris Symbionts structure* spicule types sions (^.m)

F51982 2 + 5 (periph- Strongyles 141.0-755-161 x Few sand grains Macro ery); 4 (centre) 1-1.3

F51983 2 (periphery); Subtylostyles 231-245-262 X Much coarse 4 (centre) 2.4-2.9-3.6 sand in vaguely dendritic columns

F51984 2 + 4 (periph- Strongyles and 154-/6/-168 X Few sand grains Macro and ery); 4 (centre) sub- 0.9-/. 3-1. micro (mulberry tylostrongyles type, 235-275 /on, wide, scattered)

F51985 1+5 Strongyles, fusi- 193-207-219 x Few sand grains Micro, crowded form, slightly 1.8-2.3-2.7 (clusters of 2-5 subtylote cysts, total diameter 16 x 12 ^m)

F51986 3 + 6 Styles and sub- 121-182-249 x Spicule frag- Macro tylostyles, 1.3-/. 9-2. ments concen- fusiform trated at surface. Few sand grains

F51987 3 + 6 + 7 Styles and sub- 172-2/0-238 x Spicule frag- Macro tylostyles, 2-2.5-3.2 ments concen- fusiform trated at surface. Few sand grains

F51988 2 + 5 + 8 (peri- Subtylostyles, 158-/5(5-201 x Frequent sand Macro phery); 5 fusiform 1.6-2.0-2.3 grains and spic- (centre) ule fragments scattered and clustered

•1. Fibres of variable thickness, widely spaced, dendritic, gently curved. 2. Fibres closely spaced, thin, whispy. 3. Fibres thicker, closely spaced, very sinuous and angular. 4. Interstitial spicules scarce. 5. Interstitial spicules abundant, frequently transverse, single or in bundles. 6. Interstitial spicules crowded. 7. Conspicuous peripheral umbels and felted arrangement at the surface. 8. Spongin present in some fibres. SPONGES FROM BASS STRAIT 71

sand grains, particularly in F51983. Oscules small, There are obvious similarities between Stylotella round and clustered, or slit-shaped. In some speci- inaequalis and Stylotella agminata, particularly as mens, regular round subderma! spaces covered by redescribed by Hallmann (1914b: 349; pi. 21 figs sieve-like membrane. In other specimens, includ- 1-5, pi. 22 fig. 2; erroneously cited in description). ing largest one, fine reticulation of dermal mem- These concern growth form and skeletal structure brane conspicuous, with ostia in meshes, and their range of variability. However, inclusions accentuated by concentration of foreign spicule of foreign destritus were not described in Stylotella debris in dermis of two specimens. Dense, finely agminata and its synonyms. Tangential arrange- collagenous choanosome, brownish in transmitted ment of dermal spicules, not exclusively developed light. Dermis lighter, 15-25 *tm thick. Skeletal in S. inaequalis, only occasionally seen as a com- structure variable (pi. 26 figs 1-4, and table 2). ponent of a felted arrangement in one of our speci- Types in table 2 approximate, transitions within mens, seems to be the rule in S. agminata. The chief and among specimens common. In type 2, whispy distinction resides in the spicules: those in S. fibres may become angular, with spicules frequently agminata are always styles and sublylostylcs, the broken. Most common macrosymbionts, fre- latter with heads slightly thicker than the hastate quently concrescent tubes, 140-300 fim in diameter. shaft. Their dimensions exceed those in S. inae- Pale, clear spongin only in some fibres of largest qualis slightly in length, distinctly in thickness. specimen. Stylotella inaequalis links the hitherto misplaced Spiculation: Polarity between diactinal and genus with Phoriospongia and Psammoclema in monactinal spicules. Intermediate forms (subtylos- several features: (1) the presence of foreign detri- trongyles, styloids with the thinner end blunt or tus, with a tendency to segregation of sand grains mucronate) with dominant subtylotes, never fre- and spicule fragments, with concentration at the quent. Subtylote condition faint, thickest in fusi- surface and occasional arrangement in columns or form shaft; slight subterminal constrictions abrupt lamellae inside, but without specific or intraspecific inside elongate heads. Spicules may be mostly consistency; (2) the generally loose arrangement of straight (in F5I988), otherwise majority curved, spicules with little order except ascending tracts, bent or flexuous in variable degrees. Blackened never truely reticulate; (3) the common weakness axial canals frequent. Dimensions 121-/92-262 x of megascleres; (4) the inconsistent spicular vicar- 0.9-/. 9-3.6 urn. iance of diactinal versus monactinal megascleres, with little morphological differentiation within Remarks. Burton (1934a, 1959) misinterpreted specimens; (5) frequency of variably bent Hentschel's description. The sponges misidentified megascleres. These traits also advise against affini- by him have a coarsely conulose surface and a retic- ties with either the Suberitidae, Halichondriida, ulate skeleton regularly reinforced by spongin. Mycalidae, and Tedaniidae. They should probably be left in Strongylacidon. Cacochalina truncatella mollissima Lendenfeld Phoriospongia Marshall likewise has a reticulate skeleton. Hentschel's description agrees with the present Phoriospongia Marshall, 1880: 122. -George and material, particularly the modes of growth and Wilson, 1919: 155 (synonymised with Chondropsis). - attachment, the smooth surface, the semi- Dendy, 1924: 362 (definition, discussion, synonymised with Chondropsis).— diaphanous dermis, the skeleton consisting partly Topsent, 1928: 54 (synonymised with Chondropsis, Psaminoclwla). — Topaent, 1933: 50 of scattered spicules, partly of very loose spicular (footnote 1, resume, synonymised with Chondropsis).— de tracts of variable spacing, frequently merging when Laubenfels, 1936a: 96, 99 (synonymised with specimens have crowded. Though six of the present Psammascus). Hentschel's measure- weaker and shorter spicules, Holopsamma Carter, 1885b: 211 (part).-ncc sensu ments agree with those of F51983. Loeblich and Tappan, 1964: C792 (under The "colloscleres" in Collosclerophora arenacea Xenophyophorida, wrong type species). are here interpreted as foreign, probably microsym- Chondropsis Carter, 1886a: 45. -Dendy, 1895: 500 bionts, though of a kind not seen in the present (discussion). -de Laubenfels, 1936a: 99 (synonymised with Holopsamma). material. Globular and cystose microsymbionts, Sigmatella Lendenfeld, 1888: 195. -de Laubenfels, however, are commonly found in sandy Stylotelli- 1936a: 97, 99 (synonymised with Holopsamma). nae, and there does not appear to be an obliga- Psammoclwla Dendy, 1916b: 126. — George and relationship. Dendy (1922: 74) found tory/specific Wilson, 1919: 155 (synonymised with Phoriospongia).— identical "colloscleres" in Col- similar, though not Dendy, 1924: 362 (retained). - Burton and Rao, 1932: 334 is related to loclathria ramosa. The genus not (revision). -de Laubenfels, 1936a: 99 (definition in Stylotella, and was merged in Rhaphiduphlus (cor- Psammascidae). rectly Thalysias) by van Soest (1984: 109). Psammotoxa de Laubenfels, 1936a: 99. 72 F. WIEDENMAYER

wavy to meandering in periph- Diagnosis. Sandy Stylotellinae, variable in amount with debris in tracts perpendicular canals also wavy in por- and distribution of foreign detritus, in spiculation ery. Regular periphery, as in one of syntypes (pi. 7 fig. and in development of reticulate or dendritic spon- tions of cyanophycean algae as recorded by Carter gin fibres. Megascleres mostly frail strongyles, sub- 6). No distributed small tallo- tylostrongyles, subtylostyles, styles and subtylotes and Dendy; but regularly stuck in and just below with frequent transitions, scattered or in vaguely phyte algae, sac-shaped, specks in life). radial to plumose arrangement. Microscleres, if surface (greenish size and nature: in present, either sigmata, chelae, or toxa. Spicula- Debris tend to sorting by fragments, chiefly lithic, tion may be reduced to rare megascleres. Cortical choanosome, larger reinforcement by spicules or debris frequent. arranged in meandering tracts; finer detritus, chiefly spicule fragments, strewn interstitially. In ecto- Phoriospongia argentea (Marshall) some, coarser lithic fragments pack inner, thicker spicule fragments Plate 7 figures 3-7, plate 26 figure 5, portion; finer material, chiefly and spicules of didemnid tunicates, fill thinner sur- plate 38 figure 6, text-figure 48 face layer. Proper spicules mostly arranged radi- in Dysidea argentea Marshall, 1880: 107, pi. 7 figs 6-11 ally, few scattered randomly. Some crowded (Australia, probably Bass Strait). vague ascending tracts, occasionally in plumose Chondropsis arenifera Carter, 1886a: 45 (as Chrondop- fashion. sis, lapsus, Port Phillip Heads, Vic. (.-Carter, 1886b: Spiculation: weak strongyles, frequently flexu- 122,-Dendy, 1895: 255. ous, 101-/29-156 x 1-1.3 tim. specimen (F51989). Material examined. Station KG 6, one Remarks. The new specimen being small and not Holotype of Dysidea argentea (PMJ). Syntypes of Chon- very typical, the holotype of Dysidea argentea and dropsis arenifera (BMNH). two syntypes of Chondropsis arenifera are figured Diagnosis. Cushion-shaped or massive, bulbous or for comparison. Salient features are the tuberose lobate. Firm, moderately compressible. Cream to shape, mostly smooth, glabrous surface, firm, light beige in life. Surface smooth, glabrous, to fleshy choanosome, and slightly recessed oscules vaguely rugose and verrucose. Oscules conspicu- of irregular outline. Marshall did not mention ous, rounded-polygonal, often slightly recessed. proper spicules, but admitted to having examined Regularly ectosome 0.3-0.5 thick, armoured mm his specimen in a cursory way. Still, it is not grains finer detritus on surface, coarser deeper. unlikely that proper spicules my be absent in some Choanosome fleshy, with radial-meandering tracts specimens. For comparison with similar congeners, of debris and canals in periphery. Macro-and see Remarks on Phoriospongia kirki, below'. microsymbionts common. Debris in tracts often coarser than interstitially. Proper spicules weak Phoriospongia carcinophila (Lendenfeld) flexuous strongyles, mostly radially arranged. Plate 7 figures 8, 9, plate 26 figure 6, Description. Asymmetrically cushion-shaped, sinu- plate 27 figures 1, 2, text-figure 49 ous margin, 4.5 by 2.5 cm wide, 0.5 cm to 1 cm Sigmatella high. Margin with fine branching algae and carcinophila Lendenfeld, 1889a: 66 (in key). -Lendenfeld. 1889b: 615, pi. 41 fig. 8 (Port Jack- hydroids. In life and in alcohol cream to light beige, son, NSW). fine greenish specks in life. Firm, moderately Chondropsis carcinophila. — Burton, 1934a: 552 (Great compressible. Barrier Reef). Surface smooth, almost glabrous in life, rugose Material examined. in alcohol. One oscule, 2 mm wide, with a rounded- Station BSS 179, one specimen (F51990); station triangular recessed margin. BSS 181, two specimens (F51991, F51992). Type specimens of Sigmatella carcinophila (AM Fine, indistinct regular ostia. Armoured ecto- and BMNH). some 320-500 jj.m thick. Choanosome fleshy, firm, Diagnosis. Contorted-lobate, often small, frequently attached to shells or crabs. Spongy, soft to moderately firm, easily torn. Dull orange brown

48. Phoriospongia argentea, spicules, Figure a. Stron- Figure 49. Phoriospongia carcinophila, spicules (strongyles, x392. b. Enlarged end, X1165. glyles), from F51990, x306. SPONGES FROM BASS STRAIT 73

to purplish brown in life. Surface finely rugo- Detritus in ectosome coarser than in smaller vermiculate to verrucose. Oscules of 2 sizes not specimens, almost entirely lithic fragments, fre- above 2 mm wide. Detritus in ectosome and quently scattered. Detritus in choanosome scanty, choanosome ill sorted, variably dense, in meander- foreign spicules rare. Fragments in tracts frequently ing tracts below ectosome. Proper spicules varia- encased by spongin. Proper spicules abundant; in bly frequent, scattered in and around tracts of choanosome, interstitially and within fibres, mostly debris, and parallel to plumose in short ascending at random. Some fibres or their portions almost tracts below ectosome. They may be concentrated free of detritus, but always with proper spicules. in ectosome, vaguely umbellar to confused. Proper spicules particularly abundant in peripheral choanosome and at surface. In periphery frequently Description. F51990 (pi. 7 fig. 8) elongate, irregu- in closely spaced subparallel to plumose tracts. In larly lobose, somewhat compressed, 8 cm long, ectosome, arrangement vaguely umbellar to con- 2.5-3 cm high, 1-2 cm wide. Partly overgrows two fused, almost felted. Pecten shells and supports delicate branching algae, Spiculation: strongyles, straight, or bent and hydroids and bryozoans, some incorporated. flexuous to variable degree in all three specimens; F51991 (pi. 7 fig. 9) one-half of contorted-lobose strongest in F51990, weakest in F51992. Dimen- specimen growing on crab's back, 2x2x1 cm. sions forF51990, 151-756-208 x 1.3-2.3-3.1 ^m; F51992 compressed, rounded, margin and upper F51991, 156-767-176 x 1-7.6-2.2 ^m. sides with crowded, stubby lobules mostly compressed in same plane; 2 x 2.5 x 1-2 cm. Smaller Remarks. This species is most similar to Psam- specimens in life dull orange-brown; surface 5 YR tnoclema densum (described below) in external 6/6 for F51991, 10 YR 6/4 for F51992; choano- shape, mode of attachment, surface, consistency colour. Phoriospongia carcinophila is com- some 5 YR5/10 for F51991 . Spongy, soft to moder- and ately firm. monly firmer, has a better organised ectosome and spicules Surface smooth, finely wrinkled (in alcohol); peripheral choanosome, including proper microscopically Finely conulose to tuberculate, tips in ascending tracts, which are always lacking in P. Phoriospongia kirki, 200-400 iim apart, frequently connected by slight densum. See also Remarks on ridges. Fine ostia regularly crowded, more conspic- below. interstices. Oscules of two uous in slightly depressed Phoriospongia kirki (Bowerbank) kinds: small circular ones, about 200 tan wide, fre- 10-12, text-figure 50 quent; larger ones up to 1.5 mm wide, infrequent, Plate 7 figures elongate, irregular in outline. circular or Dysidea Kirkii Bowerbank, 1841c: 161 (Wollongong, chie- Armoured ectosome around 100 /im thick, NSW). -Bowerbank, 1844b: 68, pi. 6 fig. 9.- fly of spicule fragments, with fine lilhic detritus Bowerbank, 1864: 212. -Carter, 1881a: 374 (South - interspersed. Choanosome dense, brown in trans- Coast). -Carter, 1885b: 216 (Port Phillip Heads, Vic). mitted light, granular and fibrillate. Detritus in Lendenfeld, 1885a: 23. Hyatt, 1877: 539, pi. 17 fig. 19 choanosome coarser than at surface, ill-sorted, of ISpongelia Kirkii.- (Phillip Island, Vic). foreign spicules, their fragments, lithic grains in Chondropsis kirkii. -Dendy, 1895: 251 (Port Phillip about equal proportions; mostly in meandering, Heads and Sorrento Jetty, Vic).- Whitelegge, 1901: 77 ascending tracts, without visible spongin, chiefly (Tuggerah Beach, NSW).- Whitelegge, 1906: 465 (off with few often in fascicles at variable intervals, Wollongong, off Narrabine, NSW). -Dendy, 1916b: 127 indistinct anastomoses and branches. No coinci- (Okha/Beyt, Kathiawar Peninsula, India). -Dendy, 1924: dence with tubercles at surface. Tracts more irregu- 363 (Spirits Bay, North Cape, New Zealand). — Shaw, Tasmania). -Burton, 1938: 20 lar, angular, and tighter in F51992. Proper spicules 1927a: 433 (Maria Island, choanosome (Maria Island, Tasmania). (more abundant in F5 1 99 1 ) in deeper Psammascus kirkii. -de Laubenfels, 1936a: 99 (trans- concentrated in and around tracts of debris. Few- fer only). short tracts of subparallel spicules interstitially, Dysidea favosa Marshall, 1880: 98, pi. 6 figs. 6-11 (Bass more frequent in peripheral choanosome. F51990 Strait).- Ridley, 1884a: 388 (Darwin, NT). somewhat different from other specimens. Purplish yellow-grey brown (5 R 3/3 to 7.5 R 3/4) in life, than in alchol (5 Y 6/4). Softer, more easily torn smaller specimens. Relief at surface, reticulation of ridges, more pronounced and coarser. Ostia inconspicuous. Small oscules in interstitial depres- Phoriospongia kirki, spicules (strongyles), from sions frequent, larger oscules (around 2 mm wide) Figure 50. F51993, x.366. conspicuous. 74 I WIEDENMAYfR

Walichondria mammillala Lendenfcld, 1888: 81 (pari, scarce or absent, strongyles occasionally also rare Port Jackson). -Whitelegge, 1902b: 277, 282 (revi- or wanting. sion). -Hallman, 1914b: 340 (revision). Description. F51993 (pi. 7 fig. 12) with rounded Sigmatelta australis var. tubaria Lendenfeld, 188.8: 197 lobes and apical oscules like var. mammillaris of (Port Jackson). - Lendenfeld, 1889b: 613, pi. 40 fig. 4, and F51995 like var. flabel- pi. 42 figs 4, 5. -Whitelegge, 1889: 185 (Shark Reef, Port Lendenfeld. F5I994 Jackson). -Guiler, 1950: 10(Blackmans Bay, Tasmania). lum Lendenfeld, the former a single thick lamella, Sigmatelta auslralis var. flabellum Lendenfeld, 1888: somewhat convoluted, with wide, truncate to 198 (Port Phillip Bay, Vic; Port Jackson). -Lendenfeld, slightly convex margin; the latter composite, with Whitelegge, 1889: 1889b: 614, pi. 41 figs 6, 9, 10, !L- two lateral, partly concrescent lamellae perpendic- (off Green Point, Port Jackson). 185 ular to main lamella. F51993 and F51995 yellow- S/gma/e//aco/7/a7/a LendenTeld, 1888: 199 (collective ish brown in life; F51994 yellow to pale orange. name, no nominotypical subspecies). — Whitelegge, 1889: Firm, crumbly as in stale bread, slightly compres- 185 (off Shark Point, Port Jackson). -Guiler, 1950 Sur- (D'Entrecasteaux Channel and Blackmans Bay, sible to rigid, conspicuously sandy and slimy. Tasmania). face thrown into polygonal meshwork of ridges and

Sigmatelta corticata var. mammillaris Lendenfeld, depressions. Meshes around 1 mm wide, commonly 1888: 199. -Lendenfeld, 1889b: 622, pi. 40 fig. 5, pi. 41 elongated radially on sides of flabellate specimens. fig. 4 (Port Phillip Bay, Vic). Ridges about 0.5 mm wide. Areas around base in Sigmatelta corticata var. papillosa Lendenfeld, 1888: F51993, and around oscules in all specimens, (St Vincent Gulf, SA; Tas.; Port Phillip Bay and 201 smooth, covered with armoured ectosome. Westernport Bay, Vic; Port Jackson, NSW).— Oscules up to 6 mm wide, round, with sharp Lendenfeld, 1889b: 620, pi. 40 fig. 2 (part, type series skeletal margin, and iris-like composite). -Whitelegge, 1889: 185 (Maroubra Bay, membrane. Com- NSW). monly smaller, crowded on truncate margins of Sigmatelta corticata var. flabellum Lendenfeld, 1888: lamellae, 3 to 4 across. 203 (Port Phillip Heads and Bay, Vic; Port Jackson, Choanosomal skeleton consists of (1) curved NSW). -Lendenfeld, 1889b: 624, pi. 40 figs 6, 8.- compressed trabeculae packed with sand, around (off Point, Port Jackson). Whitelegge, 1889: 185 Green polygonal alveoles, similar in size to meshwork at Sigmatelta corticata var. serrata Lendenfeld, 1888: 203 surface; (2) interstitially scattered proper spicules. (Port Phillip Bay, Vic; Port Jackson, NSW).- Meshwork interrupted and frequently indistinct, by Lendenfeld, 1889b: 625. irregular exhalant canals of variable diameters, Material examined. Station KG 6, one specimen (F51993); intergrading with alveoles. This pattern again inter- specimens (F5 1994, F51995). station BSS 187, two Type rupted, in mesial and distal portions of flabellate specimens of Dysidea kirki (BMNH). Holotype of specimens (and in axes of lobes of mammillate Dysidea favosa (PMJ). Type specimens of Halichondria specimens), by regular, terete to slightly tapering mammillata, Sigmatelta auslralis varr. tubaria and flabel- atria (see Lendenfeld, 1889b, pi. 40 fig. 6). lum, S. corticata varr. mammillaris, papillosa, flabellum and serrata (AM and BMNH). Hypotypes of Dysidea Spiculation: strongyles only in F51993 and

kirki: Carter, 1 885b (BMNH) and of Chondropsis kirki: F51994; no proper spicules in F51995. Strongyles 1895 (NMV). mostly bent to flexuous, 129-754-172 0.8-1 Dendy, x fim in F51993, 134-/65-191 x 1-1.1 in F51994. Diagnosis. Massive, lobate, mammillate, tubular, /( m

or irregularly and thickly lamellate to flabellate, Remarks. The species is very polymorphous, more with entire or serrated margin, occasionally so than admitted by Lendenfeld and Dendy. This concrescent-ramose, with compressed branches. is demonstrated by the many type specimens and Firm, sandy, compressible to rigid, mostly easily hypotypes examined. Lendenfeld's varietal names broken, crumbly, slimy. Abundance of sand and cover the range of external shapes only inade- average grain size variable. Dull yellow to pale quately, and several specimens seen by me are either orange or yellowish brown in life. Surface regularly intermediates or combine traits of two or more of rugo-reticulate, alveolate, with meshes around 1 his varieties. An extreme phenotype, not covered

mm, partitions 0.5 mm wide. Pattern sometimes in Lendenfeld's descriptions, is represented by one finer, may be veiled by thin armoured ectosome. of Bowerbank's specimens with compressed Oscules conspicuous, apical on lobes, marginal on branches partly fused in a coarse clathrate lamellae, with surrounding surface denser, smooth. arrangement. Choanosome cavernous, skeleton alveolate, with There is no congruence between shape, colour, contorted, compressed trabeculae packed with consistency, skeletal structure, and surface charac- sand, with little .spongin. Proper spicules scattered ters. Consistency varies between rigid and quite interstitially in variable amounts of matrix; thin compressible, as a function of relative abundance small the latter strongyles and sigmata; may be of sand (of variable average grain size), spongin. SPONGES FROM BASS STRAIT 75 and fleshy matrix. Most of Dendy's hypotypes in Loeblich and Tappan, 1964: C792 (under Melbourne (all wet) are compressible, with a fleshy Xenophyophorida, wrong type species). 214. -de Laubenfels, choanosome, and a distinct, continuous ectosome, Sarcocornea Carter, 1885b: 1936a: 30 (as synonym of Dysidea), — Bergquist, 1980b: in which the sandy armour is reticulate, in petaloid 480 (as synonym of Dysidea). fashion with clustered ostia in the slightly recessed membranous interstices. This condition cor- Diagnosis. Stylotellinae without proper spicules. responds to that illustrated by Lendenfeld (1889b, Foreign debris commonly abundant in choanosome only pi. 42 figs 4, 5) for Sigmatella australis. and ectosome. Distinct from Phoriosponia by The holotype of Dysidea favosa (pi. 7 fig. 11) the consistent absence of proper spicules. external shape, has is of the "australis-type" in Remarks. Marshall distinguished Psammoclema exceptionally pliable and regularly reticulate fibres, and Psammopemma on the grounds of differences no trace of an ectosome, and a very scanty matrix in shape (ramose versus massive), surface (detach- have been partly macerated when (but may able ectosorne/no ectosome), aquiferous system collected). (conspicuous, geometrical/lipostomous and remarked on the absence of sig- Dendy (1895) "lipogastric"), and skeletal structure (dendritic specimens. It is here surmised mata in some of his fibres/amorphously packed sand). Most of the spe- that in specimens like the new ones, with much cies here included in Psammoclema would be also megascleres may be virtually or coarse sand, impossible to distribute among two genera using supressed, in analogy to Echinoclathria completely Marshall's criteria, and introduction of new genera laminaefavosa and E. globosa (see above). in this heterogeneous group is undesirable. Sar- other better known species of Of the common, cocornea nodosa Carter, the type species, with its Australia, which are Phoriospongia in southern heavily armoured ectosome, lack of discrete fibres, to lobate, with finely uniform sur- chiefly massive and dense choanosome, clearly does not fit into is consistently tuberose or face, P. argentea Dysidea and falls easily within this group. smooth, cushion-shaped, fleshier, its surface always Psammascus could be regarded as a senior syn- choanosome dense, foreign with thicker ectosome, onym (by page priority) on the basis of the origi- disposition. P. car- debris not in reticulate/alveolar nal description. The misinterpretation by de also lacks the reticu- cinophila is generally softer, Laubenfels and by Vacelet et al. has already been the least late arrangement of debris, being noted above (Remarks on Desmacididae). Accord- species in this respect. Both organised of the three ing to Marshall's description, the monotypic genus P. carcinophila do not possess sig- P. argentea and is distinct by its tubular shape, by its discrete fibre carcinophila, above. mata. See also Remarks on P. net (anastomosing, regularly ascending primaries) is P. Yet another congener close to P. kirki charged with foreign debris, by soft, limp con- also be lamella Lendenfeld (1888). Though it may sistency, and by absence of proper spicules. Tubu- is thickly erect- irregularly massive, it commonly lar shape is not uncommon in Phoriospongia [P. lighter lamellate. It differs from P. kirki by its confoederala (Lamarck), P. syringiana straight, more colour (light grey, almost white), its (Whitelegge)], and is thus compatible with Psam- two robust strongyles (2-3 ftm thick), and by having moclema sensu novo. On the other hand, Psam- again from sizes of sigmata. It has been recorded mascus decipiens might be a Dysidea. The holotype from New South Wales by Whitelegge (1906: 465), seems to be lost (I could not find it among Mar- from Port Phillip Heads by Dendy (1895: 253), and shall's material of 1880 in PMJ and ZMB). Psam- by Vace- the reef complex of Tulear (Madagascar) mascus is therefore best treated as a nomen let et al. (1976: 63). dubium.

Psammoclema Marshall Psammoclema bitextum sp. nov.

IPsammascus Marshall, 1880: 92. -nee sensu de Plate 7 figure 13, plate 27 figure 3 Laubenfels, 1936a: 96. -nee sensu Vacelet, Vasseur and Holotype: F51996, station BSS 179 (39°03.2'S, Levi, 1976: 63. 14639.5'E, L of Wilsons Promontory; depth 55 m, Psammoclema Marshall, 1880: 109.-Lendcnleld, Lauben- muddy sand). 1889b: 124 (as synonym of Chalinopsilla). -de of Daclylia). fels 1936a: 59 (as synonym Diagnosis. Few basally united branches, partly Marshall, 1880: 113. -Dendy, 1905: Psammopemma repent, partly erect. Branches crooked, terete to -de Laubenfels, 1936a: 96 (discussion), 210 (definition), angular, 5-10 mm thick. Softly spongy, elastic; dull Loeblich and 99 (definition in Psammascidae). -nee sensu greyish yellow, partly purplish in life. Surface Tappan, 1964: C794 (under Xenophyophorida). smooth, hispid on branch tips. Oscules inconspic- Holopsamma Carter, 1885b: 211 (part). -nee sensu 76 F. WIEDENMAYhR

with slightly uous, stellate/petaloid, scattered, numerous. usually with volcano-shaped tips 0.5-1 wide. Fragments Armoured ectosome thin, discontinuous. Main recessed minute oscules, mm free individual lobes up to 1 skeleton twofold: (1) vague columns of detritus, 2 to 4 cm wide, with 3-5 thick. Delicate branching (2) irregular net of clear spongin fibres. cm high and mm algae and bryozoans partly overgrown by sponge. Description. Ramose, branches partly repent, In life and in alcohol greyish beige (10 YR7/2-4). partly erect. Longer branches crooked, round to Leathery, rough and gritty. Characteristic, strong angular in cross-section. Branches 5 to 10 mm and persistent odour in preserved specimens, pos- thick. Two repent branches 2 and 3 cm long. Erect sibly due to microsymbionts. branches 1 to 2 cm high. Specimen 3.5 X 6.5 cm Surface regularly thrown into tubercles, around greyish yellow in life (2.5 wide, 2.5 cm high. Dull 1-3 partly con- I mm high wide, mm apart. Some Y 5-6/2-4) with faint purple tinge in places. fluent and elongate, forming short ridges. Regu- Greyish yellow retained in alcohol. Softly spongy, larly packed ill-sorted sand everywhere, with fine very compressible and elastic. interstitial pores, mostly in rows, often forming Surface smooth to unaided eye, branch tips finely reticulate pattern. hispid. Oscules stellate or petaloid, 2 to 3 mm wide, Ectosome 175-860 /

Remarks. The new species is comparable to Psain- moclema ramosum in gross shape, but differs in Remarks. A regularly and conspicuously tubercu- its small stellate/petaloid oscules, and in the late surface, without relation between tubercles and presence of an auxiliary reticulate skeleton of clear skeletal fibres, also occurs in Phoriospongia car- spongin fibres. Psammoclema goniodes is also cinophila, Phoriospongia wilsoni Dendy, and is distinct ramose, but by its angular reticulation Psammoclema nodosum. The latter is sufficiently of fibres packed with detritus. See Remarks on P. distinct from Psarnmoclama callosum by its larger goniodes, below. size, massive shape, tough consistency, darker Psammoclema callosum (Marshall) colour, coarser tubercles, much thicker ectosome, and poorly organised internal sand-skeleton. Plate 8 figure 27 1, plate figure 4 There is a virtually unknown Australian species of Phoriospongia with relatively coarse surface- Dysidea callosa Marshall, 1880: 104, pi. 6 fig. 12, pi. 7 figs 1-5 (probably Australia, Bass Strait?). tubercles and a characteristic ectosome with very conspicuous skeletal ostia. It comprises some of the Material examined. Station KG 6, one specimen in 3 frag- syntypes of Sigmatella corticata var. papillosa Len- ments (F51997). (Type material not found in PMJ and denfeld ZMB.) (1889b, including the specimen figured on pi. 40 fig. 2). This species is also represented by Diagnosis. Small, compressed-lobate to contorted- two specimens (BMNH 1886.12.15.339 and frondose. Surface tuberculate. Leathery, but easily 1886.12.15.340) with manuscript names by Carter torn, greyish beige in life. Strong odour. Ectosome and Lendenfeld. rough, firm packed with sand, 175-860 /tm thick. Oscules small, recessed, on top of conical eleva- Psammoclema densum (Marshall) tions. Main skeleton an irregular, lax reticulation Plate figures of straggling, knotty, clear spongin fibres with ill 8 2-4, plate 27 figure 5 sorted detritus. Choanosome cavernous. Debris Psammopemma densum Marshall, 1880: 113, pi. 8 figs scarce in dense matrix, which is packed with 6-11 (Tas.).-Polejaeff, 1884b: 46, pi. 3. figs 3, 4 (Port bluegreen microsymbionts. Jackson, NSW). -Lendenfeld, 1885a: 23. -Lendenfeld, 1889b: 640, pi. 37 fig. 4; pi. 41 fig. 17. Description. Convoluted fronds or partly concres- tPsammopemma densum var. subfibrosa and tapering Ridley, cent, compressed lobes, the latter 1884a: 390, pi. 41 fig. h (Thursday Island, Torres Strait'). SPONGES FROM BASS STRAIT 77

Holopsamma laevis Carter, 1885b: 212 (Port Phillip tions of lobes. Ectosome contains mostly packed Heads). debris of spicules. Crowded debris in choanosome Material examined. Station BSS 179, one specimen with little organisation, ill-sorted and heterogene- (F51998); station BSS 181, three specimens (F51999, ous. Some less obstructed areas show threefold F520OO, F52001). Holotype of Psammopemma densum structure: (1) lax, irregular reticulation of straggling (ZMB). Type specimens of Holopsamma laevis (BMNH). tracts with mostly fine detritus, chiefly spicule- Diagnosis. Commonly small, incrusting, irregularly fragments; (2) vague reticulation of brown, finely lobulate to concrescent-ramose, often overgrowing granular organic matter; (3) interstitial ill-sorted other objects, coarse detritus. Moderately compres- detritus, chiefly shelly and lithic. Clear interstitial sible, easily torn. Greyish yellow to olive-brown in choanosome packed by globular, mulberry-shaped

life. Oscules elongate, sinuous, variable in size and bodies, around 13 ^m in diameter, presumably distribution. Surface almost smooth to variably ver- microsymbionts.

rucose, rugo-reticulate or rugo-vermiculate. Ill- Remarks. Of all Australian species here assigned sorted detritus mostly crowded throughout, poorly to Psammoclema, P. densum is the poorest in mor- organised in choansome, packed in little differen- phological definition, appearing almost featureless tiated ectosome. to cursory examination. It has also the most fragile Description. F51999 and F52001 small, the former consistency. See Remarks on Phoriospongia car- incrusting, the latter branching, originally attached cinophila and P. kirki (above). to chaetopterid worm tubes with various other Psammoclema fissuratum sp. nov. sponges. F52000 (pi. 8 fig. 3) irregular, with massive, somewhat depressed and lobose base, crooked Plate 8 figure 5 branches, short simple, others some and compo- Holotype: F52002, Station BSS 187 (38°32.0'S, site, bushy or flabellate, with largely concrescent I42"28.6'E, 17 km S of Warrnambool; depth 52 m, coarse branchlets. Basal mass 2x3x1 cm; single sand and shells). branches 5 mm long, 2-3 mm wide; composite Diagnosis. Complex-frondose. Fronds equally thick branches up to 2.5 cm long and 12 mm wide. (5 mm), budding laterally at right angle, regularly F51998 (pi. 8 fig. 2) massive, thrown into con- expanding, bifid to polyfid (tending to dichotomy), torted, mostly compressed and concrescent lobes. with distal incisions of variable depths and widths. Specimen 9x5x4 cm, overgrows bivalve shells, Firm, more compressible and resilient distally. ' fragments of bryozoans and large foraminifera, Olive-brown, distally reddish in life. Oscules small, and bears delicate branching algae, bryozoans and marginal on edges and tips. Surface smooth on hydroids. Largest specimen in life greyish yellow faces, frayed (micropapillate) on tips. Ectosome (5 Y 6-7/4) on surface, dull yellow (2.5 Y 7/8) thin, with regular detritus. Sand occupies half of i inside, largely preserved in 70% alcohol (10 YR choanosome in volume, in vague columns and i 6/2-4). F52000 olive-brown (2.5 Y 6/6), pale beige enclaves agglutinated by clear spongin, rarely scat- in alcohol. Firm, moderately compressible and tered. Isolated grains often connected by trellised resilient, easily torn, as in wet cardboard. thin spongin fibres. Oscules in largest specimen marginal on lobes, mostly elongate, sinuous, with rounded margins, Description. Four distally incised spatulate fronds, all in one plane, three being parallel out of edge about I mm wide and up to 4 mm long. Oscules in F52000 smaller, round, irregularly scattered in of fourth frond. Transverse frond 2.5 cm long, 2 pits and clefts. cm wide, divided by two incisions, the lower dis- Surface almost smooth in some areas (particu- tinct, the upper a notch. First lateral frond at base larly of largest specimen) but commonly tubercu- of transverse frond, stubby and bifid. Middle one late and rugose. Pattern mostly fine (rugae and 2.5 cm long, 1.8 cm wide, bifid, with incision 1 cm tubercles 0.5 mm wide), coarse and more irregular deep. Tallest frond 3.5 cm high, originally dichoto- sectioning), with in some places. Where relief most regular, partic- mous (one part cut off for median shallow lateral incisions, ularly in F52000, tubercles joined by low ridges in incision 1.5 cm deep, two mostly clathrate or polygonal pattern, with interstitial pits. outermost lobe truncate. Specimen 5 mm life, pale yellowish More often, meandering pattern of ridges and fur- thick. Olive-brown in now darker, reddish (2.5 YR 5-6/4) rows. The latter contain membrane with reticula- brown (10 YR 7/4), tall lateral lobes. Firm, little com- tion of fine sand (spicule fragments), and interstitial in distal parts of moderately resilient distally. ostia; pattern stands out where depressions wider, pressible in middle, along edges of fronds roofing regular square crypts, 0.5-1 mm wide. Oscules regularly arranged sharp-rimmed, Choanosome dense, rnicrocavernous, except and distal lobes, round or elongate, compressed atria branching inward in mesial por- 0.5-1 mm wide. 78 F. WIHDFNMAYER

resilient. Surface optically Surface smooth to unaided eye, except finely compressible and microconulose, more so on branch pitted and frazzled tips of taller lateral fronds. smooth, but obscured by sand on sur- Well-sorted sand grains conspicuous, crowded, tips. Conules somewhat conspicuous, round, 0.5-1 mm wide, with interstitial oslia not in distinct pattern. face. Oseules branches and on Armoured ectosome frequently only 1-2 sand regularly scattered along edges of frondose base. Paratype (pi. 8, fig. 6) grains thick, with much spicular detritus, chiefly one side of lobose base surmounted by two on surface. One-half of volume of choanosome 9x17 mm, has all portions compressed in one occupied by sand grains (including calcareous shell- tapering branches, in plane of compression. debris), with few foreign spicules and their frag- plane. One branch looped chaetopterid worm tube with ments, mostly packed in vague columns or irregu- Originally attached to life not recorded, now lar masses, rarely scattered. Mean grain size 175 other sponges. Colour in limp. Surface finely (im. Grains agglutinated by colourless, barely visi- cream. Softer than holotype, ble spongin; thin clear spongin fibres with irregu- eonulose, without conspicuous sand. Outermost pattern of ridges lar, trellis-like meshwork between neighbouring transverse fibres form reticulate grains. between conules. Recessed fleshy interstices with Choanocyte chambers spherical to oblong. Typi- fine secondary reticulum of spicule fragments, cal sizes 64 x 39, 38 x 29, 47 /un. Choanosome single ostia in meshes. contains scattered spherical bodies, like mulberries, Main skeleton in holotype an angular and irregu- 10-/2-15 //m in diameter, presumably lar reticulation of fibres, with little orientation and microsymbionts. distinction by thickness, packed with detritus, mostly spicule fragments; few fibres with about Remarks. Psaminoclenia fissurutum is sufficiently equal proportions of other fragments. Very few distinct from congeners by its shape, in combina- interstitial inclusions. Most spicule fragments lon- tion with the smooth, evenly microgranulate sur- gitudinally arranged, but many oriented at random. face, and the regular marginal oseules. See also Pale clear spongin agglutinating and enveloping Remarks on Psammoclema goniodeS and Psam- detritus. Thickness of fibres 23 to 68 /tm, mostly inoclenia stipitatum, below. around 40 /tm. Meshes triangular, rhombic and Psammoclema goniodes sp. nov. irregularly polygonal, rarely square, 23 to 150jttm wide. Peripheral portions with indistinct radial Plate 8 figures 6, 7, plate 28 figures I, 2 primaries and secondaries, like in irregular spider Holotype: F52060, station BSS 187 (38 D 32.0N, web. Towards surface, meshes smaller, more regu- 142"28.6'H, 17 km S of Wannambool; depth 52 in, coarse lar, less angular. Here, between choanosome and sand and shells). Paratype: F52059, station BSS 181 armoured ectosome, fibres packed by ill-sorted D <38"39.X'S, 144 18.2't:, 30 km SE of Ionic; depth 79 m, detritus, also interstitially. Spicular and other (lilhic fine sand with cliaeiopterid worm tubes). and shelly) components in about equal proportions, Diagnosis. Sparsely branched, base compressed- both interstitially and in fibres. Abundance of lobale, may be stipitate. Branches crooked, slightly coarse grains higher at surface, armoured ectosome compressed and tapering. Small specimens soft, not distinct. Matrix of choanosome scanty, around limp, with eonulose surface. Larger ones moder- irregular lacunae. Macrosymbionts of one kind. ately resilient, mostly smooth, distally finely eonu- Fibres of paratype thicker than in holotype, more 0.5- lose. Oseules numerous, round, 1 mm wide, continuous, more straggling than angular. Packed scattered on face of base, mostly lateral on detritus comprises more non-spicular components, branches. Armoured ectosome thin, indistinct. spicular fragments more often disoriented. More Main skeleton thoroughly angular reticulation of interstitially scattered detritus, poorly sorted. In straggling spongin fibres packed with well sorted periphery, radial and tangential fibres more distinct detritus, including many spicule fragments. than in holotype, primaries forming axes of conules and often curved in tips. No distinct armoured Description. Hololype (pi. 8 fig. 7) comprises two ectosome, only concentration of well sorted spic- tapering branches, strongly bent in same direction ular fragments between fibres. Density of in upper portions, 35 and 50 mm long, 5 and 7 mm choanosomal matrix higher than in holotype. wide, slightly compressed. Branches arise from stipitate, frondose base 25 mm high and up to 13 Remarks. The proximal part in the larger specimen nun wide in upper portion. Two short lobes resembles individual fronds of Psammoclema fis- (aborted branches) issue from one side below suratum (described above). Fresh material of both shorter main branch. Orange-brown in life (2.5 YR species might reveal closer external similarity of 5/6-8), now faded (5 YR 6-8/4). Firm, moderately whole specimens. The differences in structure of SPONGES FROM BASS STRAIT 79

the main skeleton (irregular distribution of sand flush or elevated. Armoured ectosome a stratified, internally, and anastomoses of clear spongin fibres collagenous cortex around 1 mm thick, with regu- between grains in P. fissuratum) are sufficient to lar, numerous erect ectochones underlain by distinguish the two species. The skeletal structure intracortieal crypts. Stratification also by sorting of P. goniodes (angular reticulation of sandy fibres) of detritus. Choanosome beige to dull ochre in life, is also different from that of two other ramose con- dense, microscopically stringy (strands of algal geners. The main skeleton of P. ramosum consists microsymbionts). Internal detritus ill sorted, in of very long, sparsely anastomosed fibres, longitu- vague, wide, dendritic and meandering tracts, some dinal in the core, curving outwards and becoming interstitially. radial below the surface. P. hitextum has a double texture of straggling columns of detritus and of Description. Semi-encrusting on perpendicular rock reticulate clear spongin fibres. face, outline round above, sinuous below. Now shrivelled and distorted preservation. Apical Psammoclema nodosum (Carter) by volcano-like group of three concrescent oscular Plate 8 figures 8-12, plate 27 figure 6, cones. Width 1 1 cm, height 10 cm, maximum thickness 4 cm. Compound oscular cone now com- text-figure 51 pressed, 5.5 cm wide basally, 2.3 cm high. Sarcocornea nodosa Carter. 1885b: 214 (Pori Phillip Externally dark smoky grey above, light grey Heads, Vic). below, now generally more brownish (5 YR 4-5/1 Spongelia nodosa. - Lendenfeld, lS89b: 670, above, 10 YR 7-8/2-4 below). Choanosome beige Dysidea nodosa. — Bergquist, 1980b: 482 (unrecognis- to dull ochre (10 YR 8/2 to 5 YR 6/6). Very firm able except as Dysidea). and tough, moderately compressible, little resilient. Material examined. Station KG 2, one specimen (F521X13). Surface bears strong and regular brain-like pat- Type specimens of Sarcocornea nodosa (BMNH), tern, now much contracted, more pronounced than

Diagnosis. Large, irregularly massive, heavy. Very in life. Tubercles elongate, confluent in meander- firm, tough. Surface tightly tuberculate (nodose) ing groups occasionally branching, mostly 2-3 mm to coarsely rugose, or brain-like; relief sometimes wide, up to 2-3 cm long, separated by dendritic and vague, or accentuated by superimposed coarser labyrinthic grooves 0.5-1 mm wide. Pattern weaker folds and meandering trenches (the whole reminis- and mostly radial on oscular cone. Central portion cent of pachyderm skin). Grey, often brownish, near base with pattern of contorted cock's combs. dark above and on exposed sides. Oscules conspic- Surface almost smooth, finely hispid in parts, uous, marginal, irregularly scattered or clustered. regularly scattered ostia everywhere, 25 /tin wide, spaced around 125 /im. Distinct, stratified cortex. total thickness 750-1250 fim. Outer cortex three- quarters of thickness, traversed by regularly spaced erect ectochones to often continuous intracortieal crypts (text-fig. 51). Outer cortex mostly packed with ill-sorted detritus, about equal parts of spicular and non-spicular components, except surface, where spicular fragments dominate, often erect, producing faint hispidation. Inner cortex more collagenous, with better sorted detritus (spicular fragments often dominating) loosely scattered to crowded, extended inward in linings of large exhalant canals. These are 1250 to 2500 /tm wide, irregular, often transversely folded. Walls with irregularly scattered mouths of finer canals, around 125 /.un in diameter. Main skeleton of moderately abundant, ill-sorted foreign detritus, in irregular fibres, or complex fenestrate to trellised systems, and interstitially.

Figure 51. Psammoclema nodosum, perpendicular sec- Matrix of choansome dense, stringy, permeated tion in reflected light, x4.l, showing cortex (with sur- throughout by wavy strands of filamentous algae. face in perspective) with erect ectochones and layer oi Remarks. Psammoclema nodosum is as close to P. intraocortical crypts. A large, much twisted cxhalant canal vansoesti (described below) as it is different from is visible from lower centre to upper left. 80 F. WIEDENMAYFR

all other congeners. In common with the former wide, irregularly rounded, now covered by pellucid is the dark colour of the surface, the firm, tough, membranes. Sand in inner, concave faces packed dense and heavy consistency, the well organised, only in ridges, elsewhere crowded to clustered.

thick collagenous ectosome (reminiscent of the Interstices, up to I mm wide, and sand grains cortex in Astrophorida), and the dense choanosome covered by pellucid dermis. Fine reticulum with containing ill-sorted detritus in vague meandering interstitial ostia over subdermal crypts in wider tracts. P. vansoesti is chiefly distinct by its vase-or interstices. Choanosome packed to crowded, bowl-shape, by the coarsely rugo-retieulate/alveo- without order, by sand as at surface. Thin super- late pattern on the sides or on the rim only, and ficial coat of foreign spicules, forming dermal retic- by the numerous minute oscules regularly dis- ulum. Most spicules entire megascleres. No tributed in the apical concavity. spongin. Choanocyte chambers regularly oblong to spherical, typical dimensions 63 X 52, 65 X 44, Psaminoclema radialum sp. nov. 74 x 64 ixm.

Plate 9 figures , 2 1 Remarks. The capricious shape, thin dermis with Hololype: F52004, slation BSS 187 (38"32.0'S, mostly whole foreign spicules, and the abundance 142°28.6'E, 17 km S of Warrnambool; depth 52 m, coarse of coarse sand without internal order (hence crum-

sand and shells). bly consistency) distinguish this species from its congeners and all other thoroughly sandy species. Diagnosis. Several half-bowl-or ear-shaped, partly concrescent lobes jutting from common lacunose Psammoclema ramosum Marshall base, distally folded radially, marginally blunt, scal- Plate loped. Very firm, slightly resilient, crumbly, 9 figures 3-6, plate 28 figures 3, 4 thoroughly and conspicuously sandy. Dull orange- Psammoclema ramosum Marshall. 1880: 109, pi. 7 figs brown in life. Thin pellucid dermis reinforced by 12-15; pi. 8 figs 1-5 (Bass Snail I. -Polejaetf, 1884b: 43, reticulum of foreign spicules fig. (few broken). Well pi, 3 8, pi. 4 fig. 1 (off Moneoeur Island. Bass Strait, sorted coarse sand grains crowded to packed inter- 70 m). nally without order. Cliti/ino/jsil/a arborea var. ramosu. -Lendcn(e\d, 1888: 112 (old record).- I.endenfeld, 1889b: 152 (old Description. Capricious in shape, like baroque record). -Guiler, 1950: 6 (Blackmails Bay, Tas.; record fountain, growing out from perpendicular rock only). face. Grossly lacunose base with 5 bowl-or ear- Material examined. Station KG 7, one specimen (F52005). shaped protuberances in three directions, on differ- Hololype of Psammoclema ramosum (PMJ). Hypolype ent levels, roughly in same plane, but somewhat of Psammoclema ramosum: Polejaeff, 1884b (BMNH). tilted and contorted. Upper faces deeply hollow to Diagnosis. Prolific long lacunose proximally, traversed by annular or trans- branches from common base, often verse ridges distally. Margins narrowly rounded, repent throughout, concrescent laterally and sinuous, occasionally disrupted by small hollows dorso-ventrally. Branches commonly curved to bent, or notches. Lower convex faces distally scalloped, slightly depressed and nodose, 0.6-1.2 cm wide. Firm, mostly by radial coarse ridges and grooves, the moderately compressible, easily torn. Dark purplish former 2-3 mm, the latter 3-5 mm wide. In por- or reddish grey. Surface smooth to finely tions of uppermost bowl, and on convex face of rugo-reticulate. Oscules numerous, conspicuous, irregularly lower protuberances, rugose pattern meandrine to rounded, often elongate axially, mostly on reticulate, occasionally finer. Specimen 8 cm in upper faces only, often on top of nodes. width, 7 cm in depth, 6.5 cm in height. Largest Armoured ectosome thin. Main skeleton of crowded, excrescence on top 7 cm wide, 5 cm deep, 2-2.5 straight, dendritic fibres, longitudinal in core of branches, curving cm high. Lower ones 1 .5 to 4 cm in width, 1 .5 to outwards in periphery, becoming radial 3 cm in depth; thickness 1 cm in middle, 2-3 mm below surface, rarelv anastomosing. at margins. Dull orange-brown in life (7.5 YR 6/4), slightly Description. Aggregate of repent, depressed more yellow (10 YR 6/4) in alcohol. Very firm, branches. Three main branches issueing from barely compressible and resilient, crumbly, conspic- common base, each repeatedly subdivided (more uously sandy. frequently below), partly concrescent along sides Outer convex surfaces exhalant, packed with and on top of each other. Three minor branches rounded lithic and shelly grains, 150-800 /mi, from base, one short and recurved, middle one mostly around 350-500 pm, with numerous oscules looped and partly concrescent with base, third free, scattered in depressions. Oscules mostly 1-2 mm with stubby secondary branch near base. This and SPONGES FROM BASS STRAIT 81

main branch gently s-curved. Main branches 5, 8, margin. Firm, tough, moderately resilient. Olive and 7 cm long, 1 to 1.2 cm wide, around 8 mm to purplish brown, mottled. Surface transversely thick. Free minor branch 7 cm long, 5x8 mm rugose on stalk, rugo-reticulate or rugo-vermiculate thick. Length of specimen 14 cm, width at base 4 on flabellum. Rugae and pits 0.5-2 mm wide. cm. Oscules small, inconspicuous, marginal on flabel- Dark grey, in life, with purple tinge above (5-7.5 lum. Armoured ectosome 0.7-1.3 mm thick. Main RP 1/5-6), dull purple to reddish below (2.5 R 2/7 skeleton an irregular reticulation of spongin fibres to 10 R 7/6) now faded to beige, with some greyish enveloping single and contiguous sand grains; hence areas (2.5 Y 8-9/2-4 and 10 YR 6-7/2). Firm, most fibres knotty, lacunose. Much spicular detri- moderately compressible and resilient, easily torn. tus scattered interstitially. Filamentous microsym- Oscules numerous, irregularly scattered on upper bionts throughout choanosome. face of base and of main branches, rare on sides Description. Specimen stipitate and distally flabel- and below, irregular in outline, especially larger late. Stem basally expanded into rhizome (now ones, elongate in different directions, drop-and eye- incompletely preserved), 4.5 cm high and 7-9 mm shaped. Maximum width 1 to 3 mm. Margins thick in its terete portion, transversely wrinkled. mostly sharp, occasionally slightly recessed. Compressed above, expanding, first to 2.5 cm, then Surface smooth over most of upper faces; finely abruptly into slightly asymmetrical flabellum, eye- pitted, honeycombed or clathrate along sides, lower shaped, 8 cm wide, 10-13 mm thick. Height of face and on branch tips. Ridges outlining pits in specimen 12 cm. these areas mostly finely denticulate. Smooth sur- In life olive on one side, purplish brown mot- face microreticulate, with polygonal osiia tled with olive on other. Shades darker in depres- interstitially. sions, lighter in rugo-reticulation, now faded to Main skeleton chiefly of dendritic fibres, packed greyish beige (7.5 YR 6-7/2-4) and dull red (5 R disorderly by moderately sorted detritus, with spic- 5-6/4 on rugae, 5 R 5/2 in depressions). Firm, ular and non-spicular fragments in equal propor- moderately compressible and resilient, tough. tions. Fibres longitudinal, straight, crowded in axial Relief on both faces of flabellum distinctive. Pat- region, of irregular cross-section, often com- tern on two-thirds of one face a rugo-reticulation pressed, 120 to 280 jim thick; curving outwards in with lighter, sandy rugae, and optically smooth, periphery, reaching surface obliquely or radially, oblong to irregularly sinuous depressions of vari- becoming thinner, straggling, more widely spaced, able size; width of rugae and depressions 0.5-2 mm. branching and anostomosing sparingly. Pale clear In remaining area, depressions vermiculate, den- spongin agglutinating and outlining fibres barely dritic and partly confluent, with rugae discontinu- visible. Some detritus scattered interstitially. Ecto- ous, beset by low conulcs. Pattern finer, relief some armoured to variable degree, chiefly by for- weaker than in rugo-reticulate area. Pattern of eign spicules and their fragments, 65-135 urn thick. other face generally more cerebroid, with narrow Choanosome dense, crowded to packed by globu- vermiculate depressions and relatively wide rugae. lar, mulberry-shaped microsymbionts, about 8 //m Relief strong and coarse in one-third of face, fine in diameter. and rather vague in remaining area. Depressions Remarks. Among the congeners, P. bilextum and contain sieves of very crowded round ostia. Oscules P. goniodes are also ramose. They both differ from small and inconspicuous, now contracted, numer- the present species in having smaller, less conspic- ous in depressions along edge of flabellum. Parallel uous oscules, and in their skeletal structures (see radial atria in marginal portion of flabellum terete, Remarks following the respective descriptions of 1-2 mm wide, 1-2.5 mm apart, linings with regu- the two species). lar annular folds. Ectosome armoured with sand grains and spic- Psammoclema stipitatum sp. nov. ule debris, not stratified, 700 to 1300 /an thick. Main skeleton of sand grains agglutinated and Plate 9 figure 7, plate 28 figure 5 encased by spongin, in irregular reticulation. Hololype: F52006, station BSS 187 (38"32.0'S, Diameters and orientation of fibres extremely vari- 142°28.6'E, 17 km S of'Warrnambool; depth 52 m, coarse able, no distinction of primaries and secondaries. sand and shells). Fibres often lacunose and trellised, some almost partly, or devoid for short Diagnosis. Stalked, distally flabellate, fairly sym- filled with debris, others Spongin often weakly and finely strati- metrical. Stalk tall, basally expanded, terete, 8 mm portions. Choanosome contains scattered thick below; compressed above, passing into flabel- lied, where stained. upper foreign spicules and their fragments, more densely, lum about 1 cm thick, with crescent-shaped .

82 F. WIEDENMAYER

but irregular, sometimes packed, in canal linings. Choanosome Diagnosis. Vase-to bowl-shaped, side radially folded andl permeated by filamentous algae in subparallel and angular, oblique. Outer crossed strands. Chaonocyte chambers indistinct, rugose, or coarsely rugo-reticulate, with polygonall irregularly oblong, 33-58 ^m long. pits and tabulate, conspicuously sandy ridges. The latter pattern may be confined to the rim in some Remarks. The only other species among the specimens. Reddish grey in life. Oscules minute, Stylotellinae, here described, to have flabellate ele- regularly scattered in the finely papillate concavity. ments with marginal oscules, is Psammoclema fis- Otherwise very similar to P. nodosum. suratum. Its body, however, is composite, with individual fronds regularly incised distally, the sur- Description. Holotype (pi. 9 fig. 1 1) an asymmetri- face on faces being smooth. There is some similar- cal vase. Basal face of attachment with sinuous out- ity between the two species in skeletal structure, but line, inclined foreword at 45° to axis. Lower the thin, clear spongin fibres connecting isolated portion, 1-2 cm above base, constricted, with axi- grains in P. fissuratum do not occur in P. ally convoluted pillars and caverns of variable stipitatum. depth. Wall of vase expands slightly to margin Psammoclema vansoesti sp. nov. along back and front, not along sides. Margin mostly rounded, fairly thick, of sinuous outline, Plate 9 figures 11, 12, plate 28 figure 6, generally sloping foreward al 30°-45° to axis. Con- text-figure 52 cavity regularly conical. Base of attachment 7 x 9 cm; constricted lower portion 5.5 x 6 cm; upper Holotype: F52008, station KG 6 (Tasmania, Winter Cove, width across axis diameter) 8 height E side of Deal Island, Kent Group; boulders with algae, (maximum cm; along 9-10 depth 3-6 m). Paratype: F52007, station KG 3 (Tasma- axis cm; thickness of margin 1 .3-2 cm; nia, East Cove, W of jetty of settlement, Deal Island, Kent upper diameter of concavity 5 cm; depth of con- Group; outcrops and boulders with algae). cavity 10 cm at back, 4.5 cm at front.

1 '""»,, '5/ Cr^~> .Mum? M GZ&S. WB& **

v **!fe

"J> . >Ji .

f ™'-- *r'*'&:p.-5k

Figure 52. Psammoclema vansoesti p. nov., perpendicular section in reflected light, X6.fr. The cortical structure is similar to that in P. nodosum. SPONGES FROM BASS STRAIT 83

Greyish red in life, more grey in depressions (2.5 packed sand grains and few foreign spicules and R 4-6/2), more red on rugae and on margin (2.5 their fragments. Interstices often correspond to fine R 6/6-8), more uniform in alcohol (10 R 4-5/2). stratified crypts, lowest layer discontinous between Firmly rubbery, heavy, conspicuously sandy on cortex and choanosome. rugae, margin and on upper concavity. Main skeleton similar to that in Psammoclema Irregular and coarse regoreticulation on much nodosum, but detritus even less sorted and less of the outer (inhalant) surface. Depressions abundant in fibres, with spongin more evident, between rugae mostly concave, 1-2 mm deep, weakly and irregularly stratified. Interstitial detri- vaguely polygonal to sinuous, some confluent in tus chiefly loosely strewn spicule fragments. Cortex

irregularly petaloid patterns, width 1 to 10 mm. contains scattered to crowded algal microsymbi- Surface in depressions generally smooth, like in onts, sinuous fibrils tapering both ways from lower part of outer wall, which may be weakly median inflation, filled with opaque granules, 7 /mi sculptured (verrucose and pitted). Rugae 0.5-3 mm in median thickness, 65 /mi in length. Choanosome wide, tabulate to slightly convex, confluent with contains same type of filamentous algae as Psam- surface of margin in sandy structure and general moclema nodosum, but far less abundantly. level above depressions. Surface of margin mostly Etymology. The new species is dedicated to Dr continuous, except few dark smooth pits. Parts of R.W.M. van Soest (Amsterdam) in appreciation of constricted base and most of sculptured area sup- his important contributions to sponge taxonomy port encrusting and branching algae, hydrozoans and to the knowledge of the West Indian sponge and bryozoans. fauna. Fine, barely visible (10 x), regularly crowded species is ostia everywhere in darker, smooth areas and in Remarks. The only close relative of this depressions. Microscopic relief of conules and Psammoclema nodosum. It differs in its irregularly ridges, passing into fine reticulum of spicule massive, never concave shape, its tightly tubercu- fragments. late to brain-like surface, and its conspicuous, Exhalant concavity smooth to weakly sculptured, irregularly scattered oscules only on uppermost partly encrusted by serpulids, pierced by regularly elevations. scattered oscules, more conspicuous in life, now Mycalidae Lundbeck, 1905, sensu van Soest, 1984 0.2-0.5 m wide, spaced at 3-5 mm. Paratype (pi. 9 fig. 12) a slightly oblique bowl, Mycale Gray with wall partly convoluted axially. Basal pillars [Esperia] Nardo, 1833: col. 522 (junior homonym of mostly continuous up outer wall, irregularly Esperia Huebner, 1825). branching into thin ridges, hence fluted top of outer [Acamas] Duchassaing and Michelotti, 1864: 95 (junior around base extensive, confluent and wall. Caverns homonym of Acamas de Montfort, 1808). surface pat- meandering. Rugo-reticulate and pitted Mycale Gray, 1867a: 533.-Thiele, 1903a: 949 (revi- tern only locally, near base, and along margin. sion! revival of name). - Lundbeck 1905: 23 (definition, Sandy structure of rugae on margin only. Prevail- discussion), 29 (redescription of type species). -Hentschel, dimensions for 112 spe- ing secondary relief on outside verrucose, with pim- 1913: 263, 265 (tables of spicular spe- apart, cies). -Topsent, 1924: 77-118 (revision of European ples about 1 mm wide and high, 1-5 mm 86f. (revision of nominate subgenus).- Van often aligned and passing into serrated ridges. cies), 83, Soest, 1984: 9-14 (definition, revision), 31 (discussion). Concavity regularly papillate, densely to 1-3 cm Aegagropila Gray, 1867a: 533. - Vosmaer and Pekel- below the rim, loosely below. Most papillae bear haring, 1898: 19. -Topsent, 1924: 78, 83 (revision as apical oscules. subgenus of Mycale).- Van Soest, 1984: 12, 16 (as subge- width at Height of paratype 7.5 to 11 cm, nus of Mycale). roughly triangular base 8-12 cm, upper diameter Grapelia Gray, 1867a: 534. -de Laubenfels, 1936a: 213 51f. 12 cm, thickness of margin 8-10 mm, depth of con- (genus dubium).-Vacelet, Levi and Vasseur, 1976: Mycale). cavity 8.5-11.5 cm. (retained as subgenus of 537. -Topsent, 1924: 83f. (revi- Preserved paratype dark reddish grey (10 R CarmiaCray, 1867a: sion as subgenus of Mycale). -dt Laubenfels, 1936a: 118 3-4/2) to dark beige (2.5 Y 7/2) externally, yellow- (definition, revision). -Levi, 1956b: 16 (discussion of ish beige internally (5 Y 8/4). Cortex of smooth lndo-Pacific species, Aegagropila implicitly syn- inhalant areas compound, stratified, 1000 to 1300 onymised).-Levi in Brien et al., 1973: 610 (definition (quarter to third of total) with 24 (as /mi thick. Outer layer as genus in Mycalidae).- Van Soest, 1984: 12, /mi parallel and perpendicular ectochones about 20 subgenus of Mycale). wide and 100 /mi apart (text-fig. 52), packed with Raphioderma Bowerbank in Norman, 1869: 333. 1874a: 235 (objective syn- spicule fragments, mostly without orientation, Raphiodesma Bowerbank, Mycale). locally erect. Inner layer contains scattered to omym of :

84 F. WIEDENMAYER

Esperella Vosmaer, 1885 (in 1887): 353, 354 (nomen Diagnosis. "Mycale without ectosomal skeleton. novum for Esperia Nardo). — Ridley and Dendy, 1887: The choanosomal skeleton consists of a coarse rec- 62. -Dendy, 1905: 159. tangular reticulation of thick spicule tracts, com- Pseudoesperia Carter, I886h: 455, 456. -Hallmann, pletely enveloped in a spongin coat. Interior 1914c: 399 (synonymised with Grapelia, subgenus of cavernous with scanty fleshy parts. Spiculation: Mycale). — c\e Laubenfels, 1936a: 121 (definition, discus- subtylostyles, anisochelae and sigmata." (Van sion). -Vacelet, Vasseurand Levi, 1976: 51 (synonymised with Grapelia), Soest, 1984: 28, for Acamasina). Primary fibres are Arenochalina Lendenfeld, 1887a: 821.-Hallmann, frequently cored by foreign debris. 1912: 252 (type species redescribed, synonymised with A/yca/e). -Hallmann, 1914c: 399-Burton, 1932a: 293 Mycale (Arenochalina) mirabilis Lendenfeld (Kallmann, 1912, confirmed; revised diagnosis errone- Plate 9 figures 8-10, 13, plate 10 figures 1, 2, ous, not new species). -Pulitzer-Finali, 1982b: 102 (discussion). plate 29 figures 1-4, text-figures 53-56 Zygomycale Topsent, 1930b: 431.- [Hallmann, 1914c: 406 (list of species of Mycale Arenochalina mirabilis Lendenfeld, pi. with isochelae, originally 1887a: 821 , 26 included in Zygomycale fig. pi. by Topsent). -Hentschel 191 I 70, 27 fig. 28 (Torres Strait). -Lendenfeld, 1888: 299, 301 (remarks on Mycale spp. with isochelae)). — Levi, 103.- Whitelegge: 1902a: 212, 213 (type slide AM G3528, 1956b: 16 (discussion, review). - Levi in Brien et al., 1973: from nolotype, re-examined; new record from Port Jack- 610 (definition in Mycalidae).- Van Soest, 1984: 16 (as son possibly M. (A.)flammula).— Pulitzer-Finali, 1982b: synonym of subgenus Aegagropila). 100, text-fig. 12 (Great Barrier Reef).

Acamasina dc Laubenfels, 1936a: 1 17 (nomen novum ? Arenochalina mirabilis. — Whitelegge, 1906: 466 (off for Acamas Duchassaing and Michelotti). — Van Soest, Shoalhaven Bight, NSW).-Guiler, 1950: 6 (Seven Mile 1984: 13,28 (discussion, definition, redescription of type Beach, Tas., record only). species). nee Arenochalina mirabilis. — Whitelegge, 1901: 76, pi. Mycalecarmia de Laubenfels, 1936a: 120. — Van Soest, 10 fig. 7 (= Mycale (Arenochalina) flammula). 1984: 10 (discussion). Mycale (Arenochalina) mirabilis. — Hallmann, 1912: 252 (schizotype AM G3538 re-examined, discussed). Diagnosis. "Mycalidae with subtylostylote "'.Cacochalina truncaiella var. laxa Lendenfel, 1887a: megascleres and palmate anisochelae; other 763 (Port Chalmers, New Zealand). -Burton, 1927a: 292 microscleres may include toxa, rhaphides, (holotype redescribed as Mycale sp.). micracanthoxea and palmate isochelae." (Van Esperella spongiosa Dendy, 1896: 16 (Port Phillip Soest, 1984: 9). Heads).

Mycale spongiOSa.— Hentschel, 1913: 263, 265 (No. 1 Remarks. This is an exceptionally large and in synopsis). -Burton, 1928: 1 19 (description, discussion, difficult genus. Various attempts at revision and Mycale fistulata Hentschel with var. macrochela syn- synopsis have been made. A resume of these and onymised). -de Laubenfels, 1936a: 1 17 (duscussion).- a new revision are found in van Soest (1984: 9f.). Thomas, 1973b: 36, pi. 2 fig. 8, pi. 5 fig. 9, pi. 7 fig. Past revisions can be classed in two groups: (1) that, 8 (description, comparative table of spicule measurements; represented by de Laubenfels (1936a) and Levi Mahe, Seychelles). -Van Soest, 1984: 13, 31 (type examined, (1956b, of Indo-Pacific species), which relies on included in subgenus Acamasina). Mycale imperfecta Baer, spiculation alone, and (2) that, represented by Top- 1906: 20 pi. 2, fig. 5; pi. 5, figs 3-8 (Zanzibar). -Hentschel, 1913: sent (1924), which is also based on other traits, 263, 265 (No. 14 in synopsis). notably of skeletal structure. The former approach Mycale fistulata Hentschel. 1911: 292, text-fig. 4 is pragmatic and leads to splitting into artificial (W A). -Hentschel, 1913: 263, 265 (No. 37 in synopsis). genera, the second is more difficult, but phyloge- Mycale fistulata var. macrochela Hentschel, 191 1: 294 netically more sound, and leads to subdivision into (WA). -Hentschel, 1913: 263, 266 (no. 24 in synopsis). subgenera. I have no doubt that the subgenera ?Mycate tylostrongyla Pulitzer-Finali, 1982b: 102, text- advocated by van Soest, based on Topsent's figs 13, 14 (Great Barrier Reef). approach as applied with minor modifications to Material examined. Station KG 1, one specimen (F52014); extra-European, chiefly West-Indian species, will station KG 7, three specimens (F52015, F52016, F52017) eventually be applicable to all Indo-Pacific species Station BSS 127 (40°00'S, 144°22'E, SE of King Island. involved. It so happens that the only 47 m, not species of covered in this report), one specimen. Holo- Mycale in the new identified type oi material clearly falls Arenochalina mirabilis, Torres Strait (BMNH 1886.8.27.587, within the subgenus named Acamasina by van dry, here on pi. 9 fig. 13). 22 wet syntypes of Esperella Soest. The proper name for this subgenus, spongiosa Dendy (NMV). however, for reasons of priority, is Arenochalina. Diagnosis. Variably lobate, massive, contorted, erect or repent, Subgenus Arenochalina Lendenfeld, 1887 sometimes with pedicel. Softly spongy, very compressible. Much slime upon col- Synonym: Acamasina de Laubenfels, 1936 lecttng, with dermis and choanosome more or less SPONGES FROM BASS STRAIT 85

Figure 53. Mycale (Arenochalina) mirabilis, range of shapes (not to scale), a. F52014. b. F52016. c. F52015. d. F52017.

°:>&

o 5%Se%v^#'0 &0 54 55

Figures 54, 55. Mycale (Arenochalina) mirabilis, details from F52014, in reflected light, x6.5. Figure 54. Largely macerated portion, in perpendicular section, showing peripheral skeleton. Figure 55. View of surface, with piercing ends of primary fibres and subdermal cavities partly open and enlarged through shrinkage.

deciduous. Dermis and choanosome mostly yellow- foreign debris. Meshwork in some specimens lax, ish cream to beige in life, with some pink and green- less regular, primaries then with few foreign inclu- ish. Fibres conspicuous, dark red to purple in life. sions. Most fibres contain filamentous algae. Vermiculate subdermal canals; dermis tends to Microscleres usually rare. crack when exposed. Oscules conspicuous, apical variable (text-fig. gener- on lobes, often with much smaller, scattered ones. Description. Shape 53), lobate, more or less massive, often contorted. Surface comulose to bristly, with commonly blunt ally developed. Largest specimen primary fibre tips piercing dermis. Some specimens Pedicel may be three concrescent with coarser, widely spaced fibres, with longer con- (F52015, pi. 9 figs 9-10), with 10 high, 6 x 7 cm wide. ules and tapering fibre tips. Other specimens clubshaped tubes, cm station 127, 9 cm high, 11 x fleshier, with thicker, more continuous dermis, sur- Specimen from BSS face then smoother. Fibrous skeleton mostly regu- 23 cm wide. Softly spongy, very compressible and resilient. lar, meshsize around 1 mm; primaries cored by F. WIEDENMAYER

Fibrous skeleton of finer form regularly reticu- straggling in late. Primaries parallel, straight or axial portion of lobes (longitudinal in linings oh atria), curving radially to surface, reached at righn angles, 200-300 ftm thick, 0.5-1 .5 mm apart. Their free tips (hispid surface) 0.3-0.8 mm long. Secon daries 50-100 ^m thick, 0.5-1.7 mm apart. In finer specimens all primaries contain foreign, detritus of variable distribution and sorting, spicular and non-spicular fragments in about equali proportions, thickly coring fibres, with spongin coating grains thickly and evenly. With poor sorting, fibre contours knotty. Spongin in coats strati- irregularly wavy laminae often tapering and Figure 56. Mycale (Arenocha/ina) mirubilis, spicules, fied, never x260. a. Styles and subtylostyles in F52014. b. Sigmata cut off. Proper megascleres in primaries in F52016. abundant, commonly oriented at random. Secon- daries cored by proper megascleres only, usually Dermis and choanosome mostly yellowish cream parallel, several abreast, a few oblique. At junc- with to beige in life, with obtuse tinges of pinkish purple tions, secondaries expand like trumpet bells, and greenish: in F52014, mostly 2.5 Y 9/4, locally coring megascleres fanned out. Few oblique spic- 5 R to 2.5 Y 9/2; in F52015, 5 YR 8-9/2; in F52016, ules (proper and foreign) protrude from primary mostly 2.5 Y 9/6-8, locally, on surface, tinges of and secondary fibres, commonly enveloped by 5-7.5 RP 9/4. Fibres in life dark red to purple (7.5 spongin. Secondaries occasionally branch and may - 10 RP 6/6- 8 to 2.5 R 4-5/2). Upon removal have free, rounded ends. All fibres contain numer- from sea, flesh contracts, with copious extrusion ous strands of filamentous, segmented algae. of slime; most noticeable in rarer phenolype with Choanosomal matrix in this form (F52015, coarse skeleton (F52016). In form with fine, more F52017) contains numerous (in the former abun- regular skeleton, thin dermis (remaining speci- dant) proper megascleres in confusion. Microscleres mens), process slower, appearing first as cracking very rare: only two weak anisochelae were seen in of dermis, red-purple colour of skeleton strongly one section (F52015). In coarsely textured form contrasting with pale dermis. Little remains of (F52016, pi. 10 figs 1-2), distances between choanosome in specimens not immediately primaries and between secondaries commonly 1-5 preserved, dermis remaining in peals and flakes. mm. Free tapering tips at surface up to 8 mm tall. In alcohol cream to pale beige, with fibres bleached. Primaries 270-430 fim thick, secondaries 55-350 /mi. In specimen from station BSS 127, and in some Scarce foreign inclusions in primaries, many algal types of Esperella spongiosa Dendy, dark greyish filaments in all fibres. Primaries filled with beige to light hazel. megascleres in confusion or in semiplumose Surface in finely textured specimens with scanty arrangement, marginal ones occasionally protrud- flesh, finely conulose to hispid, with tips of prin- ing, encased in spongin, hence thorny contour of cipal fibres generally protruding (piercing dermis) some fibres. Megascleres more common in secon- even in life. In F52016 surface spiny, with promi- daries than in finely textured form, always coated nent tapering tips of primaries. In fresh specimens by clear spongin. Choanosomal matrix with numer- of former kind, mouths of numerous perpendicu- ous megascleres, mostly aligned. Microscleres lar inhalant canals conspicuous through dermis. scarce but more numerous than in other specimens, Pattern locally interrupted by subdermal, tange- sigmata only. As in other two specimens, scattered tially meandering and dendritic canals. Both sorts microsymbionts of mulberry-shape, 12-15 ^.m in about 1 mm wide and quickly exposed after diameter. In this specimen only, ovate, deeply stain- collecting. ing structureless bodies, 10-16 )tm long. Oscules of two sorts: larger ones, 6-13 mm wide, Spiculation: (1) Straight, fusiform subtylostyles on apices of lobes, with sharp fleshy and skeletal with elongate, occasionally subcylindrical heads; rims, leading to axial atria tapering to various greatest width two-thirds below head. In F52016, depths; smaller ones, more numerous and scat- points short, often blunt (strongyloxeote to stron- tered, 3-4 mm wide, with less defined rims, often gylote), occasionally mucronate. Axial canals com- composite (confluent). Either one or both on same monly wide, eroded. Rare anisoxea in F52015. specimen. Individual measurements: SPONGES FROM BASS STRAIT 87

F52014: 160-/79-194 x 2.9-4.8-13 urn. to van Soest (1984: 31) is the abundance of sigmata.

F52015: 164-779-222 x 2-3.7-5.8 /t m. There are two other Australian species in this F520 16: 217-242-268 x 1.9-4.4-1.2 nm. subgenus: (1) M. (A.) flam inula (Lamarck) (see Microscleres: (2) two weak anisochelae in Topsent, 1933: 17), with synonyms Spongia ana-

F52015, length 19.6-23.4 jtm; width, in side view, tipes 1 amarck (see Topsent, 1930a: 35), Arenocha- 5.3-5.0 tan; length of free ends 5.7/6.1 and lina mirabilis: Whitelegge (1901: 76, pi. 10 fig. 6.0/10.0 urn. (3) Sigmata (8 measured in F52016), 7), and Spongelia elegans: Lendcnfeld (1889b: 655, symmetrical or contorted: length 73.8-86.2 /un; part, Australian hypotypes only, including pi. 39 total width 28.5-35.3 /mi; thickness of shaft 2.5-3.6 fig. 2 = BMNH 1886.8.27.215); and (2) M. (A.) /im. pluriloba (Lamarck) (see Topsent, 1930a: 28). Rare Remarks. It could be argued that the coarsely lev microscleres probably occur in both and were tured form belongs to a distinct species or subspe- usually overlooked, with exception of Hallmann cies. Among the 22 syntypes of Esperella spongiosa (1912: 252, footnote), who noted anisochelae in A Dendy deposited in the Museum of Victoria is at mirabilis: Whitelegge. least one specimen with a similarly lax and coarse M. (A) flammula has a distinct shape: pedun- fibrous skeleton, and some intermediate specimens. culate, flabellate with radial distal incisions. M. Many of Dendy's specimens are more fleshy than (A.) pluriloba consists of clustering compressed the new ones, with a thicker, continuous dermis, lobes, issuing and regularly widening from a and have a more irregular fibrous meshwork, with narrow base, dislally truncate and incised.

many free secondaries. I doubt that the fleshy con- Tedaniidac Ridley and Dendy, 1886 sistency is the result of immediate preservation. It Synonym: Myxillidae Topsent, 1928, sensu van Soest, helps to support the view that this is a very poly- 1984. morphic species. Tedania Gray In further support of this assumption are some rafaWfaGray, 1867a: 520. -Wiedenmayer, 1977a: 133 comments by Dr K. Riitzler (in litt., 7 October, (references, status of type species). — Van Soest, 1984: 49 1983) concerning his study of West Indian popu- (definition in Myxillidae). lations of Mycale (Arenochalina) laxissima Duchas- Trachytedania Ridley, 1881: 122. -Topsent, 1928: 53 saing and Michelotti. The species was recently (probably a synonym of Tedania). — Burton, 1932a: 306 redescribed by van Soest (1984: 29, text-fig. 9, pi. (synonymised with Tedania). 3 fig. 1). It is certainly closely related to M. Tedaniopsls Dendy, 1924: 366. -Topsent, 1928: 54.- mirabilis, as his discussion (p. 13, under (4), and Burlon, 1932a: 345 (synonymised with Tedania). -de p. 31, with reference to Mycale spongiosa and M. Laubenfels, 1936a: 54 (definition, revision). -Van Soesl, Hemiledania). fislulata) makes clear. While van Soest did not 1984: 53 (possibly to Paratedania Burton, 1929a: 441. -Burton 1932a: 306, speak of polymorphism in this context, Riitzler's 345 (discussion, synonymised with Tedania). -Vim Soesl remarks were very suggestive of wide phenotypic 1984: 54 (type species, misspell tarantula, possibly to manifestation. Hemiledania). Riitzler's spicule measurements (in litt., for speci- Diagnosis. "[Tcdaniidae] with onychactcs as mens from Colombia to Florida) show considera- microscleres (but without chelae or sigmata); ble variability (megascleres being generally larger ectosomal tylotes are terminally mierospined and than in M. (A) mirabilis, but also weakly minera- choanosomal styles are smooth." (Van Soest, 1984: lised). Microscleres may be rare, particularly in Dendritic spicular tracts may be present. specimens from the Bahamas. These specimens, 49.) which I have badly misidentified as Thorecta hor- Tedania anhelans (Lieberkiihn) ridus(see Wiedenmayer, 1977a: 70, pi. 8 figs 2-4, Plate I figure 7, plate 10 figures 3-7, pi. 9 fig. 1), are suggestive of the present coarsely plate 30 figure textured, conulose form. Riitzler (in litt.) has found plate 29 figures 5, 6, 1,

filamentous algae within the fibres of specimens text -figures 57-62 from Belize. In view of this polymorphism in synonymy. Mycale (Arenochalina) mirabilis, and of the occur- Selected Halkhondrta anhelans Lieberkuhn, 1859: 521 pi. il rence of subtylostrongyles in one of the new speci- fig. 6 (only style and onychaete; description composite, mens, it seems likely that Mycale tylostrongylu exterior thai of Reniera in/lulu, fide Schmidt, 1868: 28. Pulitzer-Finali is further syn- a junior synonym. A Trieste, Adriatic Sea). onym may be Gelliodes setosus Keller (1889: 376; Myxilla anhelans. -Schmidt, 1862: 72 (schizotype re- holotype: ZMB 276), known from the Red Sea and examined, overlooked tylotes found; Kvarncr, Adriatic Indonesia, whose only distinctive trait, according Sea). F. WIEDENMAYER

Figures 57, 58. Tedania anhelans. F52009, a conical specimen with a compound apical oscule, x4.1. Figure 57. From top. Figure 58. Median section. -

SPONGES FROM BASS STRAIT 89

Figure 59. Top view. Figure Figures 59, 60. Tedania anhelans. F52010, a small, cushion-shaped specimen, x4.1 60. Perpendicular fracture.

1932: 353 Tedania anhelans. -Hechtel, 1965: 38 (comparison Tedania nigreseens. — Burton and Rao, Indian Ocean).— with ignis, discussion). — Boury-Esnault, 1971: (excessive synonymy, discussion; Tedania - 312 (shape, colour, reproduction, ecology; Banyuls, W Burton, 1932a: 346, text-fig. 44 (excessive distribution). 1937: 27, pi. 3 fig. 22 (diagnosis). Mediterranean). -Thomas, 1973b: 29, pi. 1 fig. 15 (Mahe, Burton, 1862: pi. 7 fig. 11 Seychelles). -Pulitzer-Finali, 1978: 57 (description, ecol- Reniera digitata Schmidt, 75, ogy). -?Boury-Esnault, 1973: 282, fig. 37 (description, (Venice). 1884a: 417 (Torres Brazil, from Recife to S of Bahia; references to records Tedania digitata. -Ridley, 1886a: 52 (Port Phillip Heads, Vic.).- in Senegal and South Africa). -Pulitzer-Finali, 1983: 561 Strait). -Carter, 1887: pi. 11 fig. 3 (off Port Jack- (description, ecology).- Van Soest, 1984: 52 (comment RidleyandDendy, 51, 1887c: 157 (Gulf of Mannar, India). on synonymy in Burton and Rao, 1932, as Tedania son). -Dendy, (Port Phillip Heads, Sorrento, Queen- nigreseens, see below). Dcndy, 1895: 258 scliff, Vic). -Topsent, 1897: 453 (Ambon, Indonesia). Tedania anhelans var. digitata. — Topsent, 1936a: 23 1898: pi. 19 fig. 10 (Southern Viet (resume, colour, ecology, spongin, distribution in Western ?Lindgren, 299, (Azores). -Baer, 1906: 17 f. Mediterranean). Nam). -Topsent, 1904: 176 varr. nov., East Africa). -Lundbeck, 1910: 4-6 (dis- nee Tedania anhelans. -Burton, 1954: 229 ( = Teda- (3 -Hentschel, 1911: 332 (Western Australia). nia ignis fide Hechtel, 1965: 37). cussion). 1912: 348 (discussion; Arafura Sea).- T Reniera nigreseens Schmidt, 1862: 74 (Kvarner, Hentschel, 16 (Reniera nigreseens, R. muggiana, R. Yugoslavia: description that of a Reniera in the modern Topsent, 1920a: redescription of type material). - sense). digitata synonymised, 90 F. WIEDENMAYER

A C v-n C

^/k\. - :

Figure 61. Tedania anhelans. F520I1, view of base, x5. The basal membrane is draped in the large canals, but otherwise stripped. The exposed choanosome contains eggs or embryos (some embedded in the pinacoderm), and (above and below) sand.

Dendy, 1922: 99 (Indian Ocean). -Shaw, 1927a: 434 Tedania rubicunda Lendenfeld, 1888: 190 (Port (Maria Island, Tas.J.-Topsent, 1928: 247 (W. Mediter- Jackson). ranean, Cap Verde Islands). — Koltun, 1959: 159, text-fig. nee Tedania rubicunda. — Hentschel, 1911: 334, text- 117, pi. 25 fig. 1 (Japan Sea). fig. 25 (fide Hallmann, 1914). Tedania digitata var. verrucosa Carter, 1886a: 53 (Port Tedania digitata var. rubicunda. — Hallmann, 1914b: Phillip Heads, Vic). -Dendy, 1895: 258 (holotype 366, text-fig. 1 1, pi. 17 fig. 4 (syntype in AM redescribed). re-examined). Tedania rubra Lendenfeld, 1888: 191 (Port Jack- Tedania digitata var. fibrosa Ridley and Dendy, 1887: son). -Whitelegge, 1889: 185 (off Balls Head, Port 51 (off Port Jackson). Jackson). Tedania digitata forma inermis Hentschel, 1911: 333 Tedania digitata var . rubra.- Hallmann, 1914b: 371, (Western Australia). text-fig. 12 (syntype in AM redescribed). Tedania digitata forma polytyla Hentschel, 1911: 333, Clathrissa elegans Lendenfeld, 1888: 218 (Port Jack- text-fig. 24 (Western Australia). son and Port Phillip Bay, the latter unpublished). - Reniera ambigua Schmidt, 1864: 39, pi. 4 fig. 8 Whitelegge, 1889: 186 (Maroubra Bay, NSW).- (Yugoslavia). Hallmann, 1912: 146. -Hallmann, 1914a: 267, 412 (as Suberites panis Selenka, 1867: 570, pi. 35 fig. 16 (Port species dubia, type material in AM overlooked). Phillip Bay, Vic). Tedania assabensis Keller, 1891:313, pi. 1 6 figs 12 — 11, Tedania panis. Thiele, 1903a: 946, pi. 28 fig. lla-c (Red Sea). -Row, 1911: 353 (Red Sea). -Burton, 1927b: (holotype re-examined, measurements and figures of 81 (Red Sea, Suez Canal). spicules). Tedania anhelans forma assabensis. Levi, 1965a: 17, Reniera muggiana Schmidt, 1868: 28 (Trieste, Adriatic text-fig. 19 (Red Sea). Sea). ? Tedania ignis (Duchassaing and Michelotti): de SPONGES FROM BASS STRAIT 91

peripheral umbels and penicils of ascending spicular tracts. Choanosome cavernous, bread-like, with sand in some specimens. Main skeleton an irregular, often vague renieroid reticulation of spicular tracts with scarce spongin, frequently masked by spicules in confusion. Few thicker, more compact longer ^ tracts may occur, mostly dendritic. Tylotes with few distal spines. Some oxea (styloids) may occur.

Description. Small, thickly encrusting to cushion- shaped, mostly not over 6 cm in width. Smallest specimens from station BSS 181, attached to chaetopterid worm tubes. F52009 (text-figs 57, 58) conical, 2 x 2.5 cm wide at base, 2 cm high. Speci- mens from station KG 6 larger, cushion-shaped, F52012 8 x 11 x 2 cm, F52013 6 x 8 x 2 cm. Dendy's specimens more massive and larger than new ones, often irregularly lobate. Lobes often compressed and convoluted, partly concrescent, distally separated by meandering fissures (pi. 10 fig. Figure 62. Tedania anhelans, spicules from F52009. a-c. 6). Syntype of Tedania rubicunda Lendenfeld in Range and examples of styles and oxea, x 263. d. Tylotes, Sydney (pi. 10 fig. 7) also of this type. x 263. e. Enlarged end of tylote, x 790. f . Range of ony- life, chites, x263. g. Enlarged ends of onychaete, x790. In bright orange to deep vermilion on surface, light orange-brown internally. Surface in life 2.5 YR 7/12 to 5 YR 7-8/12 in F52012, 10 R - 2.5 Laubenfels, 1950a: 21, text-fig. 13 (Hawaii). — de Lauben- YR 7/12 in F52013. In drying, some areas of sur- fels, 1951: 260 (Hawaii). face turn dark greenish grey. Softly rubbery, a bit

''Tedania ignis pacifica dc Laubenfels, 1954a: 129, text- limp, to moderatley firm but compressible. fig. 81 (Caroline Islands, Micronesia). Surface in life smooth to vaguely verrucose and rugose. Common pattern of subdermal crypts set Material examined. Station KG 2, two specimens (F52009, off darker, greyish colour. Out of water, pat- F52010); station KG 4, one specimen (F5201I); station by relief dermis, KG 6, two specimens (F5201 2, F520I 3); station BSS 181, tern accentuated in by receding more two specimens (F51939, F51940). Type specimens of so in preserved specimens. Crowded polygonal pits Tedania digitala var. verrucosa (BMNH), T. digitatavar. surrounded by reticulate rugae, passing into mean- rubicunda, T. rubra, Clathnssa elegans (AM and drine, cerebroid pattern. Widths of depressions and BMNH). Hypotypes of Tedania digitala: Carter, 1886a rugae 0.4-0.8 mm. Dermis over depressions riddled (BMNH), Dendy, 1895 (NMV). by ostia, some barely visible optically. Flat speci- Diagnosis. Shape and size variable: thinly encrust- mens (F52012, pi. 10 fig. 3; F52013) have fine regu- ing, cushion-shaped, massive-lobate, digitate to lar radial striation along parts of margin, as do stubby-ramose, tubular, irregularly frondose, or specimens with compressed, erect lobes, then lon- trabeculate, often in combination in large speci- gitudinally below apices. mens. Softly rubbery, limp to moderately firm, Oscules often slightly raised, irregularly scattered compressible. Colour variable: fresh surface bright to clustered in flat specimens, more regularly on orange to vermilion, or brown, ochre, dull green, apices of rounded lobes, numerous on edges of bluish, grey, blackish. Superficially orange speci- compressed lobes. Width 1-5 mm; rim sharp, mens often with other, darker hue in periphery. mostly irregular and sinuous, often composite, with Frequently found as epibionts on algae and sessile confluent shallow atria, with mouths of exhalant invertebrates. Surface smooth to vaguely verrucose, canals between ridged partitions. rugose, or brain-like. Digitate and frondose pro- Choanosome conspicuously cavernous (text-figs tuberances often longitudinally striate to ridged. 58, 60). Principal exhalant canals predominantly Oscules conspicuous, mostly apical on lobes, often terete, straight, perpendicular to surface, 1-4 mm clustered or composite. Vestibules well developed, wide. Some branch and anastomose. Often folded paratangential and dendritic-meandering, or to septate transversely, lining distinct, with clusters ascending, palisade-like. Ectosome reinforced by of apopyles. Inhalant canals 200-500 urn wide, peripheral choano- tangential bundles of tylotes, or by extensions of regularly arranged radially in 92 F, WIHDENMAYFR

1965: 38). The appearance of dai some, in palisade-like pattern, particularly in mas- ignis (Hechtel, exposure to air of Ba> sive specimens. Some locally arranged tangentially, greenish grey patches upon corresponding to striation noted above. F52011 Strait specimens may point to subtle chemic in Mediterranean populations a (text-fig. 61) contains much sand in choanosome, changes, which also eggs or embryos, deep red in life. more easily developed in life. I recall, in this coi Main skeleton an irregular reticulation of pale, text, the greenish olive-drab hue noted by m clear spongin fibres, cored (thinner ones) or packed (Wiedenmayer, 1977a: 133) in the fresh peripher, 7". by styles. Fibres straight to flexuous, 18 to 90 /mi choanosome of ignis. Tangential dermal tylott

thick, branching and obliquely anastomosing with and detachable dermis have been observed i 2> little order. Meshsizes 40 to 350 /mi. Interstitial Mediterranean specimens by Schmidt (1868: megascleres (thin styles, tylotes and onychaetes) description of Reniera muggiana). The wide ect irregularly scattered to crowded, occasionally logical affinities in all three biogeographic group obscuring meshwork. In some areas, interstitial are very similar: typical members of the foulin megascleres, single or in pairs, take part in mesh- community (e.g. Dendy's specimens from Sorrent work as anastomoses. Ectosome about 350 /tm and Queenscliff, harbours inside Port Phillii thick, with umbellar, dendritic or feathery arrange- Heads), occasionally extending to slightly brack- ment of tylotes. Median branches and stems of ish water (mangrove community for T. ignis umbels occasionally recurved, festooned. Schmidt's capriciously shaped types of 7?. digital' Speculation: (1) Styles, occasionally oxea, from Venice), but also found on a variety of rock 120-777-203 X 1.1-3.2-6.5 /mi (144-766-188 x to muddy substrates of the Mediterranean (l-7( 1.1-2.0-2.9 /tm in F51939). (2) Tylotes with few m fide Bourv-Esnault, 1971; Pulitzer-Finali, 1978- distal spines, immature ones (in choanosome) 1983). almost strongyles, 158-202-239 x 1.5-2.7-5 (Oa The variability in sculpture of onychaetes in J (169-755-213 x 1.5-7. 9-2.3 /tm in F51939). (3) anhelans (as digitata), from almost smooth to Faintly sculptured onychaetes, 69-725-168 x spinulate, has been discussed by Topsent (1897 0.9-1.8 /mi. 454) and Lundbeck (1910: 5). Lindgren's (1898

record of T. digitata is here treated with reserve Remarks. The new material studied here differs because of ihe exceptionally large spicules (styles from specimens of T. ignis (Duchassaing and 300 x 16 /tm; tylotes, 240 x 6 /tm; onychaete* Michelotti, see Wiedenmayer, 1977a: 133; van almost hastate, 336 x 5 /tm). In conclusion, van Soest, 1984: 49) primarily by the lack of dermal Soest's remark (1984: 52) is still valid, that large specialisation (tangentially strewn tylotes and ony- series of specimens of this species complex from chaetes in T. ignis), and by the presence of a different parts of the world need to be examined peripheral "palisade" of inhalant canals (not men- before such distinctions are meaningful. Tedania tioned by van Soest, nor noticed by me in Baha- cammixta Ridley and Dendy (1886: 335; 1887: 52), mian specimens). The spicules in the new specimens originally described from Moncoeur Island, Bass from Bass Strait are generally shorter that in T. Strait, 70 m, recorded again by Dendy (1895: 258) ignis (Wiedenmayer, 1977a: styles, 217-242 x 4-6 from Port Phillip Heads, seems to be a good spe- /tm; tylotes, 220-246 X 2.5-4 (ira; onychaetes, cies. Topsent (1904: 176) recorded it from the 40-220 /tm; van Soest, 1984: styles, 202-281 x 4-9 Azores (98 m, gravel, sand, broken shells). It is dis- /mi; tylotes, 180-248 x 2.5-4.5 /tm; onychaetes in tinct by its creamy yellow surface, consistently high 2 sizes, 30-95 (im and 154-247 /mi) and agree with content of foreign inclusions, and subtylotes Dendy's measurements for T. digilata from the without distal spines. Both styles (300-400 x 4-6 Indian Ocean (1922: 100: styles, 200 x 6 /tm; /tm) and subtylotes (325-350 x 3-4 /mi) are longer tylotes, 200 x 4 /mi; onychaetes, 140 /tm). Such than in T. anhelans, and comparatively slender. differences are not meaningful on the species level. Measurements cited by Pulitzer-Finali (1978, 1983) for specimens from the western Mediterranean Forcepia Carter (styles, 205-280 x 4.5-11 /mi; tylotes, 185-270 X 2.3-4 /tm; onychaetes, 60-190 /tm) are very simi- Forcepia Carter, 1874: 248, pi. 15 fig. 47. -Topsent, lar to those of T. ignis. 1904: 177 (definition, review). -Lundbeck, 1905: 19 (discussion), 198 - The colour variants of T. anhelans other than (definition), 210 (discussion, review). Dendy, 1922: 91 (definition, discussion). -Van Soest, red and orange, noted in the diagnosis above, apply 1984: 66 (definition, Ecloforcepia as subgenus), 69 chiefly to Mediterranean records, and have been (discussion). recorded only sporadically from the lndo-Pacifie Ecloforcepia Cabioch, 1968: 232. -Bourv-Esnault, region. Red to orange is remarkably constant in T. 1973: 280. -Van Soest, 1984: 66, 69. SPONGES FROM BASS STRAIT 93

Diagnosis. "[Tedaniidae] with choanosomal monactinal or diactinal (tylole) megascleres and smooth and/or acanthose forcipes, isochelae and sigmata as microscleres." (Van Soest, 1984: 66.) Sigmata may be absent. Choanosomal styles may- be smooth or spined.

Remarks. The only diagnostic trait separating the two subgenera, according to van Soest, is the presence/absence of styles. This subdivision, though probably desirable, is here avoided for a nomenclatorial reason: Forcepia colonensis Carter, as type species of Forcepia, would automatically be the type species of the nominotypical subgenus. This West Indian species, however, is only known from isolated large acanthose forceps, and its placement in either subgenus would be conjectural. Van Figure 63. Forcepia biceps, spicules, a. Range of tylotes, Soest (1984: 66) pointed out that the same type of X 68. b. Enlarged ends of tylote, x263. c-d. Arcuate forceps occurs in a West Indian sponge, described isochelae: c, range, x263; d, example, x790. e. Range

by him as Forcepia (Ectoforcepia) trilabis Boury- of forceps, x263. f. Enlarged forceps, x788. Esnault, but that synonymy was uncertain.

Forcepia biceps (Carter) No styles. Forcipes spined, massive, with curved apex. Arcuate isochelae, no sigmata. Plate 10 figures 8-11, plate 30 figure 2, Description. Cushion-shaped, with sinuous outline text-figure 63 and pinched marginal lobes, some upturned; 1 1 x

Forcepia colonensis.— Carter, 1885a: 110, pi. 4 tig. 2 15 cm in width, 3 cm in height. (Port Phillip Heads, Vic). -Carter, 1886a: 53 (old In life deep red on the surface (7.5 R 5-6/12), record). -Dendy, 1896: 24 (outside Port Phillip Heads; paler, more orange in choanosome; now beige with — synonymised with Suberiies biceps Carter). Lundbcck, yellowish and greyish tinges. Firmly rubbery, easily 1905: 210 (in discussion: Australian record distinct, pos- torn, slimy in life. Mucus preserved in alcohol as sibly not in Forcepia). flaky to stringy sediment. nee Forcepia colonensis Carter, 1874: 248, pi. 15 tig. Surface partly smooth, glabrous, partly wrin- 47 (Colon, Panama; isolated forceps from sediment). kled. Wrinkles locally radial, or in vaguely rugo- Suberiies biceps Carter, 1886b: 117 (Port Phillip small, 1-2 Heads). reticulate to cerebroid pattern. Oscules Forcepia michaelseni Hentschel, 1911: 336, 337, text- mm wide, in sieve-like groups on marginal lobes. 'fig. 27 (for Forcepia colonensis: Carter, 1885; Rottncst On surface nol coveted by encrusting and branch- Island, WA). ing algae, fine ostia virtually ubiquitous, conspic- Material examined. One specimen, station KG 6 (F52018). uous only in dendritic depressions, chiefly around IHolotype of Suberiies biceps (BMNH). Hypotypes of margin, often clustering in interstices of fine dermal 'Forcepia colonensis: Carter, 1885a (BMNH). Dendy, 18% reticulum over subdermal spaces. (NMV). Choanosome riddled by exhalant canals, •Diagnosis. Cushion-shaped to irregularly massive, straight, terete, 1-3 mm wide, tangential and per- lobate. Lobes marginal and apical, tending to be pendicular to surface. They bear widely spaced radial, in conical. Firmly rubbery, easily torn, slimy in life. annular sphincters. Subdermal crypts with sparse Surface deep red, internally pale red-orange in life. palisade-like zone about 5 mm deep, Surface partly smooth, glabrous, partly wrinkled anastomoses. Numerous embryos, deep red in life. to angular tracts in parallel, radial and reticulate patierns. Oscules Main skeleton of tylotes in sinuous /*m) spacing, clusterd, sieve-like, on lobes. Vestibules conspicu- of variable thickness (45-175 and with oblique ous, numerous, radial, in palisade-like peripheral infrequently branching and merging spicules or loose fascicles. zone about 5 mm deep. Ectosome mostly thin, up anastomoses of single Structure frequently obscured by megascleres in to 1 mm, contains reticulum of paratangential tylotes. inner choanosome cavernous, bread-like. confusion. thin, comprising vague reticu- 'Main skeleton of tylotes only, in vague reticulation, Ectosome often tylotes over spaces traversed •with ascending tracts commonly better developed, lum of paratangential

tracts, but up to 1 'frequently obscured by megascleres in confusion. by umbellar ends of main mm .

94 F. WIEDENMAYER

1984: 60 (type species re- thick, with tylotes packed in contusion around and monotypy).- Van Soest, synonymised with Lissodendoryx). irregular alveoles. examined, Damiriana dc Laubenfels, 1950a: 13 (type species: D. Speculation: (1) Tylotes, mostly somewhat flex- hawaiiana de Laubenfels, 1950a, by monotypy. uous, terete to slightly fusiform, with distinct spherical heads, length 393-457-467 jtm; width of shaft, Diagnosis. "[Tedaniidae] with monactinal ordiac- 5.6-10.6 jam; width of tyles, 9.4-13.5 /mi. (2) Arcu- tinal, smooth or acanthose choanosome ate isochele, 16.4-19 x 5.4-7.5 /mi. (3) Acanthose megasderes arranged in a renieroid [subisodictyal] forceps with slightly waisted apex and gently reticulation of single spicules [or bundles]; outward-curved, divergent capped arms, length microscleres always include arcuate isochelae and 56-73-98 urn; total width, 25-29-49 /tm; proximal sigmata." (Van Soest, 1984: 54). Sigmata may be width of arms, 5-6-7.1 /*m. absent. Longer tracts, sometimes dendritic, may be present. The arcuate chela is the only real distinc- Remarks. Of the other two congeners known from tion from Myxilla, which always has anchorate southern Australia, F. crassanchorata Carter chelae. (1885a: 111, pi. 4 fig. 3; known only from a dry specimen from Port Elliot, SA) has often subty- Remarks. In his revision and discussion of Lis- lote styles next to tylotes; small, frail, faintly spined sodendoryx, van Soest (1984) emphasised reticu- forcipes with almost pointed apex and straight, late skeletal structure, tylote ectosomal spicules, little diverging arms; isochelae of two sizes: (1) and arcuate chelae and sigmata as micro-scleres. stout, with strongly arched, dorso-ventrally com- The stylote versus diactinal morphology of

pressed shaft, (2) much smaller, frail, normal choanosomal megasderes is dismissed as having

shape; frail, parabolic sigmata. Forcepia carleri little significance. This is based chiefly on

Dendy ( 1 896) is thoroughly sandy, and has proper choanosomal strongyles in his L. strongy/ata and spicules only at the surface. These are only forcipes on the undifferentiated tylotes in the choanosome

and stout chelae, both similar to those in / . cras- of L. sigmata (dc Laubenfels). In consequence, van sanchorata, but the chelae have only half the size Soest also included species with choanosomal oxea

of the larger ones in Carter's species. (Zetekispongiu) in Lissodendoryx. I concur with the inclusion of the two West-Indian species described Lissodendoryx Topsent by van Soest next to L. isodictyalis, as I have pre- - Lissodendoryx Topsent, 1892a: 63, 96, 97. 1 undbeck. viously (Wiedenmayer, 1977a: 137) interpreted 1905: 153 (definition, discussion). — Wiedenmayer, 1977a: mucronate modifications of tylotes in L. sigmata 135 (references). -Van Soest, 1984: 54 (definition), 59 as transitional to some types of subtylostyles found (discussion). in L. isodictyalis. Dendoricella Lundbeck, 1905: 126 (type species: D. More meaningful is the occurrence of rhopalum Lundbeck, 1905, by subsequent designation in choanosomal strongyles and oxea in Myxillinae de Laubenfels, 1936a: 53). -Topsent, 1928: 52 (in review of Myxillinae). from other parts of the world, several of which Paramyxilla Dendy, 1905: 233 (in footnote; type spe- have been misplaced (in the wrong genus) in the

cies: Halichondria frequens Carter, 1881a, by past. I should stress, before the following review,

monolypy). — Topsent, 1928: 53 (synonymised with that I follow Topsent's diagnostic treatment of the Dendoricella). Myxillinae (Tedaniidae with chelae), except for Hamigera sensu Dendy, 1922: 90. — nee Hamigera dropping the distinction diactinal/monactinal Gray, 1867a: 536. choanosomal megasderes. The basic distinction, Damiriellu Burton, 1935: 404 (type species: Dumiriu adopted by Topsent and most other systematists cavernosa Topsent, 1892c: xxii). — Pulitzer-Finali, 1978: 56 (type species redescribed).- Van Soest, 1984: 60 (except Dendy in his later work), of forms with (discussion). arcuate chelae versus forms with anchorate chelae, Jones dc Laubenfels, 1936a: 79 (type species: Myxilla was introduced by Lundbeck (1905). The only amaknakensis Lambc, 1894, by original designation and difficulty here is the exception in Myxilla? decepta monotypy). — Bakus, 1966: 499, 513 (synonymised with Kirkpatrick (see Koltun, 1964: 52), which has both Lissodendoryx) sorts of chelae. As will be justified below, I do not Merriamium sensu de Laubenfels, 1936a: 83 (part: not regard sigmata mandatory in diagnosing Lissoden- type species M. tortugasense de Laubenfels, 1936a).— doryx, thus also following Lundbeck (1905) and de Laubenfels, 1939: 4 (part, not M. roosevelti). Topsent (1928). IWaldoschmittia de Laubenfels, 1936a: 95 (type species: Crella schmidti Ridley, 1884a, by original The genera in question, which need clarification, designation). are: Zetekispongiu de Laubenfels, 1936b: 446 (type species: (1) Dendoricella Lundbeck. The type species, D. 7.. zonea de Laubenfels, 1936b, by original designation rhopalum, has the spiculation and skeletal struc- SPONGES FROM BASS STRAIT 95

lure of Lissodendoryx (without sigmata). The only nosa, with most "strongyles" being styloids (one end significant diagnostic trait of this genus, for Lund- manubriate, the other mucronate), a few with beck, was exclusive presence of oxea or strongyles residual spination. The chelae are said to be arcu-

in the choanosome. De Laubenfels (1936a: 53) was ate chelae of two sizes. While this is no doubt true mistaken in characterising this genus with the for the large kind, the small one is like a pluriden- presence of "odd spiny palmate" isochelae. Only tate unguiferous chela with c-shaped, smooth shaft.

in D. rhopalum, and here only in one specimen Topsent correctly compares it with the peculiar known, are odd arcuate chelae dominant, in which small "chela" of Lissodendoryx indistincta (see the compressed shaft bears a high, thin dorsal keel, Lundbeck, 1905: 162, pi. 16 figs. 3 f, g). This which may be again modified, in few chelae, by differs from the one in L. cavernosa only in having being little lobate. In most specimens, weakly thin, finely lobate rims along both sides of the keeled modifications are subdominant to equally shaft. Both have minute teeth barely visible in frequent with more normal types having lightly smallest examples. Dendy (1921: 119, fig. 31) com- compressed shafts. The only peculiarity (but only pared the one in L. indistincta with the dentate sig- diagnostic for this species) is the acuminate shape maspires in some spirophorids (Craniella disigma of the alae. Otherwise these are clearly arcuate and Chrotella amphiacantha Topsent, 1904). I chelae, lacking the proximally separate lateral alae doubt that these types are homologous, but I agree and the fimbriae of anchorate chelae. (Incidentally, with Dendy that those in Lissodendoryx are neither a separate new genus for Myxillinae with odd spiny true chelae nor sigmata. It is significant that the chelae would be more justified for Myxilla inso- boreal sibling species, L. indistincta, has true sig-

/e/?s Koltun (1964: 52, text-fig. 11).) Of the other mata and normal choanosomal styles varying from species inlcuded by Lundbeck, Crella schmidti smooth to weakly spined. The specimens described

Ridley is the type species of Waldoschmirtia (see by Pulitzer-Finali (1978: 56) have as choanosomal Damiriana, below). Damiria cavernosa Topsent is megascleres chiefly asymmetrical strongyles, with the type species of Damiriella (see below). Des- one or both ends slightly swollen. The species was macidon abyssi Topsent (1904: 204) has unguifer- recorded again by Pulitzer-Finali (1983: 560). Lis- ous chelae (without fimbriae) according to sodendoryx cavernosa in Rutzler (1965: 31) is acanthostyles in the Topsent's figure, and is probably a Myxilla. Den- doubtful, as he mentions only doricella obesichela Lundbeck (the last included choanosome. On the other hand, his sponge iden- from its ecol- species, known only from a fragment) is similar to tified as Lissodendoryx isodictyalis, gravel, might be D. rhopalum in having oxea in choanosome and ogy and packing with fine (L. caver- dermis, but has normal (stout) arcuate chelae and cavernosa, though it was whitish in life as yellow by Topsent, 1936a, sigmata. It is thus close to Zetekispongia zonea nosa was described Pulitzer-Finali, 1983), and Rutzler men- (which has dermal tylotes). In conclusion, I see no and by grounds for separating Dendoricella from Lis tioned anchorate chelae ("Isancora"). other sodendoryx. (4) Jones. The type species differs from Myxillinae in having sparsely spined oxea as dermal (2) Paramyxilla Dendy. The type species has megascleres. De Laubenfels therefore included the choanosomal acanthoxea. I follow Topsent in syn- Crellidae. It is otherwise like Lis- onymising this with Dendoricella (and thus with "genus" in the also Koltun, 1959: 135). Lissodendoryx), in analogy with the spicular vicar- sodendoryx (see De Laubenfels apparently iance of smooth/acanthose choanosomal styles in (5) Merriamium. chiefly to include "atypical" spe- Myxillinae. intended this genus assigned species by cies of Lissodendoryx and Ectyodoryx (he (3) Damiriella Burton (1935: 401 ; type "Tedaniidae" and "Myx- monotypy: Damiria cavernosa Topsent, 1892c: the "typical" species to the illidae", respectively) lacking sigmata. The type spe- xxii). Van Soest (1984: 60) mentioned this as being West-Indian M. tortugasense (ending here close to Lissodendoryx, but lacking sigmata. The cies, the to conform with de Laubenfels' (1939) use type species, from Banyuls, has only been briefly emended with all other species included) .diagnosed by Topsent (1892c), as having of the neuter gender Phorbas amaranthus and small is a junior synonym of i choanosomal strongyles, dermal tylotes, and Michelotti, according to van Soest isochelae with curved shaft, and pectinate (comb- Duchassaing 86) who examined the holotypes of both spe- shaped) ends ("a bouts pectines"). The latter state- (1984: Merriamium is thus a junior synonym of ment was unclear to Lundbeck (1905); Burton cies. Phorbas (ex Anchinoe) in the Anchinoidae. (1935), in his generic diagnosis, interpreted it as Hymeniacidon paupertas Bowerbank has been unguiferous (pluridentate?) isochelae. It was placed in Ectyodoryx (Burton, 1932a: 313; followed redescribed in detail, from fresh material, by Top- by Koltun, 1964), Hymedesmia (Lundbeck, 1905; sent (1936a: 19, fig. 4), as Lyssodendoryx caver- 96 F. WIEDLNMAYLR

figs 15c, c') are ambiguous, but that ii 113; Topsent, 1928: 57), and more recently, in 19, Desqueyroux-Faundez (1982, fig. 42), based on on. Anchinoe (correctly Phorbas: Borojevic et al., Topsent's four specimens, clearly shows a tridem 1968: 23; Pulitzer-Finali, 1983: 564, 610). It is clear of anchorate chela, deviating from the usual shap. that the species does not belong in the Tedaniidae. tate only by the foreward inclination of the lateral teeth M. roosevelti, the only species added by de Lauben- descriptions and figures of isochelae in Lev fels (1939, review with key of the species included The fig. 25; 1965a, fig. 17), and Tsurnama in 1936a), from the Galapagos, is only briefly (1958, to the arcuate type. There described, microscopically only as to spiculation. (1969, pi. 2, fig. 4) refer distinct possibility that two species and twc It probably belongs to either Hymcdesmiidae or is a This should Anchinoidae. genera are involved in this complex. re-examination of the older material, Of the remaining 12 species included, those eight be settled by Crella schmidti Ridley. If the with only one kind (size) of choanosomal particularly that of out to possess anchorate chelae, megasclere should be left in Lissodendoryx: L. latter should turn be available for it and any sophia and L. lobosa (see Lundbeck, 1905), L. certa Waldoschmittia would chelae. This genus name, (Topsent, 1892a), L. mollis (L.indgren, 1897; de other records with such ultimately fall in synonymy to Laubenfels' replacement name lindgreni is super- however, would would be exactly parallel to fluous), L. buchanani Topsent (1913a), L. lund- Myxilla, since the case becki Topsent (1913b; has sigmata according to the Lissodendoryx/ Damiriana. in the original description), L. styloderma Hentschel The presence of diactines or monactines be interpreted as. (1914), and L. kytna de Laubenfels (1930). Four choanosome of Myxillinae can species (clavigera, atlantica, maculate, ramilobosu) homology of the two basic types, more commonly vicariance, should be left in Ectyodoryx. expressed as an interspecific spicular with both types present within one spe- (6) Zetekispongia de Laubenfels differs from occasionally variable typical Lissodendoryx by possessing oxea only as cies, then commonly in individually choanosomal megascleres. proportions (e.g. acanthostrongyles next to acan- in Myxilla brunnea, Stelodoryx flabellum, (7) Damiriana de Laubenfels. The type species, thostyles probably according to the original description and text-figure see Koltun, 1959). The spicular vicariance differentiation of ectosomal 7D, has arcuate chelae of two sizes, it is probably extends to the synonymous with Zetekispongia zonea. According megascleres. It is expressed in the frequently simi- to Topsent (1897) and Burton (1935), Damiria aus- lar dimensions, occasionally in the morphological traliensis Dendy (1896) is a junior synonym of similarity or even identity of choanosomal and Crella schmidti Ridley (1884a: 432). Levi (1958: 30) ectosomal megascleres. Hereby, the rule seems to synonymised Damiria australiensis Dendy. 1896, be invasion of the choanosome by ectosomal Damiria schmidti: Topsent, 1897, Damiria aus- megascleres, and by the virtual or total expulsion tral iensis: Lindgren, 1897, 1898, Dendoricella of typical choanosomal types. Total invasion of the schmidti: Hentschel, 1912, and Dendy and Freder- choanosome by tylotes is the case in the West- ick, 1924, and Damiriana hawaiiana de Lauben- Indian L. sigmata (de Laubenfels), virtual expul- fels with Crella schmidti Ridley, and described a sion of choanosoml styles exists in L. dendyi (Shaw, sponge from the Red Sea as Damiriana schmidti see end of Remarks on L. isodictyalis. below). It (Ridley) (again in Levi, 1965a: 16). He was followed is unlikely that the Antarctic L. styloderma by Tsurnamal (1969: 153), Desqueyroux-Faundez represents an inversed trend (invasion of the ecto- (1981: 741), and Pulitzer-Finali (1983: 609). some by choanosomal styles), as choanosomal According to this view, the single species would be styles in this species are distinctly larger and are widely distributed (Australia, Indo-Malayan frequently modified as subtylostyles. This spicu-

province, Red Sea, eastern Mediterranean) and lar vicariance is corroborated by the variable characterised by dermal tylotes, choanosomal oxea, proportion of asymmetrical and even stylote arcuate chelae, and sigmata, all of variable dimen- modifications of dermal diacts in several species

sions. Van Soest (1984) regarded Damiriana as a (e.g. Lissodendoryx lundbeck i, L. diversichela, L. probable synonym of Lissodendoryx. fragilis, L. stipitata, see Koltun, 1959). A parallel The nature of the chela in Crella schmidti is not is seen in Tedania: occasionally oxea are present clear from the original description, nor from those in the choanosome of T. anhelans from Bass Strait of Dendy (1896), Topsent (1897), and Thiele (as noted above); in T. oxeata Topsent (see Koltun, (1903a: 954, who mentioned "tridentate isochelae" 1964, 1966a), choanosomal styles are completely of 2 sizes). Ridley's figure (1884a, pi. 41, fig. aa) replaced by oxea. In the latter species, dermal could be interpreted as representing an anchorate megascleres are also oxea, but decidedly smaller chela. The relevant figures in Lindgren (1898, pi. and verging on tornotes. Another, more distant SPONGES FROM BASS STRAIT 97 parallel, regarding choanosomal megascleres, may sodendoryx jacksoniana (Lendenfeld, 1888, with be seen in Axinellida, e.g. in Trachycladus small strongyles next to the normal styles in the laevispirulifer, described above. choanosome) in this genus should be reconsidered, The ease is not always clear-cut in Lissodendo- in favour of a new genus, distinct from Chaetodo- ryx: choanosomal strongyles may be slightly asym- ryx by the lack of echinating acanthostyles. I do metrical, as in L. cavernosa (see above) and in L. not think that the interstitial "oxychaetes" of the buchanani Topsent (1913a). A parallel spicular type species of the latter (C. richardi Topsent, 1927, vicariance is the better known one of spiny versus see Topsent, 1928: 242; by monotypy) are really smooth choanosomal styles. The presence of either distinct from the onychaetes of the Tedaniinae. acanthose or smooth styles has been used to Their size (225-245 x 2-3 /*m) is well below that segregate Myxilla and Lissodendoryx, respectively, of the largest onychaetes in Tedania (up to 760 X last by Dendy (1922: 88, 90, with Hamigera used 4.5 fim in T. tantula (Kirkpatrick) fide Koltun, as a senior synonym of Lissodendoryx). He rejected 1964: 59). Since some onychates are smooth, others the presence of either anchorate or arcuate chelae almost tornote, their homology with the small inter- as a diagnostic criterion (referring to both as stitial strongyles of "L." jacksoniana is likely. "tridentate isochelae") on the ground that they are These considerations lead to the case of indifferent in phylogeny (see also Dendy, 1921: Hamigera Gray. The genus is still monotypic, with 112f., 144). Lundbeck (1905) has shown that inter- the type species Cribrella hamigera Schmidt (1862, specific spicular vicariance of smooth/acanthose cited by Gray as Hamigera ntbens because of his choanosomal styles is common in boreal species of aversion to tautonymy). Its spiculation corresponds Lissodendoryx, and that some species (L. fragilis, to that of Lissodendoryx without sigmata. There L. indistincta) combine both types in variable are basically two views on its skeletal structure and proportions, with transitions. Dendy (1922: 90) systematic position: ' acknowledged the difficulty with his system, but (1) Thiele (1903a: 353) and Dendy (1922: 90). retained a distinction of the two genera (Myxilla Both saw a "myxillid" affinity of Hamigera. Thiele, and Lissodendoryx) for a practical purpose. Several who examined a schizotype of Cribrella hamigera species of Myxilla with smooth styles are known Schmidt, thought that it was a "myxillid" with an (Dendy, 1924: 362). Lundbeck's diagnostic empha- irregular, chiefly dendritic main skeleton with min- sis on chelae types has been followed by most sys- gled diactines and monactines, distinct from Myx- tematists, especially by Topsent (1928: 52f.) in his illa and Lissodendoryx, to which he attributed a influential synopsis of Myxillinae. In L. firma regular, subisodictyal main skeleton of monactines (Lambe) from the west coast of North America, only. His attribution of ternatensis (a synonym of as redescribed by Bakus (1966: 487), smooth and isodiclyalis) to Hamigera was probably motivated sparsely spined styles of similar size occur in the by excessive reliance on the peripheral choanosome in choanosome in variable proportions between speci- (he examined only slides and a small fragment mens, often almost mutually exclusive. Spicular paraffin, left by Kieschnick), where dendritic tracts but tracts occur in the peripheral choanosome, cored with mixed megascleres are indeed common, and "very crudely echinated" by the same styles or not exclusive to all peripheries of L. isodictyalis. conclusions (nor acanthostyles. In L. kymade Laubenfels (1932: 75; Dendy did not mention Thiele's solely on see also Bakus, 1966: 491), another species from Lundbeck's, see below), and relied of C. the west coast of North America, lacking sigmata, Schmidt's unsatisfactory description bluntly accepted Hamigera as senior the main skeleton is similar, but there is an imper- hamigera. He fect (intraspecifically variable) segregation of styles synonvm of Lissodendoryx. and Lundbeck (1905: by size, small ones having a spined base. Dendy (2) Topsent (1892a: 102) hamigera Schmidt not as a (1922: 88), in his diagnosis of Myxilla, stated that 173) regarded C. an "ectyonine" related to some acanthostyles in the main skeleton may be tcdaniid, but as then relied on fresh echinating. Equally important to chelae types, as Echinodictyum. Topsent Corsica, and noticed that diagnostic criterion for genera of Myxillinae, when material from Bonifacio, the main skeleton are constituted by Topsent's system of 1928 is maintained, is the con- all tracts of and are echinated by smooth sistent presence of two sorts of choanosomal diacts (strongyles) disposition negated by Thiele, megascleres in some genera, particularly when dis- subtylostyles (a figure and description of the tinguished not only by morphology, but by size, 1903a). (Schmidt's unguiferate, like those of Myxilla veneta and by location (interstitial and/or echinating aux- chelae as of Plumohalichondria plumosa, iliary megascleres, the latter in Ectyodoryx, [a junior synonym 1932-1935: 260, 265] is obviously Ectyomyxilla, Chaetodoryx, "Onychomyxilla" fide Vosmaer, observation of a microsclere type (superfluous), and /option. The inclusion of Lis- based on faulty -

98 F. WIEDHNMAYtR

suchi then new, in which he saw only the falces, not the Koltun, 1959). It should also be noted that alae.) Lundbeck (1905) examined another schizo- megascleres in both genera are frequently asym- type of C. hamigera, and found Topsent's obser- metrical, sometimes stylote (see above). insistence on sigmata vation confirmed. Topsent (1928: 57f.) reaffirmed 1 cannot accept van Soest's the affinity of Hamigera with Echinodictyum (the always being present in Lissodendoryx. There is a

latter is now placed in the Raspailiidae) and the number of species with the usual traits of Lissoden-

Anchinoidae. The latter familial assignment is doryx but lacking sigmata. In addition to those maintained more recently by Boury-Lsnault (1971: cited above, under Merriumium, there are the fol- 322), Levi in Brien et al. (1973: 613), and Pulitzer- lowing boreal and N. Pacific species (see Koltun, Finali (1983: 566) for Hamigera. 1959): L. behringi Koltun, L. papilloma Koltun, L.

Hamigera is aberrant within the Anchinoidae in amaknakensis (Larabe), and L. iranovi Koltun having columnar spiculofibres cored by smooth ("Lissodendoryx" oxeola Koltun seems to have strongyles, echinated by smooth sublyloslyles. unguiferate chelae and may be a Myxilla); plus the Plumose columns of acanthostyles, often echinated Antarctic L.flabellata Burton (1929a). The varia- by smaller acanthostyles, are the rule in the bles in morphology and arrangement of Anchinoidae. megascleres mentioned above (monactinal/diacti-

As I have expressed in my addition to van Soest's nal and regularly/irregularly spined/smooth diagnosis of Lissodendoryx, this author has placed choanosomal spicules; tylote/strongy- loo much emphasis (also in his diagnosis of the lote/oxeotestylote dermal spicules; Tedaniidae) on reticulate structure of the main presence/absence of dendritic tracts and semi- skeleton. Short dendritic portions occur in the main plumose columns) occur in the genus regardless of skeleton (chiefly peripheral) of some massive Lis- presence/absence of sigmata. The sibling pair L. sodendoryx (and of Tedania, as noted above), but indistincia -L. cavernosa is especially instructive in some stipitate and ramose species (L. stipitata in this context. A further point is that no sys- Lundbeck, 1905: 170, see also Koltun, 1959: 133; tematise to my knowledge, has consistently main- L. buchanani Topsent, 1913a: 626) the dendritic tained a generic separation of asigmate species of style is more pervasive, though commonly sup- Myxilla (see review in Dendy, 1924: 362), of Eorce- plemented by interstitial single spicules of the same pia (see above), and of Ectyodoryx. The partial kind. A parallel situation exists within Myxilla (see exception is de Laubenfels' unsuccessful attempt diagnosis in Lunbeck. 1905: 131; Topsent, 1913a: with Merriamium.

626), so that Steiodoryx is probably a synonym of Myxilla (pluridentate versus tridentate chelae not Lissodendoryx isodictyalis (Carter) being a reliable criterion). Plate 10 figure 12a, b, plate II figures 1, 2, Hamigera can be regarded as a borderline case between Tedaniidae and Anchinoidae. Van Soest plate 30 figure 3, text-figure 64 (1984: 71, 89) commented on other affinities Selected synonymy. between the group Myxilla, Ectyomyxilla and Halichondnu isodictyalis Carter, 1882b: 285, pi. 11 figs Ectyodoryx (Myxillinae sensu van Soest) in the 2a-e (Venezuela, Bahamas, Pacific coast of Mexico). - former, and Plumohalichondria in the latter (cited Carter, 1886a: 52 (Port Phillip Bay). as Pronax Gray, and thus mistakenly used by Myxilla isodictyalis. -Dendy, 1896: 30 (Port Phillip Buyl.-Whiteiegge, 1901: several modern authors: this is a junior homonym, 79 (Tuggerah Beach, NSW). Lissodendoryx isodictyalis. as de Laubenfels, 1936a: 63, pointed out, with the -Topsent, 1897: 456 (Indonesia). -Topsent, 1925a: 701 (Naples, usual superfluous replacement name Grayax). But Monaco; comparative spicule measurements). -Topsent, 1936a: 18 van Soest only pointed to similarity in spiculation; (discussion). -Hechtel, 1965: 38 (Jamaica; comprehen- the presence of auxiliary, mostly echinating acan- sive synonymy, description, distribution, discussion). thostyles in Ectyodoryx, and in other myxillines ?Rutzler, 1965: 31 (N. Yugoslavia; description, ecol- is relevant. Incidentally, mentioned above, more ogy). - Wiedenmayer 1977a: 135, text-figs 141, 142, pi. I find van Soest's criterium insufficient, for main- 29 fig. 2 (Bahamas; further references, type data, descrip- taining the subfamilies Myxillinae and Tedaniinae tion, ecology). -Thomas, 1973b: 31, pi. 2 fig. 3 (Mahe, (not in Topsent's sense) for groups with tylote Seychelles). - Bergquist and Sinclair, 1973: 38 (New Zealand; larval versus oxeote or strongylote ectosomal megascleres, ecology). -Van Soest, 1984: 54, text-fig. 19, pi. 5 figs 2, 3 (West Indies; further synonyms, respectively. Some such megascleres in Myxilla description, disseussion, distribution). (though indiscriminately called tornotes by Koltun) Lissodendoryx similis Thiele, 1899: 18, pi, 5 fig. 10 are tylotes and their spined or mucronate modifi- (Indonesia). -Burton and Rao, 1932:331 (Burma). -Ali, cations, while corresponding spicules in Lissoden- 1956a: 293 (Madras harbour). -Ali, 1956b: 553 (Madras doryx are often oxeote (see Lundbeck, 1905; harbour; ontogeny). SPONGES FROM BASS STRAIT 99

variable. Spiculation within populations may be 'A\ variable.

Description. Small, ovoid-compressed and bean- shaped, 20 x 18 x 12 mm, and 30 x 18 x 10 mm. Base of attachment narrow in small specimen, on most of broadside of larger one. In life light olivaceous brown (10 YR 5-6/4), now light yellowish grey to beige (2.5 Y 7-8/2-4). Softly spongy, easily torn. Surface smooth to finely scultptured. Relief a continuous pattern of vermiculate, dendritic subdermal canals, which occasionally anastomose. Canals 0.2-0.5 mm wide, slightly raised dense partitions 0.5-1 mm wide. Dermis over weak depres-

Figure 64. Lissodendoryx isodiclyalis, spicules, a. Range sions contains fine reticulum with interstitial ostia. and example of tylotes, x 263. b. Range and example of More prominent oscules conical to irregularly styles, x263. c. Range of small chelae, X788. d. Range lipped, 2-3 mm wide, angular, apical and single on of large chelae, x 788. e. Range of small sigmata, x 788. smaller specimen, subapical and contiguous on f. Range of large sigmata, x788. larger one. Smaller oscules irregularly scattered on sides, some round, others sinuous, or as contorted

fissures. Dermis detachable is flakes, with scattered Hamigera ternaiensis Thiele, 1903a: 952, fig. 18 sand grains. (Indonesia). Choanosome dense, traversed by few irregular exhalant canals of angular section, Lissodendoryx ternaiensis. -Lundbeck, 1905: 173 or like (transfer, discussion). - Burton and Rao, 1932: 331 fissures. (India). Ectosome not well defined, commonly finely

Lissodendoryx sinensis Br^ndsted, 1929: 228, text-fig. cavernous; thickness variable, mostly around 260 5 (Formosa Strait, China). -Burton, 1937: 26, pi. 3 fig. /tin; contains thick fascicles of tylotes, subtangen- 19 (retained, diagnosis). tially, obliquely or perpendicularly to surface. Sur-

Materia/ examined. Station BSS 181, two adjacent speci- face crowded to packed with microscleres. Chelae mens (F52019). Eight West-Indian Syntypes of HiSchon- concentrated in outer third to half of ectosome, sig- dria isodiclyalis Carter in Liverpool (see Wicdenmayer, mata equally numerous throughout. Main skele- 1977a: 135, 258). Living populations and 4 self-collected ton a subisodictyal reticulation of megaseleres, specimens, Bahamas (Wicdcnmayer, 1977a: 135). Hypo- singly or in bundles of up to 4-5 spicules. Occa- types of Dendy, 1896 (Myxilla isodiclyalis, Port Phillip sional longer, sinuous tracts. Structure frequently Heads and Sorrento jetty; NMV). obscured by confused arrangement. Megaseleres Diagnosis. Thinly encrusting to massive, lobate to mostly styles, fewer tylotes regularly scattered. digitate, rarely repent-ramose. Lobes commonly Abundance of chelae in choanosome irregular, conical. Common in shallow waler, fouling com- always greatest around canals. munities, often epibiotic on vegetation (mangrove- Spiculation: (I) Tylotes, fusiform, mostly roots, sea grass, algae) and various invertebrates. straight, with spherical to oblong tyles (tapering Softly spongy, very compressible, easily torn. inward) frequently unequal in size and shape, Colour variable: often greenish, but also yellow- 168-/55-207 x 2.5-2. 9-3.4 jtm. (2) Fusiform styles ish, brownish, greyish lavender, lighl purple. to styloids (anisoxea with blunt to pointed shorter Oscules conspicuous, mostly apical on lobes. Sur- ends), straight but more frequently bent at one- face smooth to finely papillate or rugose. Dermis third from the base, 139-/59-180 x 2.6-5.5-3.8 Ihin, detachable, semidiaphanous over dendritic- /tin. (3) Arcuate isochelae of two size ranges vermiculate canals and punctiform vestibules, with (smaller ones less numerous, with proportionately numerous tylotes loose or in bundles, often retic- wider profile, more strongly arched): ulate. Microsleres may be crowded at surface. 22.2-25.5-24.9 X 6.4-7.5-9.2 /mi, (chord and pro- /-1 Choanosome bread-like. Main skeleton irregu- file width), and 1 2.4-/5. 8. 1 x 3.5-5.7-7.6 /im. lar, vaguely reticulate, subisodictyal, with (4) Sigmata of two imperfectly segregated size megaseleres single or in bundles, with some longer, ranges, smaller ones less numerous, 29.3-50.9-34.4 sinuous tracts, often masked by confused disposi- pan in length (chord), and 15.7-/7.7-20.3 /un; rare tion. Abundance of microscleres in choanosome intermediates, 23.4 and 24.7 /tm. 100 F. WIEDENMAYER

supposed differences, particularly in dimen- Remarks. Dendy's hypotypes (1896) are consider- The sions of spicules, cited again by Burton and Rao ably larger, more massive than the new specimens. (1932: 331) to justify the retention of L. similis Except for RN 773, which is virtually smooth, and Thiele for Indonesian populations, have been RN 872, with strong relief, surface characters are be insignificant by Topsent (1925a, consistent with the small specimens described above shown to and with the material from the Bahamas described 1936a). Styles in the new material from Bass Strait are by me earlier. Oscules, however, are mostly con- thin (5-6 /tin being common in previously siderably larger than in the specimens from Bass unusually descriptions). But this may be an expres- Strait. Another special trait, not observed in the published of the atypical environment. new material, is the presence in Dendy's dissected sion jacksoniana (Lendenfeld, specimens, of a palisade-like zone of parallel incur- "Lissodendoryx" Port Jackson, Sydney, redescribed and rent canals, 5-10 mm deep below the ectosome, (pi. 1888), from revised by Hallmann (1914c), is a good species, 1 1 fig. 2), similar to those in Tedania anhelans and of small strongyles Forcepia biceps noted above. Three of Dendy's characterised by the addition It has been recorded specimens are from Sorrento Jetty, the remaining to the choanosomal skeleton. isodictyalisjacksoniana) ones are from depths of 11 to about 25 m. This from South Africa (as L. indicates that Australian records of this species by Button (1936a). (Shaw, 1927a, described seem to favour shallow, sheltered conditions like Lissodendoryx dendyi southern most records from New Zealand, the Indo-Malayan in Hamigera) is another good species from Region, the Indian Ocean, the Mediterranean, and Australia, and is atypical for the genus. The only from the Western Atlantic. specimen known is small, erect, compound-clavate, The question, whether biogeographic gradients with a characteristic papillate surface. The papil- sizes, osculiferous ones being larger. are apparent in the variable spicular traits of this lae are of two smaller have a single apical supposedly cosmopolitan species, is still unresolved The inhalant ones (van Soest, 1984: 56). Styles in West-Indian speci- "pseudostium" over a central porocalyx underlain mens have usually been figured or described as sub- by a separate concentric vestibule. The ectosomal tylostyles (Hechtel, 1965: 39; Simpson 1968a: megascleres are restricted to the inhalant papillae, text-fig. 11; Wiedenmayer 1977a: text-figs 141, 142; where they are regularly radially arranged, while van Soest, 1984, text-fig. 19), but Topsent (1925a: the interstitial dermis contains only crowded arcu- 701f.) recorded tapering bases in styles of his own ate chelae with atrophied alae. Most choanosomal material from Guadeloupe (Tedania lepioderma megascleres are slightly asymmetrical strongyles like Topsent, 1889), in Carter's type material (Halichon- those in the inhalant papillae. Styles are rare in the dria isodictyalis), and in his specimens from Naples deeper choanosome. There are no sigmata. The and Ambon. The presence of two categories of closest relative is L. papillata (Dendy, 1922: 90) both chelae and sigmata in Australian and Indone- from the Indian Ocean, which is less specialised, sian specimens, like in van Soest's (1984), is sig- more typical for the genus, but also lacks sigmata. nificant. Such a condition is rarely if ever reported in other descriptions (Topsent, 1925a: 703, men- Eetyodoryx Lundbeck tions two categories of sigmata in a specimen from Eetyodoryx Lundbeck, 1909: 444. -Topsent, 1928: 53, Venice) and may have been overlooked in some 55 (definition in Myxillinae). — de Laubenfels, 1936a: 84 cases. Burton and Rao (1932: 331) re-examined (definition in Myxillinae), 85 (comparison with Hymedes- Carter's (1886a) and Dendy's (1896) specimens from mia). — Levi in Brien et al., 1973: 612 (definition in Myx- Phillip and found them to agree with L. Port Bay illidae). — Van Soest, 1984: 71 (in discussion of ternatensis (Thiele). They retained the latter as a Myxillidae). valid species, to which they added L. sinensis Diagnosis. Distinct from Lissodendoryx by the Br^ndsled as synonym, the only diagnostic charac- addition of echinating acanthostyles. Sigmata may ter being the two sizes of both chelae and sigmata. be absent. Burton (1937: 26) described L. sinensis as a good species, without reference to the decision in Burton Eetyodoryx maculata Hentschel and Rao (1932). There may have been a delay in the former manuscript. This is the publication of Plate 11 figures 3, 4, plate 30 figure 5, only record of L. isodictyalis with repent-ramose shape. Chelae and sigmata are again of two sizes. text-figures 65, 66 latter trait As van Soest has shown that the may Eetyodoryx maculatus Hentschel, 1911: 342, text-fig. be present in West-Indian populations of L. isodic- 30 (Albany, WA). tyalis, there is no need for L. ternatensis and L. Merriamium maculatum. — de Laubenfels, 1936a: 83 sinensis. (transfer only). SPONGES FROM BASS STRAIT 101

which also occur radially around inhalant areoles and erect in peripheral choanosome. Main skeleton indistinctly plumo-reticulate, with smooth principal styles and often echinating acanthostyles. Chelae of two sizes, no sigmata.

Description. F52020 thickly incrusting to cushion- shaped, with sinuous margin, 3.7 x 5 cm wide, 0.5-1 cm high. F52021 massive, erect-lobate, somewhat compressed, with angular apical lobes, 3 x

4.7 cm wide, 5 cm high. F52020 orange in life, F5202 1 vermilion (5 R 5-6/11); both now cream to light yellowish beige. Sofly spongy, easily torn. Figure 65. Ectyodoryx maculata. F52020, perpendicular Surface with pattern of saucer-like depressions, section ( x 3.9, surface above) showing very cavernous 1.5-2 mm wide in F52020, about 1 mm wide in interior. F52021, with intervening raised areas about equally wide. Pattern regular except margin, in F52020; in F52021, only distinct on one side. Elsewhere, pattern obscured by dendritic and anastomosing rugae and depressions mostly longitudinally aligned. Round depressions correspond to wholly or partly closed vestibules, atria or mouths of radial canals, all of similar dimensions, communicating in thoroughly cavernous choanosome. Some rounded depressions roofed by membrane bearing indistinct ostia. In others membrane bears iris-like aperture which may be retracted to sharp skeletal rim. Hence differentiation into incurrent sieves and small oscules. Some oscules elongate to slit-like along margin of F52020 and on apices of F52021, here communicating with wider longitudinal canals with angular cross-section. Areolate (swollen) skeletal rim with radial structure in some of round depressions. Main skeleton an often irregular reticulation of fibres, with variably conspicuous spongin. Sometimes distinction of thicker, more continuous branching principal fibres, with little orientation, and mostly oblique, frequently branched anastomoses; the former around 50 urn thick, the latter about 20 /xm. Meshsizes 170 to 430 pun. Fibres cored to packed with subparallel to semi-plumose Figure 66. Ectyodoryx maculata, spicules from F52020. megascleres, chiefly principal styles, always with a. Range and example of principal styles, x 263. b. Range some acanthostyles, also as echinators. Megascleres c. Range and and example of acanthostyles, x263. also interstitially in variable amounts, occasionally x263. d. example of strongyles and subtylostrongyles, obscuring meshwork. Chelae numerous Range of massive chelae, x 789. e. Range of frail chelae, interstitially. x789. Ectosome not distinct, with crowded to packed chelae at surface, and commonly subtangential

Material examined. Station KG 1, one specimen (F52020); loose bundles of diacts. Bundles of diacts also in station KG 7, one specimen (52021). peripheral choanosome, erect or oblique to surface, around vestibules and atria, in Diagnosis. Cushion-shaped to massive-lobate. most conspicuous radial and umbellar pattern. Softly spongy, easily torn; orange to vermilion in regular Spiculation: (1) Principal choanosomal styles, life. Surface regularly areolate; inhalant and straight to curved or bent, 191-205-223 x exhalant areoles little differentiated, with raised rim 2.6-5.6-7.9 itm. Choanosomal acanthostyles, and central depression. Choanosome thoroughly (2) 100-/72-124 x 4.7-7.9-10.3 /im. (3) Dermal and cavernous. Ectosome indistinct, with crowded peripheral strongyles and subtylostrongyles, occa- chelae and subtangential bundles of strongyles, 102 F. WIEDENMAYER

sionally styloids, often with elongate inflation at Diagnosis. "[Anchinoidae] in which the one end, other end strongylate, tornote or choanosomal skeleton consists of columns of acan- mucronate, subterminally slightly swollen in some, thostyles mixed with ectosomal oxeotes, echinated

283-292-306 x 1.8-3. 7-4.7 /an. (4) Arcuate chelae by a special category of acanthostyles; microscleres of two categories, (2) larger ones of normal type, arcuate isochelae." (Van Soest, 1984: 86.). with strongly arched shaft, length (chord) Phorbas cf. lenacior Topsent 23.9-26.2-28.4 /im, total width in profile 9.5-77-12.4 /mi, thickness of shaft 3.3-4.7 /an; (b) Plate 11 figures 5a, b, plate 30 figure 4, smaller ones with weak, gently arched shaft and text-figure 67 much reduced alae, length 17.6-2/. 4-21.2 /an, total width 5.2-6.7-8.3 /an, thickness of shaft 0.9-1.9 Selected synonymy of sibling species and subspecies: /

(1924) and Lissodendoryx dendyi (Shaw , 1927a; see end of Remarks on Lissodendoryx isodiayalis, a b c d e above). n r\ Anchinoidae Topsent, 1928

Synonym. Phorbasidae de Laubenfels, 1936.

Remarks. Phorbasidae was introduced by de

Laubenfels (1936a: 62) without reference to its senior synonym Anchinoidae (the type genus of which he included in Phorbasidae, synonymising it with Phorbas). It can be surmised that the seniority of Phorbas prompted de Laubenfels' action. Phorbasidae has been adopted by some fol- Laubenfels' system, lowers of de most recently by \j Hoshino (1981), but also by van Soest (1984: 86). However Levi (in Brien ct al., 1973) and his col- laborators, Bergquist (1978) and Pulitzer-Finali (1983) have used Anchinoidae.

Phorbas Duchassaing and Michelotti

Phorbas Duchassaing and Michelotti, 1864: 91. — de Laubenfels, 1936a: 63 (definition). — Bergquist, 1961b: 180 (discussion). -Van Soest, 1984: 86 (definition, description of type species), 88 (discussion). Anchinoe Gray, 1867a: 535. -Topsent, 1928: 56, 57, 61. — de Laubenfels, 1936a: 64 (synonymised with Phorbas). -Levi in Brien et al., 1973: 613 (definition). - Figure 67. Phorbas cf. tenacior. spicules, a. Range sub- Van Soest 1984: 88 (synonymised with Phorbas). of tylotes and subtylostrongyles, with example of the latter, Suberotelites Schmidt, 1868: 12. -Topsent, 1938a: 12 x263. b. Enlarged ends of a subtylote, x 788. c. Large (discussion, type species re-examined, synonymised with acanthostyles, x d. Anchinoe). 263. Small acanthostyles, x263. e. Range of chelae, x788. SPONGES FROM BASS STRAIT 103

with crowded chelae and subtangential to irregu- shafts, 15.3-/5.4-20.7 /mi in length (chord), larly bundled (subtylo-)strongyles. Choanosome 5.1-7.9-9.3 urn width in profile, 1.1-2.2-2.5 /mi dense, collagenous, with some sand. Main skele- in thickness of shaft. ton vaguely columnar to confused, with stron- Remarks. Measurements for megascleres gyles/subtylotes/subtylostrongyles and two sizes of in Mediterranean specimens acanthostyles. Arcuate chelae of one kind. are generally somewhat higher as published, but there is a good agreement Description. Capricious elongate shape. Surface with Boury-Esnault's figures, except for only slight thrown into lacunae, elongate partly longitudinally overlap in the small acanthostyles. Their smooth partly transversely, some straight, others curved or megascleres are generally pure strongyles only, but Topsent crooked; their section round, semicircular to cir- (1936a) noted occasional weak distal swellings. cular. Raised areas rugose and lobulate, mostly in Mediterranean specimens are usually encrust- cerebroid pattern. Some portions of lacunae sep- ing, of "Hymedesmia-type", but massive, tate or traversed by thin bridges; some ends of lacu- "/4rtc7?//?oe-type" specimens were reported by Top- nae communicate through round to elongate win- sent (1925a). The colour of the surface in Mediter- dows. Length 7 cm, width at the base 3 x 3.5 cm. ranean specimens is commonly blue (deep, to Base was attached to chaetopterid worm tubes, greyish), that of the choanosome being yellowish to ochre. However, a minority tapers slightly to lobulate apex. Lacunae 0.5 to 1 of yellowish speci- cm wide and up to 5 cm long, raised areas about mens occurs according to Pulitzer-Finali. Topsent (1936a) regarded tenacior as 1-2 cm wide, with rugae and lobules 1 -3 mm wide. a variety (subspecies) Dull, greyish orange-yellow in life (10 YR 5-6/6), of coriaceus, which is also common now, in alcohol, faded (10 YR 6-7/4-6). Firmly in the same area, because the latter differs only in spongy to leathery. Walls of lacunae microscopi- lacking chelae, in having smooth megascleres more cally mostly smooth, glabrous, except some trans- decidedly as subtylotes, and in its surface being variable verse nervures and finely papillate pattern in some more in colour (yellow, purplish sienna areas. Subdermal round vestibules, 200-1000 /mi brown, grey, bluish grey, whitish, in thin crusts; wide, numerous especially in depressions between pink, pink with yellow spots, off-white, in ramose rugae and lobules, with interspersed oscules of simi- specimens). Both display the same differentiation into lar dimensions. Crowded, very fine ostia indistinct thin incrustations and a much rarer ramose morphotype. All traits in depressions, over vestibules, also in fields of other are shared, particularly yellowish irregular outline on top of rugae and lobules. Scat- the choanosome, the glabrous ("waxy") tered and clustered sand grains everywhere in surface, the thick leathery ectosome dermis. (thicker and better differentiated than in the Aus- Main skeleton chaotic, some vague strands of tralian specimen). Boury-Esnault (1971) segregated two as species the megascleres discernible, chiefly of bundled stron- the on grounds of morphologi- differences being consistent with gyles and subtylotes, with few large acanthostyles; cal ecological (depth-range) segregation, though contiguous. the latter also echinating with more numerous small obviously constitute sibling pair. acanthostyles. Strands usually meandrine to con- They a The ramose specimens of tenacior from Naples were not voluted, frequently crowded to almost confluent. described in detail by Topsent (1925a), as to exter- Megascleres of all sorts also scattered at random, those of coriaceus from as microscleres. Yellow spongin discontinuous, nal shape, but Monaco described as having sparsely divided, fleshy, enveloping several contiguous strands, without were branches, beset by low, frequently inter- clear outline. angular rupted ribs, or by irregular, low tuberosities, hence Ectosome indistinct, with crowded chelae at sur- apparently quite similar to the present specimen. face, subtangential and vaguely umbellate arrange- It seems likely that the single new record for Aus- ment of strongyles (the latter especially around tralia represents a distinct species or subspecies, but vestibules). Sand grains more numerous below sur- it seemed unwise to name it as such before any face, scattered in dense, collagenous choanosome. more material is found. Spiculation: (1) Smooth strongyles (rare) mostly modified as subtylostrongyles and subtylotes, the latter commonly asymmetrical, 212-2/9-228 x Order Haplosclerida Topsent, 1928 2.3-2.5-3.3 /tm; (2) two categories of acanthostyles, (a) larger ones often gently curved, 146-/54-166 Callyspongiidae de Laubenfels, 1936 x 7.2-5.9-10.2 (with spines), (b) smaller ones nm Callyspongia Duchassaing and Michelotti straight, 67-77-80 x 6.7-9. 3-\ 1.2 /mi (with spines); (3) arcuate chelae with strongly curved [Tuba] Duchassaing and Michelotti, 1864: 44 (junior - -

F. WIEDENMAYER 104

(synonymised will Bergquist and Warne, 1980: 24 homonym of Tuba Lea, 1833; see Wiedenmayer, 1977a: Callyspongia). 100, under Spinosellu). 789. -Burton. 57.- Euplacella Lendenfeld, 1887a: 741, Callvspongia Duchassaing and Michclolti, 1864: synonymisei 597 (definition, resume, discussion, (synonymy, references, discus- 1934a- Wiedenmayer, 1977a: 90 1936a: 59 (discus with Placochatinah-de Laubenfels, Warne, 1980: 24 (synonymy, defi- sion). -Bergquist and species).- Van Soest sion invalid designation of type resume). -Van Soest, 1980: 46, 56, 1 10 nition, discussion, (definition, retained Li and 1980: 110, text-fig. 39 (definition, discussion with subgenera Callvspongia Callvspongiidae). Spinosella). 790. Placochalina Lendenfeld, 1887a: 741, -di 1870: 35.-Lendenfeld, 1887a: Cladochalina Schmidt, ol type Laubenfels, 1936a: 59 (subsequent designation 768. -Burton, 1927a: 295 (designation of type species, species). revision of Lendenfeld, 1887a). [Siphonella] Lendenfeld, 1887a: 742, 808 (junior homo' IPlatychalina Ehlers, 1870: 21. -Burton. 1936a: 143 Siphonella Macquan, 1835). -Burton, 1934a: S«i Homoeodictya).-\'zn Soest, 1980: 60 nymol (as synonym of in (revision). -Van Soest, 1980: 118 (genus dubium (probably senior synonym of Callvspongia, subgenus Haplosclerida, wrong type species). Spinosella), Euchaliim Lendenfeld. 1887a: 742, 816 (type species: TubulodigitUS Carter, 1881a: 376.-Burton, 1934a: 536 Euchatina typica Lendenfeld, 1887a, by typonymy). (neotype of type species redescribed). Burton. 1934a: 540 (invalid designation of Euchalinu Pdtuloscula Carter* 1882b: 365. -de Laubenfels, 1936a: rigida Lendenfeld, 1887a, as type species), -de Lauben 57, 59 (definition, synonymised with Toxochalina).- 1936a: 47 (Burton's mistake corrected). -Van Soest. Wiedenmayer, 1977a: 90 (references). ffels, 118 (genus dubium in Haplosclerida, wrong typo Siphonochalinu sensu Ridley, 1884a: 401. and of 1980: species). authors (part). Chalinopsilla Lendenfeld, 1888: 104 (nomen novum for. nee Siphonochalina Schmidt, 1868: 7 (see van Soest, Chalinopsis Lendenfeld, 1886, nee Schmidt, 1870).- 1980: 113). Lendenfeld, 1889b: 124 (with Daciylia, Paraspongia, Toxochalina Ridley, 1884a: 402. - Ridley and Dendy, Psammoclema as synonyms). -Laubenfels, 1936a: 59 1887: 49. -Dendy, 1905: 139 (definition). -Topsent, (synonymised with Dactylic).— Bergquist and Warne. 1928: 66. -Van Soest, 1980: 110 (synonymised with 1980: 33, 34 (definition, distinct from Dactylia). Callyspongia').— Bergquist and Warne, 1980: 24 (syn- Pulitzer-Finali, 1982b: 123. onymised with Callvspongia). Arenosclera (nomen novum Spinosella Vosmaer, 1885 (in 1887): 342 Diagnosis. Callvspongiidae with distinct ectosomall Michelotti, 1864, ncc Lea, Duchassaing and I for Tuba skeleton, either as tangential net of primary and 1833). -Levi in Brien et al., 1973: 618 (synonymy, dis- secondary (sometimes tertiary) spongin fibres, or cussion).- Wiedenmayer 1977a: 100 (synonymy, defini- as abrupt peripheral condensation of main skele- tion, discussion).- Van Soest, 1980: 56 (definition as intense branching and intercalation of subgenus of Callvspongia), 64 (discussion). -Bergquist ton, by Main skele- and Warne, 1980: 24 (discussion). primaries and crowding of secondaries. Dactylia Carter, 1885c: 309. -de Laubenfels, 1936a: ton may contain ascending fibre fascicles ending 59 (discussion, revision). -Van Soest, 1980: 1 10 (defini- in spines at surface. Magascleres oxea or strongyles. tion, probably a synonym of Callyspongia).- Bergquist generally coring to packing fibres, occasionally also 1980: 34 (definition, distinct from and Warne, interstitial, in some species also in erect brushes at Chalinopsilla). surface. Toxa may be present, occasionally alone. [Chalinopsis] Lendenfeld, 1886e: 569Gunior homonym Spicular skeleton may be reduced or suppressed, of Chalinopsis Schmidt, 1870). and aspiculous fibres may be regularly cored by Dasychalina Ridley and Dendy, 1886: 329 (type spe- foreign detritus. Foreign inclusions may also occur cies: D. fragilis Ridley and Dendy). - Burton, 1932a: 278 (designation of type species, diagnosis, discussion). in spiculiferous fibres, interstitially and on surface. Wiedenmayer, 1977a: 100 (discussion as synonym of Remarks. Callyspongia and Spinosella, which I had Spinosella). as distinct genera in 1977a, were treated Chalinopora Lendenfeld, 1887a: 740, 764 (type species: regarded - van Soest (1980). Chalinopora typica Lendenfeld, 1 887a, by typonymy). as subgenera of Callyspongia by Burton, 1927a: 293 (invalid designation of Acervocha- Platychalina Ehlers (1870) might eventually replace

lina claviformis Carter, 1 886, as type species). - Burton, Spinosella, pending re-examination of its type spe- 1934a: 535 (ditto). cies, Spongia foliacea Esper, according to van Soest Lendenfeld, 1887a: 740, 770. -Burton, Chalinella (1980: 60). Esper's type material, however, is very 1927a: 296 (revision, synonymised with Cladochalina). probably lost according to my recent enquiries in Ceraochalina Lendenfeld, 1887a: 740, 778 (type spe- Erlangen, Frankfurt (Senckenberg Museum) and cies: Ceraochalina lypica Lendenfeld, 1887a, by Munich (Zoologische Staatssammlung), except for typonymy).- Dendy, 1905: 152 (definition). -Burton, slides in Berlin (ZMB), and 1934a: 540 (invalid designation of Cladochalina nuda some by Thiele East in Ridley, 1884a, as type species). -Van Soest, 1980: 118 the schizoholotype of Spongia papyraeea

(genus dubium in Haplosclerida, wrong type species).— London (BMNH). Both van Soest and 1 have over- SPONGES FROM BASS STRAIT 105

looked that Spinosella, if retained as subgenus, is Wiedenmayer, 1977a: 95). Though detritus tends replaced by Cladochalina Schmidt (type spe- best to interstices in spiculiferous ones, it may incrust cies: Tuba armigera Duchassaing and Michelotti, the surface of both. Different species with vestigial 1864: Schmidt, 1870, subsequent designation by the surface of both. Different species with vestigial Burton, 1927a: 295). If the subdivision of the genus spiculation are known both from the West Indian

Callyspongia were to be maintained, it would seem region and from the Indo-West Pacific region. It advisable to treat Euplacella (which van Soest, is possible that the loss of megascleres is an 1980, regarded as a distinct genus) as a third subge- acquired character within the Callyspongiidae, and nus. In this case, however, it would have to be that the presence of toxa, though facultative and replaced by its senior synonym (by page priority) individually variable, is an ancestral trait shared

Ceraochalina. The overlooked type species of the with other haplosclerids. This is suggested by the latter, C. lypica, is clearly a junior synonym of new species of Callyspongia described below, in Callyspongia serpentina (Lamarck) ( = C. com- which all fibres are aspiculous, and in which inter- munis of authors), as my survey of Lendenfeld's stitial toxa may be present in variable abundance; types in Sydney and London revealed. The distinc- but also by the species (species complex?) tion of the two subgenera Callyspongia and Callyspongia ramosa (see below), which is highly Cladochalina is not always practical with Indo- polymorphic, also with regard to some skeletal West Pacific and Australian material. If Burton's traits, particularly in the occasional presence of (1959: 225) synonymy of Callyspongiu spinosissima toxa. Toxa also occur in some species of Renieri- (Dendy) is accepted, we would have morphotypes dae ("Orina", "Toxadocia"), Niphatidae (Micro- with coarsely spiny surface {spinosissima) and mor- xina, of which Hemigellius is here considered a photypes with smooth surface (subcylindrica). synonym), and Oceanapiidae ("Biminia") (see van

Callyspongia truncate! Lendenfeld (1887a), (with Soest, 1980). It could be argued, however, that both most of his "species" of Siphonochatina as syno- loss of megascleres and of toxa (and, in analogy, nyms) likewise comprises morphotypes of both of sigmata in non-callyspongiid haplosclerids) are

sorts. Another case in point is Callyspongia tuber- due to parallelism, and occurred in lineages link- culata (Lendenfeld 1887a, comprising 4 of his 5 ing various ancestral poecilosclerids and derived, "species" of Siphonella as synonyms) in which the polyphyletic Dictyoceratida. Strong reduction of characters of the nominate subgenus (annular swell- megascleres, however, is much less common in non- ings, bullae, smooth portions) are often combined callyspongiid haplosclerids, and occurs, for with tubercles and spines. The retention of Cer- instance, in the West-Indian Amphimedon com-

aochalina as a subgenus is impractical, because its planata (see van Soest, 1980: 31), and in the Aus- only diagnostic trait (the presence of erect spicule tralian Amphimedonl crassiloba (Lamarck)

brushes at the surface) is not consistently developed (choanosome only).

in at least one species: it may or may not be present Presence absence of spicules and of foreign in the polymorphic Callyspongia ramosa. Bergquist detritus are here interpreted as poorly correlated, and Warne (1980) distinguished Chalinopsilla and diatoxonic traits and as differences of degree within Dactylia on the grounds of foreign material coring the Callyspongiidae, with Chalinopsilla sensu Berg- the primary fibres of the latter, but being com- quist and Warne, and Arenosclera Pulitzer-Finali pletely absent in the former. This distinction has as extremes. Paraspongia Carter, which Lenden- a major nomenclatorial drawback: the type species feld (1889b: 145) regarded as an "obsolete" syno- of Chalinopsilla (by subsequent designation in nym (relating to Lendenfeld's nomenclatorial Bergquist and Warne, 1980, not by monotypy), C. idiosyncrasies, see Wiedenmayer et al., in press) of dicholoma (Lendenfeld, 1886e), has foreign bodies his Chalinopsilla (over which it would, as such, (chiefly foreign spicules), both in primaries and have priority), is very poorly known. I suspect it secondaries, according to Lendenfeld (1889b: 143, to be a junior synonym of Phoriospongia. Dasychalina, as revived by Burton (1932a) for pi. 3 fig. 1 1). But neither such a distinction within Ridley and Dendy, and P. aspiculous species, nor a separate genus for Pachychalina fragilis aspiculous callyspongiids seems necessary to me validissima Thiele (1905), merely represents structure of from a taxonomic point of view. In my experience extremes in the variable skeletal diagnosis above. with Australian callyspongiids, including many type Callyspongia outlined in the new Callyspongia specimens, foreign detritus occurs in both Fibres packed with spicules occur in Bergquist and spiculiferous and aspiculous species, to a variable conica (Br^ndsted), C. irregularis and C. slellala Bergquist and Warne (Berg- degree. I had earlier noted detritus in the fibres of Warne, 29, 32). All three spe- Callyspongia pallida (as C. fallax forma debitis in quist and Warne, 1980: 27, 106 F. W1EDENMAYER

In dermal skeleton, primaries around cies have abundant interstitial megasclercs in the ous sediment. around 20 fim, tertiaries choanosome, as has C. persculpla described below. 45 nm thick, secondaries around 8 /mi. Corresponding meshsizes around 1 Callyspongia asparagus (Lamarck) mm, 170-350 /im, and 45-85 /tin, respectively. with incomplete segregation of Plate 11 figure 6, plate 31 figures 2, 3, Main skeleton primaries, 95-130 /tin, spaced 580 to 1200 urn, and text-figure 68 secondaries, 47 to 62 urn thick, spaced 290 to 870 distributed Spongia asparagus l.amarck, 1814 (1813-1814): 447 /an. Spicules scanty, confined to fibres, (Australian Seas, Peron and Lesueur collection). irregularly, more numerous in some fibres, almost Chalina asparagus. - Ridley, 1884a: 405 (schizotype in lacking in others; indistinctly aligned or randomly BMNH mentioned, comparison with Pachychalina oriented, coring (then often broken) or occupying macrodactyla). whole thickness of fibre; clearly visible or barely Clarlochulina asparagus. -Topscnt, 1932a: 106, pi. 6 discernible. fig. 7 (description of type fragments, discussion). Spiculation: Strongyles transitional to tornotes one specimen 1.5-2./-3.1 Material examined. Station BSS 187, and strongyloxea, 63.5-72.2-85 x fim. (F52024). Remarks. See Callyspongia pergamentacea, below. Diagnosis. Sparsely branched; branches knotty, with blunt lips, 0.5-1 cm thick. Firmly spongy, Callyspongia bilamellala (Lamarck) pink in life. Oscules numerous, unilateral or Plate 1 1 figure 7, text-figure 69 bilateral, around 1 mm wide. Surface smooth, with dense tangential reticulum. Main skeleton lax, Spongia bilamellala Lamarck, 1814 (1813-1814): 434 (part, typical form only, not var. Australia (as Southern coarse, irregular. Choanosome scanty. Spicules li; Ocean), Peron and lesueur collection). small, scanty, irregularly distributed in fibres only. Cavochalina bilamellala, — Carter, 1885d: 287 (part; Description. Three unequal branches (one now dis- Port Phillip Heads). sected) divergent from one point and curved in one Pachychalina bilamellala. -Dendy, 1895: 242 (Port - plane. One branch proximally enveloped by Phillip Heads). Whiteleggc, 1902b: 287. Ctadochalina bilamellala. —Burton, 1927a: 509. cerebroid to lactulose, coarsely sandy, slightly com- Cribroehalinu bilamellala. — Topsenl, 1932a: 68 pressible mass, possibly an undescribed Psain- (redcseriplion of Lamarck's types of typical form). moclema. Same mass supports frondose bryozoan. nee Spongia bilutuelluta var. /} Lamarck, 1814 (fide Branches of Callyspongia mostly compressed, Topscnt, 1932a: 69, correctly Eehinodictvum somewhat knotty, 0.5 to 1 cm thick; one terminally mesenlerinum). more compressed, almost spatulate. nee Echinodiclym bilamellalum. — Ridley in Ridley and

Pink in life now faded, mostly to yellowish beige, Duncan, 1881 (Spongia bilamellala var. (3 redescribed

partly still pale red-brown. Firmly spongy. from schizotype). — Topseni 1932a: 69 (for above). Surface smooth, also microscopically, finely Euplacella australis Lendenfeld, 1887a: 789 (Torres Strait: junior secondary homonym of Dactylochalina aus- mottled by vestibules in peripheral meshes. Oscules tralis Lendenfeld, 1886L). -Burton, 1934a: 598 (Great numerous, confined to narrow upper face, round Barrier Reel). to elongate, 0.5 to 1.5 mm wide. Superficial retic- Pachychalina australis. — Whitelegge, 1906: 457 (off ulum continuous except on branch tips, here with C'oogee, off Bulgo, off Wollongong, NSW). — Burton, coarse main skeleton exposed. Finer superficial 1927a: 509 (revision).

meshwork mostly obscured by fine clacareous sedi- Euplacella frondosa Lendenfeld, 1887a: 789, pi. 21 fig. ment, only darker primary tangential fibres stand- 36 (Port Jackson, NSW; Gulf St Vincent, SA). ing out. Pronounced contrast between dense Euplace/la mollissima Lendenfeld, 1887a: 790, pi. 21 surface reticulum and lax, open, relatively coarse fig. 37 (Westernport Bay, Vic.).- Whitelegge, 1902a: 215 and irregular meshwork of interior. Flesh scanty, tympanising some of meshes, contains fine calcare-

czr ^zj

Figure 68. Callyspongia asparagus, spicules (strongyles Figure 69. Callyspongia bilamellala, spicules, x782 and strongyloxea), x742. Oxea in fibres, b. Interstitial oxea. SPONGES FROM BASS STRAIT 107

(coast of NSW). — Burton, 1934a: 598 (synonymy, types Surface optically smooth except relief mentioned in BMNH re-examined; Great Barrier Reef). above. Oscules inconspicuous in concavity, minute, var. dura Lendenfcld. Placochalina pedunculata 1887a: not above 200 /tin in width, numerous, regularly fig. 38 (Port Jackson, NSW). -Lendenfeld, 791, pi. 21 scattered. Outer surface with coarse primary tan- 1888: 91.-Whitelegge, 1889: 182 (Coogee, Bondi, gential fibres just below surface, outlining rounded NSW). -Whitelegge, 1902b: 279, 287. meshes subdivided by indistinct finer fibres; secon- Placochalina pedunculata var. poculum Lendenfeld, daries and tertiaries mostly indistinct. Pattern 1887a: 792, pi. 21 fig. 39 (Torres Strait; lllawarra and obscured by (spicular Port Jackson, NSW).- Lendenfeld, 1888: 91.- crowded microconules Whitelegge, 1902b: 279, 287. brushes) and fine sediment. Placochalina pedunculata var. mollis Lendenfeld, In margins, regular radial primaries curve gently 1887a: 792, pi. 21 fig. 35 (Gulf St Vincent). -Lendenfeld. from mesial plane, meeting surface almost perpen- 1888: 92.-Whitelegge, 1902b: 279, 287 (revision of 2 dicularly, branching frequently in curves; some as syntypes). sudden intercalations; spacing 140-350 /xm, thick- Pachychalina punctata (nee Ridley and Dendy); ness 35-130 fim. Secondaries generally more Whitelegge, 1889: 182 (Port Jackson and Bondi Beach, crowded, spaced mostly around 125 ^m, up to 430 NSW).-Whitelegge, 1901: 69 (Tuggerah Beach, itm, parallel to surface; few branching or connected NSW). -Whitelegge, 1906: 458 (off Barranjoey, NSW). by short radial anastomoses. Thickness as for Material examined. Station KG 9, one specimen (F52025). primaries. Spongin abundant, meshes commonly Euplacella australis, E. frondosa, E. Type specimens of rounded, with radial widths often equal or below mollissima, Placochalina pedunculata varr. dura, pocu- those of fibres. All fibres conspicuously cored by lum, and mollis (AM and BMNH). Hypotypcs of spicules, core occupying fifth to third of thickness Cavochalina bilamellata: Carter, 1885d (BMNH); of in primaries, with at least 6 spicules abreast, in Pachychalina bilamellata: Dendy, 1895 (NMV); of 1-4 spicules abreast. Spic- Euplacella australis and E. mollissima: Burton, 1934a secondaries commonly (BNMH); of Pachychalina australis: Whitelegge, 1906, ules aligned to semiplumose, frequently fanning at Euplacella mollissima: Whitelegge, 1902a, Pachychalina junctions, occasionally disheveled, especially in punctata: Whitelegge, 1901, 1906 (AM). periphery. Peripheral skeleton frequently less regular, less Diagnosis. Stalked, calculate. Cup inverted- segregated from main skeleton where thick, spic- bilater- conical, often asymmetrical, compressed, ules dominating over spongin. Where more distinct, Outer sur- ally deeply incised and hence bilamellate. equal to 1-3 spicule-lengths in thickness, spicules inner face commonly coarsely rugo-reticulate, even more abundant, in vaguely umbellar to can- Externally sculptured surface concentrically ridged. cellate bundles, reminiscent of periphery in cups almost lipostomous, smooth ones with more Cribrochalina. conspicuous, numerous and regular oscules in con- Main skeleton interrupted in irregular intervals pinkish cavity. Rubbery, tough; dull light red to by similarly condensed /ones parallel to present grey in life. Surface smooth, but often microhispid. marginal surface, reflecting earlier growth stages. Ectosome with erect spicular brushes, umbellate to Spiculation: Fusiform oxea in fibres, curved to finely cancellate. Main skeleton regular, bilateral- symmetrically bent, 60.9-65-71.8 x 2.3-5-4 ,mi. primary symmetrical in fronds. Spongin abundant, Interstitial oxea and strongyies, 38.1-45.9-54.5 x fibres well cored by spicules, secondaries with thin 0.7-0.9 itm. spicular cores. Small oxea, smaller interstitially, Remarks. There is a pronounced affinity, in the with strongyies. ectosomal skeleton of this species, to the genus Cribrochalina. In the latter, however, erect spicu- Description. Stalked asymmetrical cup, roughly form continuous palisades along the inverted-conical. Stalk crooked, knotty, 2-2.5 cm lar brushes of the outermost secondaries, while in wide and 16 cm long, with rhizomatous base. edges Callyspongia bilamellata erect spicules at the sur- Height of cup 27 cm on higher side, 10 cm on oppo- face are grouped in isolated tufts. The synonymy site side. Wall mostly 0.5 to 1 cm thick, longitudi- suggested above should be qualified with regard to nally folded, with concentric ridges on inside, Euplacella australis, E. frondosa, and Pachycha- coarsely rugo-reticulate relief on outside. Rim punctata sensu Whitelegge. These specimens sharp, sinuous or obtusely notched. line, faint lina differ from the typical specimen described above radial striation throughout inside, only 1-2 cm deep in two traits, and could be regarded as a distinct below rim on outside. Dull light red to pinkish grey 6-8/4-6 subspecies or form. The two traits are: absence of in life (10R 7/2-8 to 10 R 8/8 inside, 10 R the rugo-reticulation on the outside (or convex side outside), now greyish yellow-brown (7.5-10 YR of single fronds) and presence on the inside or con- 6-7/2-4). Rubbery. -

108 P. WIEDENMAYER

cave side of conspicuous round oscules (width 1887a: 784 (part: specimen from Port Jackson only, fide

around 1 mm). Both forms, however, agree in the Burton, 1934a: 541). Pachychalina multiformis var. munaarensis Dendy, dominantly bilamellate to caliculate, stipitate 1889d: 79 (Tuticorin Pearl Banks, Gulf of Mannar). growth form, colour and consistency. Burton Ceraochalina multiformis var. munaarensis. — Dendy, (1934a: 498) distinguished the two as species (call- 1905: 154, pi. 7 fig. 2 (Gulf of Mannar, Sri Lanka). ing the typical form Enplacella mollissima, yet una- Dendy and Frederick, 1924: 500 (Abrolhos Islands, WA). ware of Topsent's redescription of Spongia ISiphonochalina crassifihra Dendy, 1889d: 82 (Tutico- bilamellata), but at the same time doubted that the rin Pearl Banks). latter was really distinct from E. austratis (correctly Chalina pulvinatae Lindgren, 1897: 481 (Java).— frondosa). Some growth forms of frondosa resem- Lindgren, 1898: 295, pi. 18 figs. 2, 16; pi. 19 fig. 7 (spelled ble Pachychalina punctata Ridley and Dendy, pulvina/us, description, discussion). Callyspongia pulvinatae. — van Soest, 1982: 90, text- which is a typical Cribrochalina. It differs from fig. 9 (Hong Kong). frondosa externally by having thicker, more irregu- Ceraochalina retiarmata Dendy, 1905: 152, pi. 10 fig. lar and lumpy fronds, with thick rims and frequent 4 (off Chilaw, Sri Lanka). perforations, larger, more widely spaced and T Ceraochalina cey/onica Dendy, 1905: 154, pi. 7 fig. irregularly scattered oscules, and harder con- 3, pi. 10 fig. 6 (Gulf of Mannar, Sri Lanka). sistency. Several hypotypes of Cavochalina bilamel- Chalina pulmata. — Dendy and Frederick, 1924: 499 lata: Carter belong to Cribrochalina punctata. (Abrolhos Islands, WA; nee Spongia patmata Lamarck, nee Chalina palmala: Ridley and Dendy). Callyspongia diffusa (Ridley) Material examined. Station BSS 187, one specimen Plate 11 figures 8-10, plate 31 figures 4, 5, (F52026). Syntypes of Cladochalina diffusa (BMNH). text-figure 70 Type specimens of Chalinopora lamella, Pachvchalina manus (AM, BMNH). Provisional synonymy. Diagnosis. Ctadochalina diffusa Ridley, 1884a: 672 (caption only), Variably branched, with branches sparse pi. 41 fig. D (Singapore). -Ridley, 1884c: 183 (descrip- or prolific, stubby or long and tapering, frequently tion, discussion). angular, knotty, moniliform, or compressed, erect Callyspongia diffusa. -Burton, 1934a: 541, text-fig. 6 or repent, anastomosing; or massive, lobate, tubu- (synonymy, discussion; Great Barrier Reef). -de Lauben- lar, flabellate. Firmly spongy, occasionally softer, fels 1950a: 12, text-fig. 6 (Hawaii). -Guiler, 1950: 6 (Tas- slightly viscid. Gold to mauve, or dark brown in mania). -de Laubenfels, 1954a: 86, text-fig. 51, pi. 4b life. Numerous oscules often raised on knots and (Caroline Islands, Mariana Islands). - Burton, 1959: 224 lobes, apical and marginal (Gulf of Aden, Zanzibar area).-Bergquist and Warne, on lamellae. Surface generally smooth, but 1980: 27, pi. 13 a-d (description, discussion: New hispid on elevations. Zealand, intertidal-2 m). Ectosomal reticulum variable in density, subdivi- Acervochalina finitima.- Ridley, 1884a: 399 (nee sion and depth, with erect spicular brushes. Main Schmidt, part: specimen from Port Jackson, NSW, only, skeleton irregular. Spicular core in all fibres thick, fide Burton, 1934a: 541). dense, many fibres packed with oxea. Chalinopora lamella Lendenfeld, 1887a: 757 (Port Jackson, NSW). -Lendenfeld, 1888: 85.-WhiteIegge, Description. Few, sparsely subdivided, straggling 1902b: 278, 285 (2 syntypes redeseribed). branches, generally knotty and angular, more so Ctadochalina lamella.- Burton, 1927a: 294 (transfer proximally, where almost serrated; tips tapering, only). almost smooth, terete to slightly compressed. Ser- Pachychalina manus Lendenfeld, 1887a: 777 (Port rated appearance through numerous oscules (width Jackson). -Lendenfeld, 1888: 88. - Whitelegge, 1889: 182 0.5-1 mm) on top (Shark Reef, Port Jackson; Maroubra Bay, NSW).- of tubercles and cones, mostly aligned longitudinally, Whitelegge, 1902b: 278, 284 (3 syntypes re-examined, 2 hence polygonal cross- removed). section of branches proximally. Total height 35 cm, Cladochalina manus. — Burton, thickness 1927a: 508, 510 of branches 1 cm proximally, 3-5 mm dis- (revision). tally. Dark brown in life, now faded (7.5 YR Ceraocha/ina multiformis var. digitata Lendenfeld, 5-6/4-6). Firmly spongy, slightly viscid. Surface, exept elevations mentioned, smooth, also microscopically. Very fine ectosomal reticulum apparent only with intermediate magnification. Primary meshwork wider in depressions, with indistinct subjacent vestibules forming speckled pattern. Over elevations, primary meshwork dis- Figure 70. Callyspongia diffusa, oxea, x943. tinctly denser, darker, mieropapillate. In ectosome, SPONGES FROM BASS STRAIT 109

only tangential primaries and secondaries can be onymy chiefly following Burton's manuscript indi- distinguished. Primaries 13-17 fim thick, cored by cations on the labels with the syntypes in London 4-6 spicules abreast, enclosing meshes around 175 (BMNH). Siphonochalina crassifibra Dendy /zm wide. Secondaries 8-9 /tm thick, cored by 1-2 (1889d) was merged in Callyspongia diffusa by spicules abreast, with meshsizes around 60 ;im. Burton (1934a), but this may be a synonym of Spicule brushes echinate surface at nodes and inbet- Callyspongia bullata (Lamarck). The synonymy of ween. Nodes frequently capped by bulbous spon- Ceraochalina ceylonica Dendy (1905) is doubtful gin, occasionally coalescent between neighbouring because of its thick oscular tubes and strongly nodes, producing micropapillate appearance noted spined surface. above. Meshwork of main skeleton irregular, with The polymorphism of the species, as discussed primaries often straggling and secondaries com- by Burton (1934a: 543) and Bergquist and Warne monly oblique. Primaries 33 to 63 ^m thick, spaced (1980: 28), is here accepted, with the reservations 208 to 440 )jm; secondaries 18-44 /tm thick, spaced and additions given above. The erect branching

1 10 to 218 fim. Half of diameter in primaries cored habit of the new specimen is closest to Pachycha- by more or less aligned oxea, some portions almost lina manus (here figured for comparison, pi. 1 1 fig. filled. Spicular core in secondaries proportionately 10), in which the branches are more prolific, with thinner, occasionally reduced to one spicule. short, clavate ends. The variability, though to a Spiculation: Hastate to fuisform oxea, straight lesser degree, also affects skeletal measurements to gently curved or bent, 50.6-59.^-67.1 x (fibres, meshes, and spicules). 1.5-2.7-3.3 fim. Callyspongia pergamentacea (Ridley)

Remarks. The synonymy suggested above is largely Plate 12 figures 1-3, plate 31 figure 6, that of Burton (1934a), with the exception of some names of Lendenfeld (1887a): of those included by plate 32 figure 1, text-figures 71-73 Burton, Cladochalina elegans is a good species of Provisional synonymy. Callyspongia, while Chalinissa ramosa (also cited Cladochalina armigera var. pergamentacea Ridley, Euplacella by Burton, 1934a: 598, as a synonym of 1881: 112, pi. 10 fig. 4 (Hotspur Bank, Brazil). communis) is indeed a synonym of Callyspongia Cladochalina armigera (nee Duchassaing and serpentina (see below). Chalinopora lamella and Michclotti). -Ridley, 1884a: 394 (Torres Strait; Port Pachychalina /nanus are here added to the syn- Denison, Qld).

X20.6. Figure 71. View of surface. Figure Figures 71, 72. Callyspongia pergamentacea. Details in reflected light. 72. Perpendicular section (surface above). 110 F. WIEDENMAYER

Cladochalina pergamentacea. -Ridley, 1884a: 398 partly light red-purple in life, now yellowish beige (Thursday Island, Torres Strait). (10 YR 8/4) throughout. Spongy, moderately firm Chalina pergamentacea. — Rid\ey and Dendy, 1887: 27, in places. Few, irregularly scattered oscules, 0.5-2.5 pi. 5 fig. 2 (Bass Strait). -Dendy, 1895: 244 (Port Phil- mm wide, some slightly elevated. Surface smooth, lip Heads). parchment-like, irregularly speckled by subdermal

Ceraochulinu pergamentacea. — Dendy , 1924: 327 (New vestibules. Fine ectosomal reticulum not always dis- Zealand). tinct with low magnification, partly masked by fine Callyspongia pergamentacea. — Hechtel, 1976: 240, 249, sediment. 253 (Bahian region, Brazil). Chalinopora paucispina Lendenfeld, 1887a: 765 In sections, primary meshes of ectosome trian- (Queenscliff, Vic.). -Burton, 1934a: 603, 609 (as syno- gular to polygonal, 400-800 /an wide, with fibres nym of Callyspongia ramosa). around 32 /an thick, cored to filled by 5-10 spic- Cladochalina paucispina.—-Burton, 1927a: 294 (transfer ules abreast. Primary meshes profusely subdivided only). by thinner fibres, incompletely separated into Chalinopora typica Lendenfeld, 1887a: 765 (with var. secondaries, with 2-3 coring spicules, about 20 /an tenuispina and var. megarhaphis, Port Jackson, NSW, thick, and tertiaries with only one coring spicule, and (unpublished) Port Phillip, Vic, and Kangaroo 10 /(in and thinner. Smaller meshes 30-80 /im wide. Island, SA). Main skeleton lax, irregular, with primaries and Cludochalina typica.— Burton, 1927a (transfer only). Ceraochalina papilla/a Lendenfeld, 1887a: 779f. (part, secondaries commonly straggling, their spongin unwarranted collective replacement name for annigera pale. In periphery (to 1-1.5 mm below surface), sensu Schmidt and Ridley, var. pergamentacea Ridley; where ascending and transverse fibres are more dis- not varr. nov. macropora, intermedia, micropore). tinct, primaries are 57-100 /an thick, with spacings 280-950 /an, frequently branch and form fascicles. Material examined. Station KG 1, one specimen (F52027). Type specimens of Chalinopora paucispina and Secondaries 35-45 /an thick, spaced 95-480 /an, Chalinopora typica (AM, BMNH). Hypotypes of occasionally branching and with free ends. In Cladochalina armigera, Cladochalina pergamentacea: deeper portions, arrangement more irregular, with Ridley, 1884a (BMNH); of Chalina pergamentacea: fibres frequently converging in knots. All fibres Ridley and Dendy (BMNH), Dendy, 1895 (NMV). cored by oxea to half to two-thirds of diameter, Diagnosis. Irregularly ramose, branches knotty, some primaries filled. Arrangement of spicules lobate, or compressed, occasionally flabellate. often disheveled to seimiplumose. Choanosome Spongy, light and very compressible when dry. Yel- scanty, with numerous small oxea, distinctly fainter lowish beige to light red-purple in life. Oscules con- than those in fibres. spicuous, irregularly scattered. Surface smooth, Spiculation: Fusiform oxea of the fibres, gently 6- parchment-like. Ectosomal reticulum thin, very curved to bent, 67. 71. 9-80.2 x 1.3-/. 9-2. 5 /an.

dense. Main skeleton lax, irregular; fibres gener- Interstitial oxea, 38.8-55. 1-61 /an long, below 1 ally thin, with pale spongin, with spicular cores /an thick. regular to semiplumose. Choanosome scanty, with Remarks. There is some similarity with Callyspon- smaller interstitial oxea. gia asparagus in spicular dimensions and in the con- Description. Erect-ramose with branches in a plane trast between the lax main skeleton, with scanty in irregular candelabrum-like fashion. Branches flesh, and the dense ectosomal meshwork. The irregular, knotty, with bulbous excrescences, fibres in C. asparagus are more regularly arranged, lacunose in one place, 4 to 15 mm thick. Specimen straighter, generally thicker, darker in colour, and 10.5 cm high, 7 cm wide. Partly yellowish beige, the spicules' density is much lower.

Callyspongia persculpta sp. nov.

Plate 12 figure 6, plate 33 figures 1-3,

text-figure 74

Holotype (?): F52035, station BSS 187 (38°32.0'S, 142°28.6'E, 17 km S of Warrnambool; depth 52 m, coarse sand and shells). It consists of two fragments believed to have been torn off a common base by the dredge.

Diagnosis. Ramose. Branches strongly sculptured,

with spoon-or ear-shaped lobes concrescent in tiers.

Figure 73. Callyspongia pergamentacea, spicules, x777. Rubbery, slippery, viscid; reddish brown in life. a. Oxea in fibres, b. Interstitial oxea. Surface microscopically smooth to granular. Small SPONGES FROM BASS STRAIT III

peripheral spicule brushes, or with enclosed brushes ending flush, occasionally echinating. Two condensed zones curving in abruptly from inner surface below rim, parallel with outer surface, being overgrown earlier surfaces, both thicker (290 ^m) Figure 74. Callyspongia persculpta sp. nov., oxea, x 877. than present ectosome. Main skeleton between these three zones more regular than that below. In oscules in convex portions of lobes, concavities deeper main skeleton, primaries have diameters of inhalant. Ectosomal reticulum deep, regularly con- 17-44 jtm, and are cored to variable extent, mostly densed, umbellate and cancellate. Erect spicular between one-to two-thirds of diameter (2-10 spic- penicils at surface commonly enveloped by fleshy ules abreast); spacing 100-455 /mi. dermis. Main skeleton generally regular, with vari- Secondaries 14-32 nm thick, spaced 105-520 /mi, able spicular cores in all fibres. Choanosome fairly frequently cored by 1-2 spicules. Many spicules dense, with numerous interstitial oxea not differ- scattered interstitially, also between three con- ent in size from those in fibres. densed zones. Spiculation: oxea, hastate, rarely fusiform, Description. Two fragments, of slightly unequal straight or curved, 60-66.7-72.6 x 2-5.2-4.4 /^m. size, conspicuously ornamented, roughly club- shaped, subdivided into tiers of concrescent hollow Remarks. The only fractures visible are on the lobes with sharp, upturned rims. Composite excres- inside of the first tier of each branch. They are simi- cences about 1 .5 cm apart, tilted outward and side- lar but do not match. If the branches are held with ways from axis of each branch. Each branch opposite facing fractures and with the rims of all slightly depressed overall, scooping shallower, more tiers approximately horizontal, inferring position irregular along proximal broadside. Some scooped in life, it is likely that the support was elevated, tiers on distal side like human ears. Lengths of perhaps a stalk or large shell, and that the branches branches 8 and 10 cm, width (along tiers), 3-5.5 were drooping at a 45° angle, proximally opposite, cm, depths (across tiers), 1.5-3 cm. Individual distally slightly converging. hollow lobes 8-15 mm wide, 5 mm deep, with walls This species is somewhat atypical for Callyspon- 2-3 mm thick distally. Reddish brow n in life, partly gia in its ectosomal structure. The condensation is preserved in alcohol (10 R 5-6/4), partly faded to very pronounced, with prolific branching of greyish fawn (7.5 YR 6-7/4). Rubbery, slippery, primaries just below the surface, and spicular siscid. Convex, often slightly rugose and verrucose brushes throughout the surface more crowded than outer and lower sides of scooped tiers exhalant, usual in species formerly assigned to Ceraochalina with regularly scattered small oscules, 0.5-1.5 mm and Euplacella. The similarity of this type of ecto- wide, 3-6 mm apart, with rims mostly flush, occa- some with that of Cribrochalina is more sionally slightly raised. Surface finely granulate at pronounced in this species than in Callyspongia intermediate magnification. Inhalant concavities bilamellaia, where the crowding of erect penicils In speckled to unaided eye by numerous vestibules, occurs only in some areas of the surface. (secondaries) with fine ostia in meandering, anastomosing lines. Cribrochalina, however, anastomoses in the main occasionally in vaguely stellate fashion. are much less in evidence, particularly Moderately dense choanosome with rounded- skeleton. Another atypical trait of this species is enveloping lacunose canal system. Ascending fibres distinct, the unusually thick collagenous dermis species is commonly straggling, frequently branching in most of the spicular brushes. The new with Spongia cras- mesial zone of lobe wall, closer to inner, concave comparable, in several points, redescribed by Topsent surface, curving to both sides to meet surfaces per- siloba Lamarck, 1814, as 4-5; Chalina crassitoba), pendicularly. Meshwork irregular, with connecting (1930a: 21, pi. 2 figs as Bass Strait (King Island). fibres and scarse intercalated primaries generally which also came from bifacial lobes (with oblique. This also has ear-shaped sides) of firm texture, has Ectosomal reticulation about 140 /an thick, con- inhalant and exhalent skeleton at the surface, with spicuously condensed, by prolific branching of a strongly condensed internally (overgrown earlier sur- primaries, and by closely spaced secondaries, from analogous zones oxea of similar dimensions to which additional short intercalated primaries may faces), and hastate Callyspongia persculpta. The lobes in issue. Mesh sizes 33-123 ^m. All radial spicule- those of type specimen are much larger than cores expand just below surface, like brushes, at Lamarck's only erect and mostly free, concres- times forming palisades. Coriaceous, finely fibrous in C. persculpta, in a massive base. Its oscules are wider dermis (brown in transmitted light) overlying cent below 112 F. WIEDENMAYER

than in the new species, more crowded and con- (part: BMNH 1884.10.10.1 (dry) only; Port Philli spicuous, especially on the wide margins of the Heads). -?Dendy, 1895: 243.

lobes. Its main skeleton is apparently more regu- Chalina polychotoma vat. oculata Carter, I885d: 28" lar, tighter than in C. persculpta, with rectangular (part: BMNH 1886.12.15.152,-153 only; Port Philli. meshes, their width generally exceeding their Heads). Chalina polychotoma var. trichotoma Carter, 1885; height. The chief difference is the great scarcity of 115. -Carter, 1885d: 284 (Port Phillip Heads). spicules in the main skeleton of Lamarck's sponge: Ductylochalinu uustralis Lendenfeld, 1886f: 726 (part they occur in primaries only, forming discontinu- material from Port Phillip Bay and Port Jackson only, ous single rows. The peripheral skeleton, by con- not from Western Australia). — Lendenfeld, 1887a: 813 trast, is rich in spicules, its but structure is not quite Cladochalina dendroides Lendenfeld, 1887a: 769, pt clear from Topsent's description. Primaries are said 19 fig. 21 (Port Chalmers, New Zealand). — Burton to have spicular cores rapidly increasing in diameter 1934a: 605 (syntype in BMNH redescribed). below the surface, becoming packed distally (there Cladochalina muinmillutu Lendenfeld, 1887a: 769 (Por is no mention of penicils, but primary tips are said Chalmers, New Zealand). - Burton. 1934a: 605 (holotype to frequently stand out in relief), and secondaries redescribed). forming the fairly wide meshes of the tangential Cladochalina tenuirhaphis Lendenfeld, 1887a: 769, pi 19, fig. 24 (Pott Chalmers, New Zealand). superficial reticulum (on inhalant surfaces only?) -Burton. 1934a: 606 (holotype redescribed). are also charged with spicules. The specimen is dry, I'uchychulina puuetspina Lendenfeld, 1887a: 77( and any coriaceous ectosome possibly originally (junior secondary homonym of Chalinopora paucispbn. present (as in C. persculpta) might have vanished Lendenfeld, 1887a; Nett Zealand). -Burton, 1934a: 60(- through natural maceration. (holotype redescribed). This insecurity makes the generic placement of I'uchychulina ramulosu var. lamella Lendenfeld, 1887a. crassiloba speculative. Van Soest (1980: 31) dis- 777 (as lamella, junior secondary homonym ol cussed it as a possible synonym of Atnphimedon Chalinopora lamella Lendenfeld, 1887a; New complanata (Duchassaing). 1 find this very unlikely. Zealand). — Burton, 1934a: 606 (holotype redescribed).

\ Pachychaima ramulosu ar . digitals I endenfeld, 1887a: Callyspongia ramosa (Gray) 777, pi. 19 fig. 22 (as digitata, junior secondary homonym of Chalinis.su communis digitata Lendenfeld, 1887a; Plate 11 figure 11, text-figure 75 Port Chalmers, New Zealand). -Burton, 1934a: 606 (holotype redescribed). Provisional synonymy. Pachychaima macrospina Lendenfeld, 1887a: 777 (Port Spongia ramosa Gray, 1843: 295 (pari; New Zealand). Chalmers, New Zealand). -Burton, 1934a: 606 (holotype Pachychaima ramosa. -Dendy. 1898: 318, pi. 33 (pari; redescribed), pi. 19 fig. 18 (junior secondary homonym redescription of 4 syntypes, including leclotype). of Cludochulina elegans Lendenfeld, 1887a; Port Callyspongia ramosa. — Burton, 1934a: 597, 603 (syn- Chalmers, New Zealand). - Burton, 1934a: 606 (holotype onymy, diagnosis, resumes, revision, discussion of syn- redescribed). onyms). -Burton, 1938: 20 (Tasmania). -Bergquist, Pachychaima rigida Lendenfeld, 1887a: 778. pi. 19 fig. 1961a: 36 (New Zealand). - Bergquist and Warne, 1980: 14 (junior secondary homonym of Chalmissu rigida Len- 30, pis 14f, 15a-f (synonymy, description, discussion, dis- denfeld, 1887a; New Zealand). - Burton, 1934a: 607 tribution, New Zealand). (holotype redescribed). Cludochulina nuda Ridley, 1884a: 395, pi. 41 fig. i Pachychulina oculata Lendenfeld, 1887a: 778 (junior (Torres Strait). -Burton, 1934a: 609 (comment on secondary homonym of Chalina polychotoma var. oculata holotype). Carter; Mauritius). - Burton, 1934a: 608 (holotype Cladochalina nuda var. abruptispicula Ridley, 1884a: redescribed). 396, pi. 41 fig. (Torres Strait). -Burton, j 1934a: 609 Ceraochalina papilluta var. macropora Lendenfeld, (comment on holotype). 1887a: 780 (as macropora, junior secondary homonym Ceraochalina nuda. — Lendenfeld, 1887a: 781 (with of Chaline/la macropora Lendenfeld, 1887a; New varr. oxyus and oxystrongylus, nomina vana). Zealand). -Burton, 1934a: 607 (lectotype in BMNH Chalina nuda. -Hentschel, 1912: 397 (Aru Islands, redescribed). Indonesia). Ceraochalina papilluta var. intermedia Lendenfeld, Chalina polychotoma (nee Esper): Carter, 1885d: 284 1887a: 781 (as intermedia, junior secondary homonym of Siphonochulinu intermedia Ridley and Dendy, 1886; Port Chalmers, New Zealand). -Burton, 1934a: 607 (holotype redescribed). Ceraochalina papilluta var. microflora Lendenfeld, 1887a: 781, pi. 19 fig. 16; Port Chalmers, New Zealand). -Burton, 1934a: 607 (holotype redescribed). Figure 75. Callyspongia ramosa, oxea, X734. Ceraochalina levis Lendenfeld, 1887a: 782, pi. 19 fig. --

SPONGES FROM BASS STRAIT 113

19 (Torres Strait; Port Chalmers, New Zealand). fig. 3 (Port Jackson, NSW, Port Phillip Bay, Vic.; Torres Whitelegge, 1906: 406 (2 type slides in AM re- Strait, Qld; Port Chalmers, New Zealand). -Burton, examined).— Burton, 1934a: 604 (lectotype and one 1934a: 608 (syntypes from Port Chalmers, Port Phillip paraleetotype redeseribed). and Torres Strait redeseribed).

Ceraochalina microrhaphis Lendenfeld, 1887a: 783 fuchultna typtea Lendenfeld, 1887a: 817, pi. 18 fig. (Port Chalmers, New Zealand). -Burton, 1934a: 607 5, pi. 27 fig. 24 (junior secondary homonym of (holoiype redescribcd), Chalinopora typicu Lendenfeld, 1887a; Port Jackson, Ceraochalina multiformis var. maeandra l.endenleld, NSW; Western Australia; New Zealand). 1887a: 783 (Port Chalmers. New Zealand). - Burton, I uchalina puueispinu Lendenfeld, 1887a: 818, (junior 1934a: 607 (holotype redeseribed). secondary homonym of Chalinopora puueispinu Lenden-

Ceraochalina multiformis var. lamella I endenfeld, feld, 1887a; Port Jackson, NSW; Port Phillip Bay, Vic; 1887a: 783, pi. 19 fig. 15, pi. 27 fig. 25 (as lamella, junior Port Chalmers, New Zealand). - Burton, 1934a: 608 (lec- secondary homonym of Chalinopora lamella l.endenleld, totype and paralectolypes described). 1887a; New Zealand). -Burton, 1934a: 607 (holotype I uchalina macropora Lendenfeld, 1887a: 818 (junior redeseribed). secondary homonym of Chalinissa macropora Lenden-

Ceraochalina multiformis var. etegans I endenfeld, feld. 1887a; Port Jackson, NSW). 1887a: 784, pi. 19 fig. 23; pi. 27 fig. 4 (as etegans, junior Chalina macropora. — Whilelegge, 1906: 454 (type slide secondary homonym of Claclochalina eleguns l.endenleld, in AM re-examined, new record off Crookhavcn River, 1887a; Westernpori Bay, Vic). NSW). Euchalina phtilipplnensis Lendenfeld, 1887a: 819 Ceraochalina multiformis var. digitata I endenfeld, (spelled plulippensis on 757; Port Phillip Bay, Vie.; 1887a: 784, pi. 19 lie. 20 (part: not syntype from Port p. Jackson; as digitata, junior secondary homonym of New Zealand). Dactyloehalina monilatu (nee Ridley, 1884a).— Chalinissa communis digitata I endenfeld. 1887a; Port Lendenfeld's material only, Chalmers and Chatham Island, New Zealand). -Burton, Lendenfeld, 1888: 100 (part: identical with syntypes of Dactyloehalina australis; Port 1934a: 607 (syntypes from New Zealand redeseribed). lackson), - Whitelegge, 1889: 183 (off Green Point, Port Ceraochalina multiformis var. mollis I endenfeld, Jackson). 1887a: 784 las mollis, iunioi secondary homonym of Phyllospongia (Antheroplax) perforata (nee Hyatt, Cladochalina mollis I. endenfeld, 1887a; Port Chalmers, 1877). Lendenfeld, 1889b: 172 (part: dry specimen from New Zealand). Burton, 1934a: 60S (holotype New Zealand. BMNH 1886.8.27.46, only, = slide AM redeseribed). (.3711).

Ceraochalina multiformis var. dura I endenfeld. 1887a: 1 Paehychalina densa Br^ndsted, 1923: 125, text-fig. 784, pi. 19 fig. 25 (Port Chalmers, New Zealand). 7 (Auckland Islands, New Zealand). Burton, 1934a: 608 (syntype in BMNH redeseribed). Getfiodesflagelbformis Br^ndsted, 1923: 129, text fig. Ceraochalina retepiax Lendenfeld, 1887a: 785, pi. 19 10 (Auckland Islands, New Zealand). fig. 17 (Port Chalmers, New Zealand). - Burton. 1934a; Gelliodes //7//o//m.v Br^ndsted, 1923: 131, text figs. 12, 608 (holotype redeseribed). 13 (Campbell Islands, New Zealand).

Ceraochalina tenella I endenfeld, 1887a: 785 (junior Toxochallna dtfficilis Br^ndsted, 1923: 132, text-fig. secondary homomm ol (halmclla tenella I endenfeld, 14 (Auckland Islands, New Zealand). 1887a; Pott Phillip Bay, Vie.). IGelliodes biformis Brr/mdsted, 192.3: 449, text-fig. 8 l Ceraochalina euplax l.endenleld, 1887a: 785, pi. \ ) lit'. (New Zealand). 13 (junior secondary homonym of Cladochalina euplax Paehychalina uffinis Br^ndsled, 1924:455, text-tig. 13 - 608 Lendenfeld, 1887a; New Zealand). Burton, 1934a: (New Zealand). (holotype redeseribed). IPachvchahna litnue Br<;>ndsled, 1924: 455 lexl-fig. 13 Ceraochalina extrema Lendenfeld, 1887a: 785 (Port (New Zealand). Chalmers, New Zealand). - Burton, 1934a: 608 (holotype specimen redeseribed). Material examined. Station BSS 180, one ramosu Gray (BMNH Dactyloehalina mollis Lendenfeld, 1887a: 812 (junior (1-52028). Syntypes of Spongia polyehotoma van. secondary homonym of Cladochalina mollis Lendenfeld, only). Type specimens of Chalina Carter (BMNH), of Dactyloehalina 1887a; east coast). - Lendenfeld. 1888: KM). ociilala. trichotoma mamillala. C. Dactyloehalina candelabrum Lendenfeld, 1887a: 812 australis. Cladochalina dendroides, C. puueispinu, P. ramulosass.tr. (Port Chalmers, New Zealand). - Burton, 1934a: 608 tenuirhaphis, Paehychalina macrospina, P. etegans, P. rigida, (holotype redeseribed). lamella, digitata. P. pupillatu van. macropora, intermedia, Dactyloehalina conulata Lendenfeld, 1887a: 813, pi. < eiaochalina mieroporu, C. levis, C. microrhaphis, C. multiformis van. 18 fig. 2 (junior secondary homonym of Chalinopora digitata, mollis, dura, <". conulata Lendenfeld, 1887a; Port Jackson, NSW). maeandra, lamella, etegans, Dactyloeha- . tenella, C. euplax, C. extrema, Euchalinopsis dendroides Lendenfeld, 1887a: 816, pi. retepiax. t D. candelabrum, D. conulata, Euchalinop- 18 fig. 8 (junior secondary homonym of Cladochalina lina mollis, minima. Euchalina typicu, E. dendroides Lendenfeld, 1887a; lllawarra and Port Jack- sis dendroides. E. macropora, E. phillippinensis Lendenfeld son, NSW). /laucispina. E. and BMNH). Hypotypcs of Chalina polyehotoma: Euchalinopsis minima Lendenfeld, 1887a: 816, pi. 18 (AM I ,

II WlliDr.NMAYI'k

bent or flexuou.s, ( !arter, ihksu. (HMNl i), and of Phyllospongla perforata: ends, straight to irregularly

Lendenfeld, 1889b (liMNll). 67-77-86.5 X 1 .4-/. 7-2.6 ftttl.

Remarks. The synonymy suggested above is largely Diagnosis. Ramose, erect; branches terete or com- Burton (1934a: 603), with some exceptions pressed, solid or tubular; occasionally encrusting, thai of records outside New Zealand waters. with oscular processes. Firmly spongy, occasion- concerning Lendenfeld (1887a), with its varie- ally soil or almost incompressible. Mauve or dull Euchattna exigua ties, is here accepted as a good species of Cullyspon- yellow in life. Surface generally smooth, even, Chattnopora paueispina lendenfeld is here locally micioconulosc. Oscules numerous, scattered gia, regarded as a junior synonym of Catlyspongia per- or in lines. Flush Or raised. EctOSOmal skeleton vari- garrtentacea (see above). Chalitta uculata ably condensed 01 paralangential, subdivision not (1924) is, in my opinion, always distinct, may be locally absent. Erecl spie- novaezealandiae Dendy a synonym of Caliyspongia serpentina (sec below). nlai brushes may be present at surface. Main skele- Topsenl's revision of Spongia serpentina ton mostly regular, rectangular, primaries and most (1933)

I aiuarck apparently came lo Burton's attention too secondaries regularly cored by o\ca, but fibre thick- late, he (1934a: 598), like Topsenl, accepted ness and proportion of spongin varies. Oxea vari- and

I enden (eld's epithet communis as valid. Both able, especially in thickness, may also occur and C. serpentina seem to be interstilially in ehoanosome of variable density. CaltySpongla ramosa fairly in Strait, in Toxa may be present in variable proportions. common Bass and, overlapping several trails, were easily confused in the past. This Description. Single branch, regularly compressed is apparent in Chalina polyehotoma: Carter, with expanding lo spatU late end asymmetrically cleaved its varieties (all merged by Dendy, 1895: 243), which

by two notches, one 6 oilier I mm deep, mm. comprises specimens of both species; but also in

Branchlel, I long, issuing cm from base. Main the revisions of (he type series (different parts branch proximally Icicle, 4 thick, soon mm com- thereof in Dendy, 1898, and Burton, 1934a) of

pressed, expanding from 3-4 I mm to 5 mm at end. Spongia ramosa Gray, in which (wo of the seven total height 7 cm. synlypes probably belong to C. serpentina. Pale ochre in life (It) YR 6/6), now slightly The oxea in both species are very similar in range faded. Softly spongy, limp. Oscules numerous, of shapes and dimensions. There is also, occasion-

0.3 l mm wide, round lo irregularly scat elongate, ally, in both species, a more or less pronounced leicd along both narrow sides and one of faces. segregation of thick and thin oxea, thick ones in Surface smooth optically, microeonulose al inter- C. ramosa being hastate with abrupt points, and mediate magnification. Vestibules only apparent generally more common. The new specimen of C. hour Outside in upper half of main branch. ramosa is atypical in having only thin fusiform Iclosomal reticulum poorly defined even at inter- oxea. Of the specimens redescribed as synonyms mediate magnification, masked by dermis and ill- at' C. ramosa by Burton (1934a), one, the syntype sorted sediment. o\' Ceraoelialina levis Lendenfeld from Torres Meshwork of main skeleton regular, though Strait, agrees with the present specimen in (his primaries and secondaries generally sinuous. respect, except for possessing rare oxea of the Peripheral skeleton moderately condensed, by normal, sioul kind. Bergquist and Warnc (1980: crowding of secondaries and by adventitious 31) noted an inverse relationship between volume primaries beginning from fourth lo second secon- of spongin (fibre thickness) and spicule size in their dary fibre below sin face, bill also from outermost numerous specimens. The present specimen, with ones. All primaries have short, conical free ends relatively thin fibres, does no! seem to fit this slate. of unequal length, rudimentary to about lOO/un. Burlon's (1934a) synonymy was accepted by lips of conules frequently rugged, truncate, with Bergquist and Warne (1980), who noled great mor- spicules occasionally protruding. Primaries below phological variability in their material, externally periphery 26-45 /an I hick, spaced 240-380 /tin. and in skeletal features. I hey noted toxa in only Secondaries of deeper portions 23-36 /

re-examined), the chief diagnositic trait seems to 772. pi. 20 fig. M; pi. 27 figs 5. 11, 13, 15 (Port .lack be the distinctly higher condensation in the son, NSW). -Lendenfeld, 1888: 8. - Whitelegge. 1889: ectosomal skeleton of C. serpentina. In boili spe- 182 (off Green Point. Port Jackson). Chalinissa communis var. digitata Lendenfeld, 1887a: cies, however, the condition of the surface is pi. 20 intermediate between that of Callyspongia 772, fig, 30 (Port Jackson, NSW). Pachychalina communis.— Whitelegge, 1901: 70 (syn- (nominotypieal subgenus sensu van Soest, 1980) onymy; Tuggerah Beach. NSW '). -Whitelegge. 1906: 458 and subgenus C. (Euplacella sensu van Soest. (van. flabellum and digitata, Chalinissa elegans, C. elon- 1980, 1982, correctly Ceraochalina): the conules of gata, C. tenuifibris, c. serpens Lendenfeld merged, all spongin cored by oxea are obviously homologous types re-examined, spicule dimensions; coast a\ NSW, with naked spicule-brushes, and the latter seem to common). occur occasionally in both sibling species. The Cladochalina communis.— Burton, 1927a: 504, 505 specific identities of Ceraochalina pergamentacea (transfer, the 2 varieties merged). - topsent, 1933: 2, pi. (nee Ridley): Dendy, 1924, and of Cladochalina 1 figs 5, 6 (2 syntypes of typical var. and var. of Spon gia serpentina redescribed). dendyi Burton, 1929a (both cited in Burton's syn- Euplacella communis.— Burton, 1934a: 598 (transfer, onymy of 1934a) are impossible to ascertain on the synonymy: Great Barrier Reef). basis of their descriptions. Chalinissa macropora I endenfeld, 1887a: 772, pi. 20 Callyspongia serpentina (Lamarck) fig. 28 (Port Jackson, NSW ). Cladochalina macropora.— Burton, 1927a: 504, 505

Plate 12 figure 4, plate 32 figures 2-4, (transfer )

Chalinissa tenuifibris l endenfeld, 1887a: 773, pi. 20 text-figure 76 fig. 29 (Porl Jackson. NSW ) Provisional synonymy. Cladochalina tenuifibris.- Burton, 1927a: 504, 505 Spongia serpentina Lamarck. 1814 (1813 1814): 452 (transfer, possibly synonym of C. communis).

(with var. tf; King Island, BaSS Strait, lasmania. I lnilinis.su serpens I endenfeld, 1887a: 773, pi. 20 fig. Redescribed by Topsent, 1933: 2). 33 (east coasl oi Australia). -Burton, 1927a: 504, 505 Chalina polychotoma (Esper) var. oculata Carter, (synonymised with Cladochalina communis).

I885d; 284 (part: BMNH 1886.12.15.154,-156 only; Port Chalinissa elegans l endenfeld. 1887a; 773, pi. 20 fig, Phillip Heads, Vic). 27 (junior secondary homonym of Cladochalina elegans Chalina polychotoma vat. moniliformis Carter, 1885d: lendenfeld. 1887a; Broughlon Island, NSW).- 285 (south coasl of Australia). Lendenfeld, 1888: 87 (lllawarra, NSW'). - Whitelegge, Chalina polychotoma var. angulata Carter, I885d: 285 1902b: 278, 286 (2 syntypes re-examined).— Burton, (Port Phillip Heads. Vic). 1927a: 504. 505 (synonymised with Cladochalina Pachychalina elongata Ridley and Dendy. 1886: 329.— communis). Chalinissa rigida Lendenfeld, 1887a: 773 (Port Jack Ridley and Dendy, 1887: 23. pi. 6 fig. 1 (off Moncocur son, NSW). -Burton, 1927a: 504, 505 (synonymised with Island, Bass Snail, 1 as). communis). Cladochalina elongata.— Burton, 1927a: 508, 510 ( ladochalina (transfer). < 'halinissa elongata I endenfeld, 1887a: 774, pi. 20 fig. - 87.- Euplacella elongata.— Button, 1934a: 597 (transfer 34 (lllawarra. NSW). Lendenfeld, 1888: slide ex only). Whitelegge, 1902b: 278, 286 (AM-type BMNH- Chalinissa communis Lendenfeld, 1887a: 772 (collec- syntype re-examined). —Burton, 1927a: 504, 505 (syn- with Cladochalina communis). tive name for following varieties, no nominotypieal onymised subspecies; junior secondary homonym of Tubulodtgi- Chalinissa ramosa I endelfeld, 1887a: 774. pi, 20 fig, (Port Jackson). 1927a: 504, 505, 510 (syn- tus communis Carter, 1881a). -1 cndenfeld, 1888: 87, pi. 31 -Burton, onymised with Cladochalina communis). 8 (Port Jackson). -Whitelegge, 1902b: 278. 286(syntype Pachychalina ramosa.— Whitelegge, 1906: 455 (off in AM and type slide of var. flabellum e\ BMNH Coogee, Shoal- re-examined). Wata Mooli, off off Wollongong, off haven Bight, NSW). Chalinissa communis var. flabellum I .endenfeld, 1 887a:

Ceraochalina typica 1 endenfeld, 1887a: 782, pi. 19 fig. 26 (Port Phillip Bay, Vie.). Chalina oculata var. novaezealandiae Dendy, 1924: 326 (New Zealand). '.Cladochalinu dcndvi Burton, 1929a: 421 (Victoria Land, New Zealand's Antarctic Territory).

Material examined. Station KG 9, one specimen (L52029). Type specimens of Chalina polychotoma van. oculata, moniliformis, angulata Carter, I885d (BMNLI), of IWhv chalina elongata Ridley and Dendy (BMNH), of Chalinissa communis vast, flabellum, digitata, C. mac 76. Callyspongia serpentina, spicules, x 1059. a. Figure ( ropora, C. tenuifibris. C. serpens. '. elegans, C. rigida. Oxea in fibres, b. Interstitial oxea. 116 P. WILDENMAYtR

C. elongata, C. ramosu, C. lypicu Lendcnfeld (AM and Ectosomal skeleton much condensed, meshsizes BMNH). Hypotypes of Pachychuliiui rumosa: around 60 /tm. Cubic arrangement frequently Whitelegge, 1906 (AM). obscured by oblique and branching fibres. Thin- Diagnosis. Ramose, erect. Branches irregular, ner fibres without order. Fibrous conules at sur- knotty, angular, lobate, or compressed, occasion- face short, generally below 30 /*m, around 60 pm ally frondose, often prolific. Firmly spongy, rub- apart, cored by 1-3 broken spicules. Most echinat- bery, viscid; fawn in life. Oscules numerous, ing conules between nodes of reticulum. Tips of scattered. Surface smooth, glabrous to microconu- primaries mostly blunt, bifid or T-shaped. Ecto- lose. Ectosomal reticulum much condensed, chie- some contains variously abundant foreign detritus fly irregular, with short echinating fibres. (sand grains, flocculent sediment, spicules and their Choanosome cavernous, main skeleton very irregu- fragments). Foreign matter scanty in choanosome. lar. Indistinct primary fibres cored, but oxea often Spiculation: Fusiform oxea of fibres, rarely semiplumose. Many oxea interstitial, smaller. straight, commonly asymmetrically bent, often Description. Prolifically branched, with 15 broken, 46.5-54. 7-59.8 x 1.6-2-2.6 Mm. Intersti- branches distally. Basal portion a palmate frond, tial oxea, 48.7-55.3 x 0.8-1.2 pm. 8 mm thick, 6.5 cm wide distally, 10 cm high, with cleft 6 cm long on one side. Few crooked, stubby Remarks. The two syntypes of Spongia serpentina branchlets from its sides and lower face. Remain- Lamarck (typical variety), as redescribed by Top- ing longer branches from upper face and edge of sent (1933: 2), are beach-worn specimens in which

basal frond, straggling, knotty, compressed in the ectosome is mostly gone. The type of var. /3

different directions, of variable thickness, gener- (Topsent, 1933, pi. 1 fig. 6) is better preserved, but

ally with angular cross-section. Angularity height- Topsent unfortunately neglected to describe its ened by coarse pits and lacunae, often crowded ectosome. However, the shape of the two figured with common margins forming rugo-rcticulate pat- specimens, Topsent's identification and compari- tern. Most branches only partly affected by this son with Chalintssa communis as redescribed by pattern, to variable degrees. Some branches more Whitelegge, 1906, and the frequently semiplumose strongly compressed, flabellate, below bifurcation arrangement of oxea in primary fibres, leave no

or multiple subdivision, partly also due to concres- doubi that S. serpentina is the oldest available name cence of contiguous branches. Most branches of this species.

0.6-1.5 cm wide, erect ones 10-21 cm high. Total Through the type specimens listed above, I find height 31 cm. the poloymorphism confirmed, which was noted Fawn in life (7.5 YR 7/6), now duller (10 YR by Whitelegge (1906) and Burton (1927a), and 6-8/4). Firmly spongy, rubbery, viscid, residual which is apparent in the figures cited above in syn- mucus upon handling difficult to wash away. onymy. Detailed descriptions of skeletal structure Oscules numerous, scattered all over, 1-3 mm wide, are so far scanty and inadequate. Whitelegge (1906: round. Surface, except pitted areas, smooth, gla- 459) merely noted that the extreme variability also brous, also on branch tips. Many areas show dense, affects the fibrous skeleton and the spicules. He fine, weak speckling by vestibules. Very dense was not clear on the structure of the ectosome,

ectosomal reticulum with minute conules visible remarking that it is "velvet-like in appearance, and only at intermediate magnification. No clear dis- coated with a thin crust of oxeote spicules; the latter tinction of primary and secondary tangential fibres. are very closely arranged and partly conceal the Choanosome regularly cavernous, riddled by dermal network as well as the pores, which are canals 0.5-0.8 mm wide, fleshy interstices contain- scarcely visible to the unaided eye." It seems likely ing numerous weak oxea. Main skeleton quite that this refers to naked spicular brushes homolo- irregular. Little distinction between primary and gous to the spongin-conules in the new specimen. secondary fibres, but former contain more spicules, Topsent described the oxea in the fibres of the type commonly 2-6 abreast, are more flexuous, fre- material of Spongia serpentina as fairly stout, quently fasciculate, and prolifically branched. Spic- barely fusiform with short conical points, meas- ules in primaries aligned and coring, but frequently uring 70-85 x 4-6 /*m; interstitial oxea as more disheveled, semiplumose. Secondaries may be fusiform, always weaker, 70 X 1.5 pm. Spicular devoid of spicules or contain a discontinuous core measurements given by Whitelegge (1906) for the of one, rarely two spicules. No appreciable differ- types of Chulinissa communis and of the synonyms ence in thickness (about 25 fisa) between primaries recognised by him have a range of 60-80 X 4.5-6.5 and secondaries. Orientation of both, except most pm for oxea in the fibres, 60x2^ for intersti- of peripheral portion, random, mesh sizes variable tial oxea (in communis only). The principal oxea (50-600 pm). in the new specimen are smaller, but hardly signifi- . ,

SPONO] s FROM li\ss STRAIT II? cantl) so, considering the analogous case in the new Sui lace smooth to unaided eye. With low mag specimen of Callyspongia ramosa. nification, regularly conulose, lowet conules prox imal, higher ones distal. In three specimens with Callyspongia toxifera sp. nov. thin branches, ectosomal reticulum rudimentary, Plato 12 figure 5, plate 32 figures 5 6, undulating. In F52030, tangential reticulum at SUI lace, between very low conules, ineshwork dis- text-figure 77 tinctly finer than inside, its distal secondary Holotype: F52032, station BSS 181 (38 J9.8'S, branchlets like thinly branched specimens in SU! I44°18.2'E, 30 km SI of Lome; depth 79 m, verj lace. Mam skeleton an irregular ineshwork of clear, d with chaetopterid worm tubes), rhree paratypes: pale, weak spongin fibres, always free of propei F52030, station BSS 179 (39°03.2'S, 146 19.51 ,l ofWil spicules and foreign inclusions. Commonly no ills skni>. Promontory; depth ss m, muddv, sand); F52031 and miction of primaries and secondaries, all fibres l 52033, station BSS LSI sinuous to crooked, occasionally bent at almost Diagnosis. Small, sparsely branched. Branches right angles, 12 to 26 /mi thick, swelling and tapei thin, straggling. Soft, limp; greyish yellow in life, ing profusely, enclosing randomly oriented, twisted minute, lines. Oscules irrgularly scattered oi in Sur- meshes 120 ) !0 /un w ide. Some primaries vaguclv face smooth, microconulose, Ectosomal reticulum distinct by iadi.il orientation and more crooked or condensation rudimentary, Main skeleton course. In FS2031, meshwork densei than in othei irregular, with weak, pale spongin fibres withoul specimens, with more uniform meshsizes. oxea. Proper spicules onl) intei stitial toxa of vari- Peripheral condensation absent oi slightly able frequency. Scattered foreign detritus in developed, with mote icgulai meshes subpaiallel choanosome. with sin lace, and thinnei fibres (aiouud 8 /

and 160 ; length (chord) 25.6 65 2 89.4 ^m, thick dichotomous. Distance between bifurcations 1 to i less 0.8 1 .7 /(in. hie, 8 mm. I 52030 greyish yellow (2.5 Y7 8 4) in unchanged in alcohol. Othei specimens now lighter, Niphatidae van Soest, 1980 pale beige to cream. Soft, limp, easily lorn. Oscules inconspicuous anil minute, irregularly scattered, Gelllodes Ridley except 152030, where more conspicuous, I 1 .5 mm Gelllodes Ridley, 1884a: 426 Ridlej and Dendy, aligned narrow sides. wide, regularly on two 1887: -17 (new diagnosis). Dendy, 1905: 137 (defini mill), di' Laubenfels, 1936a: 53 (definition in Des macidonidae, revision), Van Soest, 1980: 106 (definition in Niphatidae), Desqueyroux-Faundez, 1984; 779f,

(definition in Niphatidae, description ol -i species).

Diagnosis. "Niphatidae in which the tangential ectosomal reticulation is [often] obscured by pro truding tufts of the choanosomal primaries, producing a rough conulose surface. Spongin fibres hugely Figure 77. Callyspongia toxifera sp. nov. Toxa from present. Secondary reduced.

F52033, 709. Microscleres abundant siginala." (Van Soesl , 19X0: 118 F. WIEDENMAYER

106.) Proportion of spicules to spongin highly vari- often epibiotic. Compressible, resilient; light yellow able. The surface, in some species, may be almost to whitish, or greyish purple in life. Surface velvety, smooth. The fibrous skeleton, or portions thereof, microconulose. Oscules round, 2-3 mm wide, flush may be isodictyal. or raised. Conspicuous meandering vestibules. Choanosome cavernous. Main skeleton irregular, Gelliodes incrustans Dendy with little differentiated primary spongin fibres cored by several oxea, often in semiplumose Plate 2 figure plate 12 figure 2, 7, arrangement; secondaries with single oxea.

plate 33 figures 4-6, plate 34 figure 1, Peripheral skeleton may be tighter, almost isodictyal. Oxea also interstitial, with numerous sigmata, text-figures 78-81 foreign detritus and microsymbionts.

Gelliodes incrustans Dendy, 1905: 137, pi. 9 fig. 6 (Sri Description. Sponge now dissected, originally Lanka). -Levi, 1965a: 20, fig. 25 (Red Sea).- ovoid, slightly compressed, small, attached to side Desqueyroux-Faundez, 1984: 782, figs 7, 53, 62, 63 (New Caledonia). of Jaspis stellifera (described above), 1.7 x 2 cm wide at base, 8 mm high. In growth position, with Material examined. Station 1 (one specimen, F52036). KG mouth of larger vermetid proximally encrusted by Diagnosis. Small, encrusting to cushion-shaped, Jaspis stellifera on top, narrower, almost conical

»u«*

^j^^^M'&m^i^, ^^zi^0k^ H ,^£^f .^^ 78

\* nXv - By >

78-80. Gelliodes incrustans. Details Figures in reflected light. Figure 78. Median sagittal section of whole specimen x 6.6. Figure 79. Surface (above) and periphery, x 20.5. Figure 80. View of surface showing the pattern of subdermal canals, x 20.5. SPONGES FROM BASS STRAIT 119

Figure 81. Celliodes incrustans. spicules, x979. a. Oxea. b. Sigmaia.

half of Gelliodes, still in place, directed upwards, Remarks. Previous records were from the Gulf of bears apical, sharp-rimmed oscule, 2 mm wide. Mannar (Dendy), the Red Sea (Levi), and New Colour in life not recorded, now, in alcohol, pale Caledonia (Desqueyroux-Faundez, with synoptic fawn to dark beige (5 YR 7-8/4). Compressible, table of skeletal measurements). The latter record resilient, velvety surface. Weakly undulating sur- is somewhat doutbful, as the sigmata are consider- face smooth to unaided eye, with meandering sub- ably larger (50-90 x 2-3 /tm). My measurements dermal canals and vestibules barely visible. With agree well with those of Dendy and Levi, except low magnification, surface regularly and finely for the somewhat thicker oxea in the latter (105-130 conulose to hispid, conules about 50 /

Choanosome cavernous, traversed in all direc- pi. 1 figs 1-7) for Gelliodes pumilus (Lendenfeld),

tions by confluent, meandering canals of rounded but that outer skeleton (to a depth of 200 /tm) is section, 350-750 /im wide. Main skeleton irregu- devoid of spongin. The inner, fibrous main skele- lar, often obscured by much scattered to cluster- ton is described as isodictyal. Most of Burton's ing detritus and macrosymbionts. Fine sediment specimens have a fairly smooth surface, but Len- dominates, especially at surface, over foraminifera denfeld's holotype (also figured) is coarsely conu- and their debris, whole branches of bryozoans, and lose. The description of this species in van Soest foreign spicules, chiefly triradiates, probably from (1982: 89, fig. 8) refers to an undifferentiated main neighbouring calcareous sponge Aulorrhiza skeleton with thick primaries and transverse thin- procumbens. Spongin dominates in fibres. Struc- ner secondaries, both packed with spicules, oxea ture of main skeleton variable, but with two types being larger than in G. incrustans. Of the massive- or zones: (1) Generally deeper inside with thicker lobose species of Gelliodes with Indo-West Pacific fibres. Primary fibres indistinct, little oriented, records, three others have fairly smooth surface. slightly thicker, irregularly spaced, occasionally G. carnosus Dendy (see Desqueyroux-Faundez, branching, with spicular cores fifth to third of 1984: 779) and G. fragilis Desqueyroux-Faundez diameters (2-5 spicules abreast, often broken), (1984: 782) both differ from the present species generally straggling; arrangement of spicules occa- chiefly by higher proportion of spicules to spon- sionally disheveled, semiplumose. Intervening gin, and consecutively more rigid or more fragile secondary mesh work, generally with single coring consistency. G. tenuirhabdus Pulitzer-Finali spicules, highly irregular in meshsizes and orien- (1982b: 1 12), known from a single specimen, seems similar the the tation. (2) Generally below surface to variable to have a skeleton very to one of depths (up to 1.5 mm), without apparent primaries, present specimen. But the proportion of spicules fibres mostly thinner, meshwork more regular, to spongin is even weaker in the former, and the strongyles, finer, almost isodictyal. Interstitial oxea in both oxea are thin (100 x 1.5 /mi), almost zones. weakly mineralised. It could be argued that these ecophenotypes, but Overall, fibre diameters 10-65 /tin, average about are differences of degree, or of Barrier Reef will have 35 /tm; meshsizes 68-285 /mi for inner zone, around more material from the Great decide tenuirhabdus 100 /im for outer zone. to be examined to whether G. G. incrustans. Spiculation: (1) Oxea, fusiform, straight to is a synonym of 1.6-2.9-4.4 /mi; slightly curved, 93-/75- 142 x (2) Oceanapiidae van Soest, 1980 sigmata of variable curvature, occasionally con- Synonym: Phloeodictyidae Carter, 1882. torted, fairly abundant, scattered to clustered inter- use stitially, 6.8-72.5-17.9 x 0.9-1.2 ^m. Remarks. Phloeodictyidae was dropped from 120 F. W1EDENMAYER

in taxonomy by Topsent (1928: 66; as Phloeodic- Strait, 70 m; off Sydney, 55-64 m; off Bahia?).- 66-71 tyinae, in the sense of Ridley and Dendy, 1887), Whitelegge, 1906: 466 (off Sydney, m). 1 Phloeodktyon putridosa (sic). -Burton, 1928: 118(16 who then preferred to place Phloeodktyon mi. E of Devi River, Orissa Coast, India, 124 m). (Oceanapia) in the Renierinae Ridley and Dendy. ? Phloeodktyon putridosum! .-Wilson, 1925: 419 Carter's senior synonym has not been used since (Philippines). then. In my opinion, ICZN Article 40 does not Phloeodktyon putridosum. — Topsent, 1933:42, pi. 3 this Soest (1980: 85, 96, apply in case. Though van fig. 8 (description of holotype, discussion). 97) synonymised Phloeodktyon and Oceanapia and Material examined. Station BSS 187, one specimen mentioned Phloeodictyinae in his historical survey, (F52037). his formal erection of Oceanapiidae (same work, pp. 80, 1 14) is not a stated replacement of Phloeo- Diagnosis. Cake-shaped, with low and stubby, dictyidae, based on such generic synonymy. irregular fistules bearing small, chiefly lateral Carter's family name, moreover, is here judged to oscules. Compressible, resilient, easily torn; deep be invalid pending an appeal to the International brownish red in life. Surface smooth to finely wrin- Commission for its suppression, and Oceanapiidae kled, with scattered sediment. Bark-like ectosome can be said to be in general current use (ICZN stratified, with 2-4 matted layers, each 1 mm thick, Article 79c). separated by floors of same height with scattered spicules and sparse spicular pillars. Choanosome Oceanapia Norman fleshy, with ill-sorted detritus and macrosymbionts.

Oceanapia Norman, 1869: 334. — Ridley and Dendy, Oxea variable, in 3 indistinct categories. 1887: 32, 36 (in Phloeodictyinae). -Dendy, 1922; 45 (defi- Description. Incomplete, torn off its base by nition in Phloeodictyinae).— Topsent, 1928: 66 (in Renie- dredge, subsequently damaged. Base probably rinae). — Bergquist, 1965: 160 (discussion in originally cake-shaped. Preserved top, with jagged Adociidae). — Levi in Brien et al. 1973: 620 (definition in

torn outline, 10 1 1 wide, 1.5 Renieridae).-Vacelet et al., 1976: 98f. (2 species x cm cm thick in described). — Van Soest, 1980: 85 (synonymy, definition), middle, almost flat on one side, thrown into irregu- 91 (discussion), 114 (definition), 115 (illustration). — lar, oblique adjoining turrets or fistules on other. Bergquist and Warne, 1980: 37 (definition and discussion One of these composite, bifid at top. Their height in Oceanapiidae). and width 2-3 cm. All three bear small, irregularly Rhizochalina Schmidt, 1870: 35. - Ridley and Dendy, scattered oscules chiefly on one side, 1-2 mm wide, 1887: xiv, 32. -Bergquist, 1965: 160 (discussion). - some clustered. One apex of composite middle Vacclet et al., 1976: 92-96 (7 species described). — Van turret bears larger oscule with contorted rim, Soest, 1980: 85, 92, 1 14 (synonymised with Oceanapia). diameter 4-5 Other half of base bears Phloeodktyon Carter, 1882a: 122. -Dendy, 1905: 165 mm. two low

elevations, 1 (definition in Phloeodictyinae). — Dendy, 1922: 47 (defi- 0.5 and cm high and wide, with one nition in Phloeodictyinae). — Wilson, 1925: 419 (defini- small lateral oscule each. Marginal surface tion in Phloeodictyinae). — Bergquist, 1965: 160 encrusted by platy calcareous alga, in turn over-

(discussion). — Van Soest, 1980:85,91, 1 14 (synonymised grown by small unidentified sandy sponge, algae with Oceanapia). and hydrozoans. Biminia Wiedenmayer, 1977a: 124 (definition in Deep brownish red in life, now faded to deep Nepheliospongiidae). — Van Soest, 1980: 1 15 (definition fawn (7.5-10 YR 6-7/6-8). Compressible, resilient, in Oceanapiidae, probably synonym of Oceanapia).— easily torn. Hoshino, 1981: 121 (in Adociidae, new species).— Surface smooth, finely Hooper, 1984b: 55 (in Oceanapiidae, new species). wrinkled over base, contains scattered sediment. Ostia visible at intermedi- Diagnosis. "Oceanapiidae in which the ectosomal ate magnification, clustered over meshes of bark-like skeleton is reinforced by spongin, this subjacent matted layer, between single tangential trend being particularly evident in the fistule walls. oxea. Underlying meshes about 100-300 /tm wide, Megascleres oxea; microscleres, when present, sigmata or [and] toxa." (Bergquist and Warne, 1980: 37.)

Oceanapia putridosa (Lamarck)

Plate 12 figure 8, text-figure 82

Akyonium putridosum Lamarck, 1815 (1814-1815): 168 (King George Sound, WA, fide Topsent, 1933). 1 1 Rhizochalina putridosa . . — Ridley and Dendy, 1887: 33, pi. 8 fig. 5, pi. 9 figs 1, 7 (Moncoeur Island, Bass Figure 82. Oceanapia putridosa, oxea, x 274. SPONGES FROM BASS STRAIT 121

often irregular in outline (composite, petaloid or [Euspongiaf] Bronn, 1859: 22(nomen vanum, see Bur- lobed). Widths of tangential fibres in same order. ton 1934a: 574).- Wiedenmayer, 1977a: 54 (further Below surface, 2 to 4 successive matted layers, synonyms). Ditela Schmidt, 1862: 24. mostly parallel, about 1 mm thick, separated by discontinuous floors with sparse single spicules and Diagnosis. "Spongiidae in which the primary fibres fibrous pillars, their height equal to thickness of are reduced in number and the highly developed matted layers. Some floors wedge out, with con- secondary network of fine, intertwined fibres tiguous matted layers merging. Inner matted layers makes up the bulk of the skeleton. Primary fibres

frequently riddled by round openings about 1 mm contain a central axis of foreign material, and are wide and 1-5 mm apart. most in evidence near the sponge surface. Secon- Remaining deeper choanosome indistinctly strati- dary fibres contain no foreign material. The tex- fied, fleshy, contains ill-sorted detritus. Main skele- ture of the whole is springy and very compressible,

ton typical for genus, with stringy terete fibres in supple and elastic. The surface is [rarely] armoured, all directions, indistinctly graded by sizes of fibres is covered with low, even conules, and most fre- and meshes, occasionally with stronger more con- quently is pigmented black, brown, or grey; the tinuous fibres in planes parallel to surface, and interior is white to beige. The form of the sponge interstitial oxea. Numerous commensal barnacles is variable, but commonly massive spherical, lamel- inside fistules. late, or cup-shaped." (Bergquist, 1980b: 450.) Spiculation: oxea, mostly hastate, with conical points of variable length, occasionally sub- Spongia hispida Lamarck mucronate, commonly curved or bent, 90-154-229 Plate 12 figures 9-12, plate 34 figure 2 x 2.5-5.9-13.8 nm. Spongia hispida Lamarck, 1814 (1813-1814): 452 Remarks. The dry holotype, as described and partly ("Southern Seas", Peron and Lesueur collection). -de

figured by Topsent, is also badly preserved and Laubenfels, 1948: 15, pi. 3 fig. 6 ( = pl. 2 fig. 6 in Top- fragmented. The spicules agree in shape and meas- sent, 1933). ICacospongia urements including an indistinct segregation into mollior.— Ridley, 1884a: 378 (nee Schmidt; see Lendenfeld, 1889b: 256; Torres Strait). three categories (holotype: intermediate ones, 140 Euspongia irregularis Lendenfeld, 1885h: 485.— x 7-8 fim, dominating; larger ones, up to 185 x Lendenfeld, 1888: 132. -Lendenfeld, 1889b: 245 (collec- 12 fairly frequent; smaller ones, 130-140 x fim, tive name for following varieties, no nominotypical 2-3 nm, scarce). My identification is corroborated subspecies designated). — Br^ndsted, 1926: 295 (New by the presence of a threefold bark layer with two Zealand). intervening hollow layers in the holotype, and by Euspongia irregularis var. tenuis Lendenfeld, 1885h: the presence of sand at the surface. Topsent 485 (Torres Strait and Long Reef, Qld; Mauritius; remarked on the differences of the "Challenger" Chatham Islands, NZ).- Lendenfeld, 1889b: 251, pi. 13 specimens, a much more fibrous structure and uni- fig. 5 (old record). Euspongia irregularis \ax.jacksonia Lendenfeld, 1885h form spicules measurements, but finally accepted 485, pi. 36 fig. 3 (Sydney Harbour). -Lendenfeld, 1888 the identification. Neither Ridley and Dendy nor 133 (as jacksoniana, old record). — Lendenfeld, 1889b Wilson (who indicated only 150-160 x 8 urn for 254, pi. 21 figs. 8, 9 pi. 22 fig. 17, pi. 29 fig. 1 (as jack- multiple the oxea of his specimen) mentioned a soniana, old record). bark layer. Pending a re-examination of these speci- Euspongia irregularis var. silicata Lendenfeld, 1885h: mens, the question as to their identity with 485 (St Vincent Gulf, SA; Sydney Harbour; Fiji; Chatham Lamarck's and the new specimen must remain Islands, NZ). -Lendenfeld, 1888: 133, 134 (old open. records). — Lendenfeld, 1889b: 255 (old records).— Whitelegge, 1897: 331 (Funafuti). -Whitelegge, 1901:95, Order Dictyoceratida Minchin, 1900 117, pi. 12 figs. 17, 17a (beaches of NSW). Euspongia irregularis var. lulea Lendenfeld, 1885h: 485 Spongiidae Gray, 1867 (Mauritius). -Lendenfeld, 1889b: 250, pi. 12 fig. 9 (old Spongia Linne record).

Euspongia irregularis var . dura Lendenfeld, 1889a: 31 Spongia Linne, 1759: 1348 (part).- Vacelet, 1959: 73 (in key). - Lendenfeld, 1889b: 251, pi. 13 fig. 4 (East coast (definition, discussion, revision of Mediterranean spe- of Australia; Sri Lanka; Madagascar). cies).— Wiedenmayer, 1977a: 54 (synonymy in part: not Euspongia irregularis var. villosa Lendenfeld, 1889a: Ceratodendron, Aphrodite ( = Hippospongia); additional 31 (in key). -Lendenfeld, 1889b: 252 (Australia; Chatham - references, definition, discussion, West Indian species). Islands, NZ; Madeira (?); Cape of Good Hope (?); Van Soest, 1978: 8, 19 (definition, revision of West Indian Karachi). — Hentschel, 1912: 435 (Aru, Indonesia). species). - Bergquist, 1980b: 450 (synonymy, definition). Euspongia irregularis var. frondosa Lendenfeld, 1889a: :

122 F. WIEDENMAYER

31 (in kcy).-Lendenfeld, 1889b: 253, pi. 22 fig. 1 (Indian toms. Relief less pronounced in life (colour slide Ocean). after collecting). Membranes pierced by small Euspongia - irregularis var. mollior (nee Schmidt). oscules, some in series, 1-5 mm wide. Lendenfeld, 1889b: 256 (in part: East Indies; Torres Strait, Almost black to dark grey, with reddish tinge in old record based on Cacospongiu moltior: Ridley, life (10 R 3-4/1), now faded (10 R 4-5/2), only 1884a). -Topsent, 1897: 484 (Ambon, Indonesia). partly preserved, dermis having disappeared near Euspongia irregularis var. areolata Whitelegge, 1901 base and along edges of some ridges. 96, pi. 12 fig. 18 (Lake lllawarra, coast near Sydney, Bottom, now NSW). as in life, yellowish beige (10 YR 8/4). Choano- Euspongia irregularis var. stiriganensis Wilson, 1925: some lighter, almost cream. Spongy, moderately 486 (Surigao Strait, Philippines). firm. Euspongia irregularis var . hisptda. -Topsent, 1933: 1, Surface on prominent parts of outer side finely pi. 2 fig. 6 (two syntypes of Lamarck described, lecto- conulose to hispid. Conules mostly composite, type designated and figured). about 1 mm high and apart, bifid to trifid, with Spongia zjmocca irregularis, -de Laubenfels, 1948: 14 barely protruding primary fibres; connected by (in review). -de Laubenfels, 1954a: 6, texi-fig. 2 ridges. (Micronesia). Hispidation less prominent on inner side. Ostia not seen. Spongia irregularis. - Desqueyroux-launde/, 1981: F52039 (pi. 12 fig. 10) much 754, figs 95. 97, 130(rcdescription of record in Topsent, smaller, digitate-ramose. Two short, parallel 1897). branches, 2.5 cm long and 8-10 mm thick, out of base of main branch about 9 cm long, 1 . 5 cm thick Material examined. Station KG 7, one specimen (F52038); proximally, 1 cm distally. Latter station BSS 187, one specimen (F52039). Type specimens much constricted, almost severed 3 o( Euspongia irregularis varr. tenuis, jacksonia, silicata, cm above base. lutea, villosa, frondosa Lendenfeld (AM and BMNH). All branches have grooves down from apex along E. and of irregularis var. areolala Whitelegge (AM). one side for about 1 cm, with smooth walls and few small oscules partly closed. Similar grooves less Diagnosis. Shape variable, massive-lobate, lamel- conspicuous further down sides. late, erect or repent-ramose, often irregularly Surface dull greyish red in life (now more brown, anastomosing. Spongy; externally black to dark 10 R 4-5/2-4); choanosome paler. Alcohol stained grey, often reddish in life. Surface conspicuously orange-brown. Conules less prominent than in and regularly conulose, hispid after collecting. F52038, with primaries occasionally protruding. Dermal membrane thin, partly deciduous after col- Dermal membrane better developed, showing retic- lecting, frequently contains some foreign inclusions ulum of collagen nervures with main lines converg- (spicular debris). Choanosome dense. Main skele- ing in conule tips, parallel with subjacent ton irregular, with straggling primaries always con- uppermost secondaries. taining detritus, with pale spongin; secondaries of Choanosome dense, opaque, more so in F52039, variable thickness, clear. Some detritus also riddled by anastomosing canals about 100-300 ^m interstitial. wide. Matrix contains abundant globular microsymbionts, of different kinds in two speci- Description. F52038 (pi. 12 fig. 9) larger, irregu- mens. Main skeleton in both with straight to slightly lar, capricious. Gross shape like upper quarter of straggling primaries, 28-47 ^m thick, rarely ovoid body. Base deeply excavated. Outer, convex branched, spaced irregularly 200 jtm to about 2 side thrown into parallel to radial, erect ridges, 5-10 mm, filled with foreign detritus, chiefly spicule mm wide, separated by grooves of similar width. fragments, many almost complete, generally well Some ridges branch downward, most obliquely and aligned with tips protruding here and there, hence irregularly dissected by anastomoses between neigh- knotty contour of some fibres. Sand grains or other bouring vertical grooves, hence a succession of debris rare. Encasing pale yellow spongin barely irregular hummocks and pinnacles. Single large visible. Debris (spicules, lithic fragments, oscule, 12 mm wide, with raised collar, cuts across foraminifera) also interstitially, loosely scattered 2-3 ridges near top. Inner, truncate side with fainter or clustered. Secondary fibres and their meshwork relief, irregularly cutting across several erect lamel- irregular, with little if any relation between thick- lae. Base excavated by 3 to 4 arches, as if sponge nesses (4-32 nm) and meshsizes (20-390 fim), except had grown over as many cobbles. From arches, thicker fibres occasionally forming ill defined fas- several lacunae, 0.5-1.5 cm wide, reach deep inside, cicles with more uniform meshes. Secondaries toward top. Grooves on outer side variable straight in to slightly bent between junctions, pale, depth, few millimetres to over 1 cm. still Some yellow, always clear, enclosing triangular to poly- covered by dermal membrane, now much receded, gonal meshes. Dermis in both specimens armoured partly collapsed, partly still stretched above bot- by spicule fragments. SPONGES FROM BASS STRAIT 123

Remarks. The species is externally highly poly- onymy accepted), 456 (type species misidentified as C. morphic, as the synonymy shows in accordance ]oliascens). with Lendenfeld's taxonomic concepts. F52038 ILendenfetdia Bergquist, 1980b: 456. does not fit any of the described varieties well, but Diagnosis. "Spongiidae in which the primary fibres can be interpreted as a more complex growth form are cored with detritus and do not run in regular of var. areolaia, in which secondary, lateral erect fashion from surface to surface of the compressed lamellae are attached at right angles to mesial body. The fibres wander at all angles to the sur- lamella bearing large oscules along its rim. F52039 faces, branch, and frequently are in weak fascicles. can be conceived as a young stage of the more com- The secondary network is very highly developed, plex var. jacksonia (stubby, digitate branches, see made up in the main of very thin, tangled fibres pi. 12, fig. 1 1) or of the typical variety (Lamarck's branching and anastomosing, occasionally fascic- specimens: longer, tangled branches). ulate. Fine tertiary fibres are apparent in some spe- As in Carteriospongia (see below), the compo- cies. Secondary fibres are clear of detritus. The sition of the detritus filling the primaries has no surfaces of the sponge are [commonly?] reinforced systematic importance. The almost exclusive con- by a sandy cortex which is often quite thick; the tent of foreign spicules (used by Lendenfeld to surface itself is usually marked by fine ridges and separate var. silicata), of grains only, and of mixed grooves. The texture is compressible, and the shape types seem to occur indiscriminately. of the sponge is variable, but the body is always The armoured dermis, not previously recorded thin, compressed lamellate or cylindrical." (Berg- in this species, necessitates the slight modification quist, 1980b: 454.) of the generic diagnosis (above) with respect to Bergquist's (1980b: 450). Remarks. There is still some confusion about the The affinity with Cacospongia mollior, claimed taxonomy of this genus, particularly about its type

by Lendenfeld ( 1 889b), can be excluded, given the species, the synonymies and correct names of the differences in skeletal structure (Bergquist, 1980b: species assigned to it, and to whether it is really 460; Pulitzer-Finali and Pronzato, 1977: 92, fig. distinct from Phyllospongia. Burton (1934a: 572) 3). The affinity with Spongia zimocca, claimed by designated Spongia foliascens Pallas as type spe-

de Laubenfels, is doubtful. The latter species is cies of Carteriospongia, stating that this is a senior primarily East-Mediterranean, and Australian and synonym of the first species described in Carteri- East Indian records should be re-examined. The ospongia by Hyatt (1877), Spongia otahitica Esper true nature of the skeletal structure of topotypes (as Carteriospongia otahitica Hyatt). 1CZN Article

of S. zimocca is still not firmly established (see 69a (v) would apply here if Burton's designation Vacelet, 1959: 78). were valid. He overlooked the fact, however, that already Ridley (1884a: 385) had effectively desig- Carteriospongia Hyatt nated as type species the first species described by Hyatt in his new genus, but in another sense than

Carteriospongia Hyatt, 1877: 540 (definition and dis- Burton. Ridley (loc. cit.) stated: "Hyatt formed the cussion in Phyllospongiadae, 5 species included, another genus Carteriospongia nominally for a species one questionably).— de Laubenfels, 1936a: 21 (definition called by him otahitica, Esper, which is, however, in Spongiidae). - Bergquist, 1965: 132 (discussion, apparently lamellosa, Esper, to the plate of which 1980b: implicitly merged in Phyllospongia). -Bergquist, he refers". This statement is sufficient to consti- Spongiidae, discussion, 454 (synonymy, definition in tute a subsequent designation (ICZN Article 69a revision). (iv)), and as such has priority over Burton's. It is [Carterispongia] Ridley, 1884a, b: 385 (nomen vanum; also clear, however, that Ridley regarded Hyatt's synonymy, discussion, valid subsequent designation of first described species as misidentified, a fact cor- Carteriospongia otahitica sensu Hyatt (non Spongia otu- roborated by Hyatt's explicit description, which hitica Esper) as type species, subject to decision by Inter- national Commission, description of Australian records), leaves no doubt that he had before him Carteri- 594 (description of Indian Ocean records). -Burton, ospongia lamellosa (Esper) (the junior synonyms 1934a: 572 (invalid subsequent designation of Spongia of which are Spongia polyphylla Lamarck, Spon- foliascens as type species), -de Laubenfels, 1948: 126 (dis- gia laciniata Lamarck, Carteriospongia radiata cussion as nomen vanum). Hyatt, Mauricea lacinulosa Carter, and Carteri- de Laubenfels, 1948: 126 (nomen [Carterospongia] ospongia pennatula sensu Ridley et auct.). nullum; merged in Phyllospongia). The type species of Carteriospongia cannot be Poly]ibrospongia Bowerbank, 1877: 459. -Bergquist. cited as Spongia otahitica Esper (ICNZ Article 49), 1980b: 454 (merged in Carteriospongia). nor can it presently be cited as Spongia lamellosa Mauricea Carter, 1877: 174. -Ridley, 1884a, b:385, 594 Article 70b). Pending a decision by (merged in Carteriospongia).- Bergquist, 1980b: 454 (syn- Esper (ICZN 124 F. WIEDKNMAYER

Phvllospongia (Antheroplux) ealkiformis. -Lenden- the International Commission in this matter, the (Weslcrnport Bay, Vic.; fcld,'l889b: 182, pi. 5 fig. 1 type species of Carteriospongia has tor now to be Sydney Harbour). regarded as void. An eventual acceptance of Spon- Phyllospongia calyeiformis. - Bergquist and Skinner, gia lamellosa as type species would not affect the Australia). 1982, pi. 4 fig. 1 (colour, southern pen- scope of Carteriospongia, since this species (as Phyllospongia ulienu Wilson, 1925: 481, pi. 51 fig. 10 Bergquist natida) is also included in this genus by pi. 52 figs 1,3, 5 (Philippines). (1980b). Phyllospongia and Carteriospongia were ICacoSpOngiapoCUlum Selcnka, 1867: 567, pi. 35 fig. apparently kept distinct, though diagnosed together, 7 (Port Phillip Bay). Levi (in Brien et al., 1973: 625), but later merged by Material examined. Station KG 3, two specimens by Berg- (Vacelet et al., 1976: 106). As diagnosed (F52040a,b). Syntypes of Carteriospongia ealkiformis quist (1980b: 454), the two genera seem sufficiently Carter (BMNH).

distinct, but it remains to be demonstrated whether conical cups with single newer specimens of the species complex Phyl- Diagnosis. Wide-mouthed Smaller specimens smooth or lospongia papyraeea (with holdsworthi, madagas- or multiple stalk. sculptured on both sides, larger specimens carensis, distorts, coriacea) always hold up to weakly coarsely rugo-reticulate on Bergquist's diagnosis. Until such lime, Carteri- with regular sculpture, outside, more meandrine or vermiculate in con- ospongia is here retained. For possible synonymy frequent in concavity of of Lendenfeldia, see below, remarks on C silicatu. cavity. Secondary cups larger specimens. Oscules small, regularly scattered specimens. Carteriospongia ealiciformis Carter in concavity, inconspicuous in large Firmly rubbery, more resilient along rim; purplish 1-7, plate 34 figure 3, Plate 13 figures or pinkish brown, with lighter patches in life. Sur- side text-figures 83, 84 faces micropapillate. Ostia on inhalant outer scattered to crowded, tending to segregation by 2 ectosome 150-500 thick, Carteriospongia ealiciformis Carter, 1885b: 221 (Port sizes. Armoured i±m Phillip Bay, Vic). - Bergquist, 1980b: 454 (good species). underlain by crypts on outside. Choanosome caver-

co o o o

^ o 83

Figures 83, 84. Carleriospongia ealieiformis. F52040a, details in reflected light. Figure 83. View of inhalant surface, x 11.7. Figure 84. Perpendicular section, x6.3. Fxhalanl surface above SPONGES FROM BASS STRAIT 123

nous, canals often of 2 sizes. Interstitial detritus. reticulation on parts of surface, with oscules in pits. Main skeleton typical for genus, dense, with closely Meandering and dendritic subdermal canals with spaced, distinct primaries. crowded smaller oscules particularly in smoother areas with fewer single oscules. Large portions of Description. Wide-mouthed calices. F52040a outer surface, particularly in F52040a, with distinct, larger, cup somewhat compressed, asymmetrical, irregular reticulum of subdermal canals about 200 sinuous margin descending to about half height of /tm wide, meshes 1-3 mm wide, riddled by larger opposite wall on one broadside. Opposite radii of ostia, 135-330 /an wide, irregularly scattered. Inter- inner wall, from bottom to rim, 10.5 and 8.5 cm stices pierced by crowded smaller ostia, around 65 along longer axis of ellipse, converging at 120"; /an wide. Where reticulum indistinct, little segre- those along shorter axis are 9 and 6-7 cm, converg- gation of ostia by size, spacing and areas (text-fig. ing at 110° (inclination steeper along lower wall). 83). Armoured ectosome of F52040b 125-375 /an Width at mouth 12x17 cm. Asymmetrical pedicel thick on inner surface, mostly 150 /mi thick on below conical bottom inclinded 30° away from axis outer surface, below which a zone of extensive of cup, slightly bent, expanded below into sprawl crypts. Choanosome riddled by complex canal ing holdfast with perpendicular base. Pedicel 3 cm system, with two size ranges, intergrading occasion- long in middle, 1-1.4 cm thick; elongate base of ally: (1) canals 250-750 ^m wide, predominantly holdfast 3x6 cm wide. Cup wall 6-7 mm thick transverse; (2) 65-125 /un wide, reticulate, com- near bottom, 2-3 mm at sharp rim. F52040b a simi- monly oblique (text-fig. 84). Main skeleton of larly shallow, compressed cup with asymmetrical F52040b comprises closely spaced primaries rim. Inner radii 3.5 to 7.5 cm, width across rim 8.5 (350-750 /an apart), knotty, 100-135 /an thick, x 1 1 cm. Five thin pedicels from horseshoe-shaped occasionally with lacunae, charged with foreign ridge surrounding concavity. 2 cm wide, at bottom debris (mostly spicule fragments, poorly aligned, of cup. Pedicles inclined outwards radially, con- sometimes discontinuous). Irregularly scattered torted, wedging out into little expanded holdfasts; detritus also intcrstitiaily. Secondaries 18-65 /'in proximally 5-10 mm wide, 2-4 cm long including thick, (angled and crowded, mostly oblique in all holdfasts. directions, with indistinct meshes. Here and there, Purplish to pinkish brown in life, lighter in some particularly between closely spaced primaries, areas, now mottled beige, with greyish, brownish, ascending and transverse secondaries evident, with yellowish and orange tinges. Firmly rubbery across meshsizes around 100-200 /an. All secondaries with rim, little compressible across cup wall, less across darker axial filament; thicker, more continuous pedicles. ones (more deeply stained by basic fuchsin) dis- Outer surface of both cups and of pedicels tinctly stratified in periphery. Thinner, shorter ones smooth in general, but minutely granular. Stronger, have trumpet-shaped junctions cut off by continu- coarser granulation on inner surface. On both sides ous wall of perpendicularly joined thicker fibre, as of rim, granules mostly aligned radially, to 1-3 cm if grafted. Base of such junctions frequently drawn below rim on inside, up to 5 cm deep on outside. out to join base of neighbouring transverse secon- Individual granules of inner, exhalant surface fre- dary, forming additional layer on wall of joined quently confluent, in short meandering and branch- thicker fibre. ing ridges, 350 to 500 /an wide, interstitial grooves Inhalant ectosome in F52040b comprises outer slightly narrower. zone, 100- 160 /an thick, with packed spicule frag- In F52040a, small oscules regularly scattered over ments, pierced by ostia, and inner zone, 230-350 inner surface, but almost lacking in bottom (width /an thick, with scattered to crowded spicule frag- 4-5 cm), spaced about 3 mm, single or composite ments, traversed by primaries, generally free of (clustered), 200 /mi to 1 mm wide, now partly secondaries, containing crypts. closed, in shallow depressions of vaguely rugo- Main skeleton below exhalant surface irregular, reticulate pattern. Oscules often at centre of radi- tangled, with primaries indistinct, fasciculate, ally confluent subdermal canals about 200 /an wide. oblique and branching, ending in elevations (gran- In bottom, similar canals arranged in dendritic to ules) of surface; spaces below intervening depres- loosely reticulate pattern, often pierced by crowded sions almost devoid of secondaries and contain oscules about 100 /an wide. Similar pattern, but small atria. More interstitial detritus than below more radial, below rim, coinciding with interstices opposite surface, with many lithie grains. of aligned granules. In F52040b, oscules more irregularly and widely Remarks. Fxpanding juctions of secondaries spaced on inner surface, mostly at 5 mm, but scarce enveloping primaries agree with what Wilson (1925) in some areas. Coarser, more prominent rugo- described and figured for Phyllospon^ia aliena. The 126 F. WIEDENMAYER

cementation of a crossing secondary of similar 1893.3.4.24, Great Barrier Reef; BMNH 1893.3.4.63, -69, unpublished). thickness by a peripheral layer of the crossed fibre Bass Strait, latter locality (Wilson, 1925, pi. 52 fig. 3) was also observed, as Diagnosis. Asymmetrically saucer-shaped, rim were some knobby ends of free fibres, however often elliptical, with short stalk. Surfaces smooth overgrown by a new outer layer extending into a or sculptured, pattern of latter strongly and fibre of thinner diameter continuing in the same coarsely rugo-reticulate, rugo-vermiculale or tuber- direction beyond the knob of the first stage. Car- culate, except for rims, which have a weaker, teriospongia foliascens may be quite similar to the radial, scalloped sculpture. Surface otherwise present species in shape, though its cup-shaped wall micropapillate, lipostomous in life; papillae radi- is often spirally coiled. The main distinction there, ally aligned around rims. Rubbery; dark red to red- however, is the presence, on the inner surface, of brown on exposed parts, pale vermilion in depres- a regular and prominent primary pattern of sions, in life. Armoured ectosome not over 150 /mi crowded papillae or radially oriented vermiculate thick, often patchy, evanescent or lacking. Dry ridges with narrow interstices. This pattern (see specimens hence often with irregular whitish Bergquist, 1965: 131; Levi in Brien et al. 1973, fig. patches, surface otherwise radially striate (align- 428) is much coarser and more prominent than the ment of primary fibres in periphery). Deeper main granulation of C. caliciformis, which is also men- skeleton more irregular, primary fibres indistinct, tioned by Wilson (1925) for C. aliena. Superim- lumpy. posed on the rugo-vermiculale pattern of C. Description. Like a very shallow goblet (pi. 13 figs foliascens, especially in larger specimens, is often 8-10), elaborately and symmetrically decorated. a more irregular secondary one of mamelons and Rim slightly upturned from base, oval outline 6 x lobules (see Bergquist, 1965, fig. 3a; Vacelet, Vas- 7.5 cm in width. Peripheral width of 1-1 .5 cm deco- seur and Levi, 1976, pi. 5 fig. 0- An analogous, rated above by radially arranged, straggling and but different secondary surface relief occurs in C. sparingly branched ridges and grooves. Ridges caliciformis: it is the coarse rugo-reticulate pattern around 1 mm wide, grooves 1-2 mm. Similar pat- already noticed on the inside of the smaller new tern, but much weaker, on underside of rim. Flat specimen. It is much more pronounced in the larger bottom of cup (33 x 52 mm) with much stronger of the two synlypes of Carter here figured for com- pattern of partly anastomosing lobules and ridges, parison (pi. 13 figs. 6-7), where it also characterises 1 to 4 mm wide, up to 5 mm high. Similar, more the outer surface. Both syntypes of Carter, further- rugo-reticulate pattern on base. Compressed, short more, show the tendency, in this species, for the pedicel vertically from base, somewhat off-centre, excrescence of an additional frond or cup from the 12 mm high, thick in bottom. 6x9 mm middle, expanding below into two sprawling rhizomes at right The occurrence of C. foliascens in southern Aus- angles, both about 2 cm long. Total height above tralia is doubtful and needs to be verified. The spe- pedicel 2.5 cm. Thickness of rim 4 mm proximally, cies is common in shallow reef environments in the 3 mm distally. Thickness near centre, including whole Indo-West Pacific Region. Records from erect lobules, 1 cm. Upper face in life dark red- Victoria, New Zealand and New South Wales are brown (10 YR 3-4/2-4) on elevations, pale vermil- based on synonymies and records in Lendenfeld ion (10 YR 7/8) in depressions. Underside was deep (1888: 177; 1889b: 196), whose identifications, par- dark red (5 R 6/3) to dull vermilion (10 R 8/6), ticularly with many supposedly conspecific speci- now faded superficially to uniform beige (10 YR mens, are always suspect, in my experience with 7-8/3-4). Choanosome in centre brownish orange his type series in Sydney and London. (5 YR 6-7/8). Rubbery. Carteriospongia silicata (Lendenfeld) Surface in depressions with fine granulation, barely visible on underside, more distinct on upper Plate 13 figures 8-12, plate 14 figures 1-5 face. Granules fo both sides aligned radially along Phyllospongia (Carteriospongia) silicata Lendenfeld, outer half of rim. Granules on upper side 500-600 1889a: 26 (in key). -Lendenfeld, 1889b: 195, pi. 7 fig. lim wide, occasionally confluent, with narrow inter- 8 (Torres Strait). — Bergquist, 1980b: 456 (in review, as stices. Numerous oscules, 100-600 /mi wide, irregu- unrecognisable). larly scattered, mostly round, occasionally Carteriospongia elegans. — Burton, 1934a: 600 (nee elongate. Underside completely riddled by very Phyllospongia elegans Lendenfeld; Great Barrier Reef). regular fine ostia, barely visible at low magnifica- Material examined. Station WB, one specimen (F52048). tion, with narrower interstices. Holotype of Phyllospongia silicata (BMNH). Three hypo- Armoured ectosome 100-135 /im thick, with types of Carteriospongia elegans: Burton, 1934a (BMNH packed to crowded mixed detritus in variable SPONGES PROM BASS STRAIT 127 proporiions: spicule fragments dominate in some wet syntypes of Mega/opastas arenifibrosa Dendy areas, lithic and skeletal grains in others. Ostia and Frederick (which I did not examine) the ecto- around 85 /tin wide, interstices around 45 /tm. some "sometimes contains a good many broken Peripheral zone in main skeleton with primaries spicules", according to the original description. Of more or less erect, 75-120 /tm thick, spaced Burton's (1934a) three specimens of "Carteriospon- 400-1000 /*m, knotty, lacunose, often discontinu- gia elegans" (Saville Kent coll., two of which are ous below, passing into usually fasciculate secon- identified , on the label (by Burton?) as Phyllospon- daries. Periphery also comprises lax secondary gia silicaia) only the one from the Great Barrier reticulation of clear, pale fibres, crypts, and inter- Reel' (the other two are from Bass Strait) has exten- stitially scattered spicule fragments. sive patches of whitish ectosome still preserved on Deeper main skeleton highly irregular, primaries the lower side. In the other two, and in the holo- thicker (up to 250 /tm), very lumpy, with single very type of Phyllospongia silicaia, the armoured ecto- large lithic grains (mostly quartz) almost exclu- some is reduced to traces. sively, completely enveloped in .spongin. Primaries The salient diagnostic characters in Bergquist's may form irregular meshwork, with meshes (1980b: 456) definition of Lendenjeldia are: (1) 320-560 /tm wide. Many "primaries" merely lumps absence of a sand cortex, (2) absence of grooves of 1-3 grains enveloped in spongin and joined on on the surface, and (3) presence of a tertiary fibre 2-4 sides by secondaries. Secondaries 25-60 /tm net. All three seem to be of doubtful value, prob- wide, mostly straight, either crowded or vaguely ably consitituting differences of degree only. fasciculate, particularly below periphery, or form- A common growth form in Carleriospongia ing lax angular meshwork, with meshes up to 420 silicaia is that of wide, shallow cups with a short /tm wide. pedicel, with the concavity (and to a lesser degree All fibres of deeper choanosome rusty in trans- the outside) usually decorated by more or less regu- mitted light, in shagreen-like pattern, like leopard- lar protuberances. The concavity of large specimens skin with high magnification, corresponding to also contains irregular frond-like excrescences in brownish orange central zone noted above. Its the bottom. According to J.N. A. Hooper (personal boundary not always that with peripheral skeleton, communication), numerous specimens of this spe- may reach deep inside, or almost to ectosome. cies in the Great Barrier Reef are without obvious

Microscopic pattern occasionally on two successive pedicel, and lie flat on the substrate, or raised only fibre layers, frequently overgrown by clear spongin. slightly above it. Some thinner fibres form short anastomoses between adjoining, mostly parallel secondaries, Carleriospongia cf. vermicularis Lendenfeld generally 12-18 /tm wide, possibly tertiary fibres, but intergrading with secondaries in diameter. Their Plate 14 figure 6, plate 34 figure 4 "grafted" junctions trumpet-shaped (as noted in C. ?Phyllospon%ia (Carleriospongia) vermicularis Lenden- caliciformis), usually accentuated by rusty layer. feld, 1889a: 26 (in key). -Lendenfeld, 1889b: 201, pi. 15

fig. 5 (west coast of Australia). Remarks. 1 have fomerly assigned this species to ICarteriospOngia vermicularis. — Burl on, 1934a: 573 Lendenjeldia (Wiedenmayer et al., in press), rely- (Eagle Island, Great Barrier Reef). ing on macroscopic characters only. In examining ? Phyllospongia vermicularis.— Br^ndsted, 1934: 26 dry material of the type species of Lendenjeldia in (Bandanaira, Indonesia). London (BMNH), i.e. L. plicala (Esper) (with syn- Material examined. Station BSS 180, one specimen onyms lamellaris, torresia, dendyi with varieties, (F5204I). ridleyi with varieties, polyphemus and areni- Single straggling branch, 4.5 cm long fibrosa), I noticed the common radial alignment Description. of primaries and their tips (conules) near the rim (without base, now dissected), variably terete to on both sides of the fronds, often affecting the slightly compressed, proximally 6 to 3 mm wide, whole outer half of the frond surface. This was also tip subclavate, 15 mm long, 7 mm wide. Partially noticed on the dry holotype of Phyllospongia incorporated lacy bryozoans and fragments of clam silicaia and in Burton's dry specimens mentioned shells, including an almost complete Pecten valve. in life (2.5 8/2-4), now slightly above, where it obviously corresponds to the radi- Yellowish beige Y ally aligned granulation in the new specimen. A darker (10 YR 7/4). Spongy, resilient. Surface similar pattern, though less pronounced, was noted finely conulose to hispid, with conules around 300 above for Carleriospongia caliciformis. /tm high and 1 mm apart. Tips of primaries pro- No trace of an armoured ectosome was noticed trude here and there, especially around apex, for in the dry specimens of L. plicala. However, in the up to 300 /tm. Surface contains conspicuous, seal- I

128 F. WILOENMAYER

According to tered delrital grains and foraminifera. Surface in cal to somewhat flattened branches." communication), more upper hall' riddled by round orifices about 100-200 J.N. A. Hooper (personal with Indo- fim wide, around 200-300 /tin apart, now mostly than one thinly ramose CarteriospOngia closed, presumably oseules. Special, presumably Malayan and Australian distribution occur, hence pending availability of the inhalant field on lower half of one side of subcla- I prefer an open name vate lip, free of conules, with polygonal orifices lost holotype or of similar lopotypes. of similar widlh in regular meshes of reticulum formed by armoured eetosome. Few lacunae in sur- Coseinoderma Carter face, about 0.5 mm wide, one forming longitudi- Coseinoderma Carter, 1883c: 309. -Bergquist, 1980b: nal vermiculate groove 15 mm long, with smooth 456 (definition, discussion, revision). fleshy wall. Spots devoid of armoured eetosome, flesh below shrunk, pripheral main skeleton visible. Diagnosis. "Spongiidae in which the primary fibres Transverse sections near base show scarce strag- are cored and the secondary elements arc clear, gling primaries more or less radial from axial region extremely fine, numerous, and intertwined. Carter's to surface, 55-80 pan thick, knotty, occasionally analogy with 'whorls of wool' was very apt. The branched, often fasciculate, half filled with spic- surface of the sponge is invested with a sand ule fragments and lithic grains in disorderly armour, but the texture remains soft, spongy, and fashion, often off-axis and protruding. Secondaries exlemely compressible. The sponge body is fiabel- of pale beige spongin 10-55 /tm thick, straight to late, pyriform, massive, or pedunculate, with apical bent or curved, in lax meshvvork with meshes or marginal oseules." (Bergquist, 1980b: 456.) 100-400 foo wide. Outer spongin layer occasionally visible on thicker secondaries; "grafted" junc- Coseinoderma pesleonis (Lamarck) tions of thinner ones not as distinct and frequent text-figures 85-88 as in C, caliciformis. Periphery contains lacunae Plate 14 figures 7, 9, 500-600 in main skeleton, /mi deep, traversed by Spongia pes bonis Lamarck, 1814 (1813-1814): 379 primaries only. Eetosome packed with ill soiled (Southern Seas, Peron and Lesueur collection). mixed detritus (spicule fragments, skeletal and lithic Coseinoderma pes leanis.— Topsent, 1930a: 33, lexi- grains up to 700 ^m wide), about 200-300 /

which is now rearranged systematically. There are, Diagnosis. Stalked or compressed-pedunculate, dis- however, still some boxes of unsorted exotic speci- tally globular, pyriform, or thickly flabellatc. mens from the old collection of Gray's and Bower- Oseules commonly elevated, forming apical (and bank's time, which might contain the holotype. lateral) crest, but also irregularly scattered on sides. Burton's specimen has branches up to 10 mm thick. Surface conspicuously sandy in life, with skeletal Br

.4 . °Vl ftwm

o°

0° 00 ^0

^0 o

Details in reflected light. Figure 85. F52042, view of coarsely sandy surface, Fieures 85-88 Coscinoderma pesleonis. perpendicular section (surface on right margin), X6.8. Figure 87. F52043, perpendicular x6 8 Figure 86 F52042, F52043, view of surface (ostia only), x6.8. section x4 1 Figure 88. no !•. WILDENMAYLK

Description. F52042 fairly symmetrical, with Remarks. This is so far the only species of almost globular, slightly compressed top, 5 x 6 cm Coscinoderma recorded from Australia. The West- wide, 6 cm high; inclined pedicel, proximally com- Indian Coscinoderma lanugo de Laubenfels (1936a) pressed and transversely corrugated, 5.5 cm tall, was admitted as second species by Bergquist 7-15 mm wide, slightly expanded at base. Apical (1980b: 458), but is still poorly known. Van Soest crest, 5.5 cm long, formed by series of concrescent (1984: 30), who examined a schizotype, merely elevated oscules. Few raised oscules irregularly scat- stated that it has a thick coat of sand and a skele- tered on either side, another two, little prominent, ton "much more slender" than Coscinoderma! below each end of crest. musicalis (Duchassaing and Michelotti), also West- F52043 more elaborate, with larger, more com- Indian. From van Soest's description and text-

pressed top, 4 x 7 cm wide, 8 cm high, deep figure of the skeleton of the latter, I conclude that

oblique concavity on one side. Apical crest with it is not congeneric with C. pesleonis. oscules less distinct, extended downward on side C. pesleonis is often externally similar to with concavity, curved, branching. Strongly com- Tliorecla latus (Carter; see Remarks on Thorecla pressed pedicel 4 cm long, 4x10 mm wide, ends choanoides, below). The peduncle of the latter is in rhizomatous base. often less individualised, shorter, its oscules are F52044 has top elongated crosswise, higher and flush or slightly elevated, and, if apical, not aligned slightly thicker at one extremity; 72 mm across, 27 in a crest as in C. pesleonis. The skeletal ostia are to 35 mm high, 24 to 28 mm thick. Without crest, more widely spaced in C. pesleonis, punctiform, oscules small and flush, top thrown into irregular whereas they are commonly polygonal or com- ridges outlining shallow contorted depressions. pound, in the meshes of a finely rugo-reticulate or

Compressed pedicel short (1 cm), not expanded at clathrate armour in T. latus. The chief difference

either end, torn off its base. is in the skeletal structure, which makes C. pes- F52045 smallest; terete, transversely corrugated leonis softer, more compressible. pedicel 23 mm high, 4 mm wide; slightly compressed narrow main part gradually expanding to Leiosella Lendenfeld width 23 4 above pedicel, then maximum mm, cm leiosella Lendenfeld, 1888: 120. -Lendenfeld. 1889b: slightly constricted below two unequal distal 201 (part). — dc Laubenfels, 1936a: 21 (definition in Spon- oscules. Total height 9 cm. giidae, invalid subsequent designation of type species).— In life and in alcohol light grey to beige with faint de Laubenfels, 1948: 59 (erroneous reference to first pub- pink or orange tinge. Compressible and resilient, lication, wrong type species). — Bergquist, 1980b: 458 surface firm and slightly rough. (definition, discussion, revision, wrong type species and Weakly magnified surface variable between date). specimens, in average size of sand grains, width and Diagnosis. "Cup-shaped, lobed, or flabellate Spon- spacing of ostia (text-figs 85, 88). Ostia 100-350 giidae with a skeletal network in which the secon- /mi wide, 0.5-1 .3 mm apart in F52042, 150-600 /un dary elements become very dense. The primary wide and equally spaced in F52043. fibres are lightly cored and tend to become fascic- Armoured ectosome 700-800 /tm thick, subja- ulate, either where they arise out of the dense secon- cent zone of crypts 200-400 (im thick (text-figs 86, dary network or just below the surface. The 87). Choanosome traversed by conspicuous, closely secondary fibres also frequently contain some spic- spaced radial canals 0.3 to 1 mm wide, ule detritus, usually as a single discontinuous line

anastomosed at right angles, communicating with of fragments. At the surface there is a thin but 2-2.5 canals 100-200 /tm wide. Few deeper canals, ordered sand armour. The texture is firm." (Berg- mm wide, mostly parallel with surface, probably quist, 1980b: 458.) Oscules small, numerous, regu- extensions of the atria, similar in width. Walls rugo- lar, often on papillae. reticulate. Remarks. It has been overlooked that Leiosella Primary fibres straggling, regularly spaced elegans cannot be the type species, 850-1000 /tm, 55-80 /tm thick, knotty, with spic- not having been among the originally included ule fragements and scarce other debris disorderly species. Euspongia levis Lendenfeld is herein designated type species. enclosed, many protruding, rarely filling whole The species was left in the fibre. Secondaries clear, pale, very long, tangled, genus by Bergquist (1980b) so that the oversight has no further isodiametric around 32 /tm, with faint axial fila- conse- quence in taxonomy. ment; connection by scarce short anastomoses, 15 /an thick, barely expanded and "grafted" at junc- Leiosella caliculata Lendenfeld tions, occasionally torn off (cf. Topsent, 1930a: 34, text-fig. 3). Plate 14 figures 8, 10, plate 15 figures 1-5 SPONGES EROM BASS STRAIT 131

[Euspongia compacta] Carter, 1882a: 106 (junior purplish, greenish, orange-brown tinges, in life. primary homonym of Spongia compacta Sowcrby, 1806; Oscules chiefly on convex side and margins, occa- [in part, not specimen from Port Fairy]; Wollongong, sionally in concavity. Ostia cribriporal or in NSW; "S. Australia"; Vic.). - Lendenfeld, 1885h: 527 (old petaloid groups. records). Surface smooth to micropapillate. Ectosome and choanosome as in generic diagnosis. Leiosella compacta. - Lendenfeld 1888: 120 (East coast). -Lendenfeld, 188: 120 (East coast). -Lendenfeld, Description. Thick asymmetrical frond (pi. 14 fig. 1889b: 210, pi. 13 figs 7 (syntype of Euspongia infundibuliformis Carter), 9 (Fremantle, WA). 8), deeply folded radially, with wide back roughly in Spongia compacta. - Bergquist, 1980b: 458 (in review, one plain, folded forward on both sides, free left as good species). margin, right side completely folded around form- Spongia compactella de l.aubenfels, 1948: l8(nomcn ing oblique cup. Main concavity deeply scooped novum for Euspongia compacta Carter). longitudinally, with faint transverse ridges. Outer [Euspongia infundibuliformis] Carter, 1886g: 374 wall bears faintly scooped or rugo-reticulate pat- (junior primary homonym Spongia of infundibuliformis tern in places. Linne, 1759; Westernport Bay, Vic.). Greater width of main frond 7 cm outside, 5.5 Spongia infundibulis de Laubenfels, 1948: 20 (nomen cm inside, mostly truncate rim 8-14 wide novum for Euspongia infundibuliformis Carter). mm (equal to thickness of wall). Transverse width of Leiosella calyculata Lendenfeld, 1889a: 28 (in key, main concavity, nomen imperfectum). from truncate bottom to back, 1 .5

Leiosella caliculata Lendenfeld, 1889b: 221, pi. 21 fig. cm (jutting folds) to 3 cm (receding folds). Secon-

1 (nomen correctum, for Euspongia infun- dary cup 3 cm long along its inclined and slightly dibuliformis! Carter, undescribed hypotypes; Port bent axis, its compressed mouth 12 X 25 mm wide, Phillip Heads, Vic.).- Bergquist, 1980b: 458 (in review, outer width 3.5 x 4 cm. Two pedicels below as good species). bottom of main cup, their axes 2.5 cm apart, one Coscinoderma concentricum Kirkpatrick, 1903b: 254, below base of small cup 5 mm long, other one, pi. 5 figs 19, 19a (off Port Elizabeth, South Africa). - under free margin of main cup, 2 cm long, obli- de Laubenfels, 1948: 61 (as synonym of Euspongia quely compressed. Greatest width of specimen 10.5 mathewsi Lendenfeld). cm, greatest Carteriospongia concentrica.-Hcvgquhl, 1980b: 458 height, along longer pedicel, 6 cm. (in review). Dark brown-grey with purplish tinges in life, some Euspongia foliacea. — Ridley, 1884a: 378 (nee Spongia areas greenish and rusty brown. Choanosome was foliacea Esper; Torres Strait). lighter brown-grey. Concavity now dull olive- Leiosella foliacea. -Lendenfeld, 1889b: 219, pi. 12 fig. brown (2.5 Y 5/2-4), outside light greyish yellow 12 (syntype of Euspongia infundibuliformis); pi. 20 figs (2.5 Y 6-7/4). Softly rubbery, now firmer. 15, 17-18; pi. 21 fig. 5 (including Ridley's specimen and Outer surface now smooth to finely granular and including Euspongia infundibuliformis Carter as syno- pitted. Circular oscules regularly distributed in nyms; East Indies (new record)). some reas, lacking in others, mostly with regular Lendenfeldia foliacea. — Bergquist, 1980b: 456, 458 (in collars (typical genus), particularly review). of some, on Phyllospongia vasiformis. - lendenfeld, 1889b: 180 truncate rim, flush; width 0.5-1 .5 mm (up to 3 mm across collars). Relief inside finely (part: pi. 28 fig. 1 only). on granulate to ? Leiosella elegans Lendenfeld, 1889a: 27 (in key).— reticulate, somewhat stronger than on outside. Lendenfeld, 1889b: 212 (Lremantle, \VA).-de Lauben- With low magnification, smooth areas of outer side fels, 1948: 60 (resume). bear very low regular conules, higher, with tips of primaries protruding, in granular areas. On inner Material examined. Station KG 6, one specimen (F52046). Type specimens of Euspongia compacta Carter, Euspon- side, relief that of periphery of main skeleton, with gia infundibuliformis Carter, Leiosella caliculata Lenden- ectosome much receding between obtuse tips of feld, Coscinoderma concentricum Kirkpatrick (BMNH). primaries, across meshes formed by outermost Hypotypes of Leiosella compacta: Lendenfeld (AM and secondaries. Occasional vague stellate or petaloid foliacea: Ridley, Leiosella folia- BMNH), of Euspongia patterns visible with higher power, centered in cea: Lendenfeld, Phyllospongia vasiformis: Lendenfeld meshes, not like cribripores in Lendenfeld (1889b (BMNH). Unpublished dry specimen from Victoria in pi. 20 fig. 15). Primaries almost straight, or moder- Dendy Collection (BMNH). ately straggling, or in zig-zag course; mostly spaced Diagnosis. Shape irregular: radially folded (partly 300-400 fim, 30-80 /tm thick, sparsely branched, open) cups, hollow fronds, or concrescent ear- occasionally fasciculate, knotty; contain spicule shaped lobes, with short, single or multiple pedun- fragments almost exclusively, rarely filling whole cle. Outer, convex side may be regularly sculptured, fibre. Secondaries around 25 jim thick, straight to rugo-reticulate, vermiculate or tuberculate. Softly slightly curved, clear, pale yellow. Meshes irregu- to firmly rubbery; Basically dark brown-grey, with lar, 100-525 fim wide. Some spicule fragments scat- 1

132 F. WIEDENMAYER

levis (Lendenfeld) tered interstitially. Dermis around 330 fim thick, Leiosella armoured with scattered to crowded spicule Plate 15 figures 6, 7, text-figures 89, 90 fragments. Euspongia levis Lendenfeld, 1885h: 536, pi. 36 fig. 2 (Sydnev Harbour; Broughton Island, NSW). Remarks. I have examined the only extant syntype Lendenfeld, 1888: 121 (Perth WA; of Coscinoderma concentricum Kirkpatrick 'Leiosella levis.- Tas.). -Lendenfeld, 1889b: 213, pi. 12 fig. 14, pi. 15 fig. (BMNH 1902. 1 1 .16.43, dry), next to the types and Whitelegge, 1901: 94 6, pi. 20 fig. 14 (old records).- hypotypes indicated above. Though they exhibit a (Lakelllawarra, NSW). -Bergquist, 1980b: 458, fig. 7a, wide range of external shapes, I have no doubt that b. they are conspecific. Concave, rounded fronds, Leiosella laevis.- Whitelegge, 1889: 183 (Maroubra, mostly partially fused, transitional to usually asym- NSW). metrical cups are the rule, also combination of cups [Euspongia compacia] Carter, 1882a: 106 (in part, only; Port Fairy, Vic). and fronds. The outer side is often thrown into a specimen BMNH 1877.5.21.1900 relief vermiculate or reticulate ridges which may of Material examined. Station KG 3, one specimen (F52047). be combined with mamelons or papillae. The inside Type specimens of Euspongia levis and hypotypes of is more commonly smooth, but may bear a scooped Leiosella levis: Lendenfeld (AM and BMNH). Syntype pattern, with radial and transverse ridges. Kirk- of Eusnongia compacta Carter (BMNH).

patrick's sponge is somewhat unusual in showing Diagnosis. Shape complex: prolific, contorted, a more prominent relief in the convity, but so does compressed lobes and lamellae, concreseent and the new specimen. The unpublished large, dry anastomosing, sprawling, with indistinct pedicel(s). specimen of this species from Victoria, in the Firmly rubbery; in life, dark grey-brown where Dendy Collection, here figured for comparison (pi. exposed, dark yellowish beige below and in recessed 15 figs 4-5), shows a regular, relatively fine deco- portions. Oscules flush or raised on papillae. Sur- ration on the outside, similar to the one in the speci- face micropapillate above, smooth below. Ostia in men in Lendenfeld (1889b, pi. 28 fig. 1). The pairs, petaloid clusters, or meandering lines. thickness of the dermis is also subject to consider- Armoured ectosome about 1 mm thick, with two able variation. layers. Choanosome with regular canals of two Oscules may also occur in the concave side in sizes. Main skeleton almost devoid of primaries. some specimens, as one of the syntypes of Leiosellu Detritus .scanty in fibres, abundant interstitially. caliculata (BMNH 1955.4.7.3) demonstrates. This

is not surprising in this genus, since ostia always Description. Complex shape, of four elongate seem to occur over the whole surface, i.e. on both lamellae transversely folded and partly coalescent convex and concave sides in flabellate and cup- (pi. 15 fig. 6). Complex, rhizomatous pedicel below

shaped specimens. It may even occur, if the descrip- centre of first lamella, oblique to one side. If tion of Leiosella elegans in Lendenfeld (1889b: 212) oriented in growth position, with base of pedicel

is accepted, that oscules are present only on the flat on bottom, rhizomes facing forward-left, concave side; though (as in his description of lamellae are increasingly inclined backwards, Leiosella caliculata) Lendenfeld may have simply almost recumbent, staggered, completely coalescent overlooked oscules on the outside. on right side, partly coalescent (backs to lower Leiosella elegans (accepted as a good species by fronts) in centre and left. First interstice hollow, Bergquist, 1980b: 458) is likely to be a senior syn- with small round lacuna through base; second inter- onym (by page priority) of L. caliculata. The deco- stice with three elongate lacunae, third open on left ration on the outside of the holotype of the former, side; third interstice with continuous meandering

as described by Lendenfeld, is not incompatible lacuna from open left side to centre. Some lamel- with the range of patterns mentioned above for L. lae, particularly first one, have transverse ridges caliculata. On the other hand, large specimens of on front, some more prominent, incipient secon- Carteriospongia caliciformis may develop a simi- dary lamellae. Base, beyond pedicel, has only slight lar decoration (see above), and the synonymy of rounded swellings and hollows, except open lacu- L. elegans with Carter's species cannot be excluded nae on one side, and two closed small lacunae at present. The holotype of L. elegans could not off-centre. be found during my survey of the dry type collec- Width of specimen (length of the lamellae) 10-1

tion in London (BNNH), but it may eventually be cm, depth in centre (across lamellae) the same; found among the dry exotic sponges of Bowerbank height of main body (perpendicular) 3-3.5 cm;

which are as yet unsorted. However, there is a type pedicel 1.5 cm high, its base 7.5 x 2.5 cm. slide in Berlin (ZMB 6818) which might decide this Dark grey-brown in life, largely preserved in issue. alcohol (10 R 4-5/1-2). Base dark yellowish beige SPONGES FROM BASS STRAIT 133

a * v-r » *~

j/$ -*W%,>' « Mpt.., 89

•• v# .-' # : - v.

Figure 89, 90. Leiosella levis. Details in reflected light. Figure 89. Perpendicular section (surface on top and right), X6.2. Figure 90. View of surface showing ostia frequently grouped in petaloid fashion, x 12.5.

(2.5 Y 7-8/2-4). Firmly rubbery. Oscules on all and regular on lower side, where some areas upper, dark surfaces, including pedicel, more smooth, with single ostia or none. numerous on fronts and ridges of lamellae, always Canal system in periphery regular (text-fig. 89), collared, some conical, with transverse ridges on with primary, ascending canals, 400-670 fjtn wide, backside, occasionally connecting neighbouring spaced 2-2.5 mm, and anastomoses around 400 /im oscules. Collars and ridges distinctly paler. wide, spaced 0.8-1.6 mm. Secondary canals 65-130 Surface between oscules finely granular to /irn wide, frequently fasciculate. unaided eye, that of base smooth. With low mag- Main skeleton almost devoid of primaries. Few nification, upper surface shows regular primary portions of thicker fibres contain debris. Secondary reticulation of sandy ridges (meshes 0.5-1 mm skeleton in great disorder, with tangled fibres 15-30 wide). Conspicuous ostia in depressions, mostly in nm thick, twisted, interlocking, barely discernible pairs or in petaloid clusters (text-fig. 90), or in meshes 35-440 /an wide. Few vague fascicles in meandering short rows, with flush sandy interstices different directions. Much mixed detritus scattered forming secondary meshwork. Primary meshes fre- interstitially, crowded around walls of canals. quently indistinct. Petaloid groups more frequent Armoured ectosome of two layers: (1) outer one, 134 I-. WIEDENMAVF.R

320-420 /tm thick, packed with debris, including filaments. These are usually terminally knobbed, many coarse sand grains; (2) inner one, 320-580 of variable length (order of one to several mm) and

/am thick, with numerous crypts, crowded mixed thickness (below I pm up to 29 ^m, commonly 5-15 debris (chiefly spicule fragments). ^m). They may be scattered or bundled, often forming a dermal or subdermal reticulum, and may Thorectidae Bergquist, 1978 be scarce in some species. Surface conulose, tuber- Synonyms. culate, or pitted. Consistency tough to firm, some- Irciniidae Gray, 1867 (nomen oblitum). times brittle. Stelospongiinae Lendenfeld, 1889a, 1889b (nomen oblitum based on genus dubium). Remarks. I do not think it is realistic to separate Stelospongiidae: Topsent, 1928 (nomen oblitum). Psammocinia as a genus or even subgenus from

Remarks. Irciniidae was published in one of the Ircinia. It is not uncommon in this group (Ircinia most influential early papers on systematica of "sensu lato") to find specimens (species) with an intermediate taxa of sponges. It is regrettable that armoured ectosome, in which choanosomal detri- Bergquist disregarded the availability of this senior tus occurs in primary fibres only, or sparsely also

synonym. Thorectidae, like Oceanapiidae (see in secondaries. This is the condition of most West

above), but unlike other family names, found Indian ircinias (van Soest, 1978), and it charac- general acceptance in the short period since its terises the Australian Ircinia lendenfeldi (senior syn- introduction. Stelospongiidae was originally onym of /. jacksoniana). In Ircinia arenosa Len- applied (as subfamily) to Sielospongia sensu Len- denfeld, a good species of Psammocinia accord- denfeld (species of Fasciospongia, Smenospongia, ing to Bergquist (1980b), sand is present only in Cacospongia, Dysidea, Thoreaa, Hyatiella, the ectosome and in the fibres, not interstitially, Ircinia, "Psammocinia", fide Bergquist, 1980b), if Lendenfeld's figure (1889b, pi. 36 fig. 3) and and Ircinia sensu Lendenfeld (mostly Ircinia s.l). description can be trusted. The presence of very Thoreaa, Thorectandra and Aplysinopsis were large sand grains, "coated individually with spon- included in the Aplysininae by Lendenfeld (1889b). gin, and linked to the skeletal fibres by spongin Topsent (1928: 70) included in the Stelospongiidae strands", mentioned by Bergquist (1980b) in her species ranged in now Cacospongia, Hyrtios, diagnosis of Psammocinia, is probably a monotypic Ircinia s.l. Most species included by Bergquist character, of no importance on this systematic (1980b) in the Thorectidae were assigned by de level. A similar condition is illustrated by Lenden- Laubenfels to the (1948) Spongiidae, partly to the feld (1889b, pi. 36 figs 4, 5) for Ircinia lendenfeldi Spongiinae (those now in Hyrtios, Ircinia s.l.), (as irregularis), regarded as a good species oUrcinia partly to the Verongiinae (those now in Cacospon- by Bergquist (1980b: 466, as jacksoniana). gia, Thorecla, Fasciospongia). Vacelet (1959: 89) regarded Sarcotragus as a subgenus Ircinia Nardo of Ircinia conditionally: valid only in the event that the congruence of thin filaments (below Ircinia Nardo, 1833, col. 519. -Vacelet et al., 1976: 2 nm) and pithed fibres without detritus in two 102, 103 (discussion of widely distributed Indo-Wesl Mediterranean species (/. spinosula and /. musca- Pacific species). — Wiedenmayer, 1977a: 60 (synonymy rum) could be extended to other species, in other with further references; for Euircinia read Euriciniu: parts of the world. Bergquist's diagnosis Sar- nomen negatum [for Ircinia, read Jrciniu\, nomina vana; of cotragus is clarification of type species, revised diagnosis, West weakened in this respect by admitting primary Indian species). — Van Soest, 1978: 33, 41 (definition, dis- fibres with scarce detritus and filaments cussion, revision of West Indian species). — Bergquist, up to 5 jrni thick. She included, besides spinosula 1980b: 464 (synonymy, definition, discussion, revision). and muscarum (both recorded by Lendenfeld from Sarcotragus Schmidt, 1862: 35. — Wiedenmayer, 1977a: Australia, but remaining to be verified as such), 60 (as synonym of Ircinia, further references). — Bergquist, the Australian /. australis (senior synonym of /. 1980b: 466 (as good genus). arbuscula). This, according to Lendenfeld (1889b) Psammocinia Lendenfeld, 1889b: 579 (as subgenus of has detritus only as a discontinuous core in Ircinia). — Bergquist, 1980b: 468 (as good genus). primaries, and filaments 6 /on thick. Four West Diagnosis. Thorectidae in which the primary fibres Indian species described by van Soest (1978) have frequently attain great size by being woven into filaments of 6 pm in diameter and below (down to complex fascicles. Sand extremely variable in abun- 1 jim in /. strobilina), armoured dermis, and detri- dance, coring or charging primaries, or all fibres, tus in most fibres, sparse except in /. felix. Ircinia often occurring also in matrix of choanosome and arenosa was described by Lendenfeld as having fila- in ectosome, occasionally rare or absent. The ments 2-2.5 nm thick. matrix is commonly charged with fine collagenous In conclusion, the characters purported as diag- SPONGES FROM BASS STRAIT 135

nostic for Sarcotragus and Psammocinia by Berg- Material examined. Station KG 5, one specimen (F52049). quist (1980b) are either differences of degree, Syntypes of Hircinia caliculata Lendenfeld (BMNH). mono/oligotypic, or incongruent with her distri- Diagnosis. Thick-walled, bution of species, and thus diataxonic. blunt-rimmed cups, frequently open on one side, or concave fronds, with Ircinia caliculata Lendenfeld multiple pedicel. Firmly rubbery, tough; variable shades of greyish red to orange in life. Strong rugo- Plate 15 figures 8-10, plate 34 figures 5, 6, reticulate sculpture may be present on outer text-figure 91 (convex) side. Whole surface finely and regularly papillate. Small oscules numerous in convavity. Hircinia calyculala Lendenfeld, 1888: 180 (nomen Ectosome imperfectum; West coast, WA; lllawarra and Sydney Har- armoured, 250-400 jim thick* packed bour, NS\V).-\Vhitelegge, 1889: 184 (off Shark Island, with detritus. Choanosome dense, with much inter- Sydney Harbour). stitial detritus. Main skeleton lax, irregular, little Hircinia (Sarcotragus) calyculala. -Lendenfeld, 1889a: fasciculate, thicker fibres generally loaded with 62 (in key). detritus. Filaments numerous, tangled, with irregu- Hircinia (Sarcotragus) caliculata. - Lendenfeld, 1889b: lar, indistinct heads. 572 (nomen correctum, old record). Hircinia caliculata. - Whitelegge, 1901 : 116 (Tuggerah Description. Erect concave frond supported by Beach, NSW). strong, two- or threefold pedicel attached to granite Hircinia campana. — Lendenfeld, 1888: 178 (pan, nee cobble (pi. 15 figs 8-9). Frond broadly tongue- Lamarck, Australian record only; Sydney Harbour).— shaped, 5.5-6 cm wide at base, folded around on Whitelegge, 1889: 184 (off Green Point, Sydney sides, more so on right (viewed Harbour). from concave face), forming widely open compressed cup. Almost flat Hircinia (Sarcotragus) campana. — Lendenfeld, 1889b: 569, pi. 28 fig. 3 (part, nee Lamarck; old record). central part of frond tilted backwards, 12 cm in Ircinia campana caliculata. -de Laubenfels, 1948: 71 greatest height, along its axis. Greatest width,

(resume). across lateral folds, in upper third, 1 1 cm. Margin Ircinia caliculata. - Bergquist, 1980b: 466 (in revision, of frond has three round notches, one on left, two as good species). plus round hole on right. Thickness of frond 1.5

Figure 91. Ircinia caliculata. Perpendicular section in reflected light, showing armoured ectosome, canal system, and part of the skeleton, x8.3. 136 F. WIEDLNMAYER

wide, 0.3 to 2 mm. Dis- cm near base, about 1 cm near rim (at one point canals, 300-1000 /xm spaced only 4 mm). continuous subcortical crypts. Bottom of concavity juts foreward slightly. Main skeleton lax and irregular meshwork of Complex pedicel below this expands slightly thick, knotty, straggling fibres, mostly 100-250 /*m towards base, comprises stout, somewhat com- thick, with meshes around 1-2 mm wide. Com- pressed pillar on left, 17 x 22 mm thick in middle; monly little distinction of primaries and secon- more compressed twofold pillar on right, with daries, as almost all fibres contain much mixed round hole in centre, 7 mm wide in front, 13 mm detritus, but ascending fibres often more knotty in back, 38 mm on side, across hole. Arch separat- and lacunose. Fibres occasionally in loose fascicles. ing main pillars in front horseshoe-shaped, 15 mm Few thinner fibres, around 50 /im, with few or no wide and high. Space between pillars at back much foreign inclusions. higher, about 6 cm to where pillars merge with Numerous filaments (pi. 34 fig. 6) tangled, often inclined back of frond. Bases of pillars extended grouped in strands, 4 to 6.8 /*m thick; their heads into irregularly curved rhizomes, anastomosed at scarce, irregular, often angular, 9-14 ^m wide. Fila- back, clinging to cobble 8.5x4x4 cm. Massive ments almost always coated with brown, finely base, between upper concavity and arch between speckled pattern, similar to that noticed on inner pillars, penetrated obliquely from left by deep, fibres of Carteriospongia silicala. Length of rare almost terete concavity, now partly dissected. clear filament 540 /*m. Upper part of back of frond decorated by conspicuous rugo-reticulate pattern, with sharply out- Remarks. Of the two descriptions in Lendenfeld lined meshes mostly 5-10 mm wide, narrower (2-3 (1889b), the one under Hircinia campana fits the mm) just below rim. Pattern irregular below and present specimen better. The other one, under H. on outside of stronger lateral frond, comprising caliculata, mentions oscules 0.3 mm wide, "scat- shallow pits and meandering grooves about 7 mm tered over the outer surface of the cup-shaped wide. sponge." He possibly confused the outer with the Colour in life complex: dull greyish red in con- inner surface. However, the dry syntype of H. cavity (10 R 6-7/2), greyish orange on upper back caliculata from Illawarra, here figured for compar- and sides (2.5 YR 7/2-4), lighter, purer orange on ison (pi. 15 fig. 10) resembles the new specimen lower back, sides and pedicel (5 YR 8/8 to 7.5 YR sufficiently in external shape. 9/4). Choanosome was dark brownish orange (YR Several species of lrcinia with Australian records 6/8). Colours changed little in alcohol. Firmly rub- have been accepted as valid by Bergquist (1980b: bery, tough in life and in alcohol. Most of surface 466; see Wiedenmayer et al., in press, for further

(including rugae and depressions on back) regularly data). Of those common in southern Australia, /. papillate (conulose), like goose-skin or somewhat rubra Lendenfeld (common in New South Wales) denser. Pattern weaker in parts of concavity, much is dark red in life, commonly small to medium- of upper margin almost smooth. Conules around sized, cup-shaped or lamellate, but, in contrast to

1 mm wide and up to 1 mm high. /. caliculata, always thin-walled, soft, and has detri- Concavity riddled by oscules, more regularly tus only in the primary fibres. /. australis Lenden- arranged and smaller above and on left, often feld (with /. arbuscula Lendenfeld as junior clustered near bottom and in right fold. Their width synonym) is multilobed, each lobe with an apical 0.6-1.2 mm above/left, 0.2-0.5 mm below/right, oscule, the surface covered by dense, pointed con- interstices mostly 2-5 mm. ules; with little detritus, in primaries only. /.

With low magnification, fine rugo-reticulation irregularis (Polejaeff) (with synonym /. gigantea all over outer surface and peduncle, nervures occa- Lendenfeld) is massive, often cake-shaped, large, sionally forming indistinct radial pattern over dull orange to fawn in life, with dark brown oscules conule tips, fine ostia crowded in meshes. Exhalant clustering on top, and coarse, blunt conules in life, concave surface contains much smaller oscules and is internally very cavernous. It is common in (about 50 /itn wide) regularly scattered between northern Australia, New South Wales, and St Vin- larger ones. cent Gulf, South Australia (Bergquist and Skinner, Armoured ectosome 250-400 /*m thick (text-fig. 1982, pi. 2 fig. 1). 91), with packed spicule fragments dominating in lrcinia lendenfeld7 de Laubenfels (1948: 77; for outer half, coarse sand grains crowded below. Simi- /. irregularis Lendenfeld, nee Polejaeff, with syn- lar segregation in choanosome: spicule fragments onym /. jacksoniana Bergquist) has been recorded larger crowded around walls of canals, coarse sand only from New South Wales. It is massive-lobate, grains and few spicule fragments scattered inter- with blunt conules connected by ribs, "dirty light stitially. Choanosome riddled by terete, annulate brown" externally, "intensely orange-yellow" inter- SPONGES FROM BASS STRAIT 137

nally (fide Lendenfeld, 1889b: 579). The filaments a regular, almost perfectly rectangular meshwork are described as "very stout" (10 ^m thick). The in which the spaces between the fibres may be quite ectosome is described as armoured, with detritus large (up to 4 mm) and primary fibres are up to forming a banded reticulum, the oval membrana- 0.3 mm thick. The surface is always armoured, and ceous meshes being cribriporal. may be smooth, conulose, or thrown into ridges Ircinia halmiformis, as described by Lendenfeld and hollows. Shape commonly stalked, globose to (1889b: fig. 586, pi. 27 9) from the West Coast, fiabellate, but also tubular, lobose, or caliculate. has the shape of a folded lamella, with folds being Mucus production common, especially in species radial and dendritic. The surface seems to be pitted with widely spaced fibres. (Combined after Berg- (in the photograph of a wet specimen; Lendenfeld quist, 1980b: 468, 472.) described it as smooth). The sand grains are in Remarks. The definitions columns, but individually coated by spongin and given by Bergquist (1980b) for Thorecta and Thorectandra differ only connected by short, straight fibres. by degrees. External shape and mucus production Ircinia arenosa Lendenfeld (1889b: 583, pi. 36 are hardly useful in practice as diagnostic criteria. fig. 3) was described as regularly cup-shaped, often This is best exemplified by two of the species tall and pedunculate; with walls around 1 cm thick, described below, one new, and one misidentified surface smooth, except for irregular rugae on the (as being a haplosclerid) in Bergquist's revision. In outside of large specimens; colour in life brownish both, the dimensions of the fibre reticulation and pink; with sand in ectosome and all fibres, but not the rugo-reticulation of the surface are intermedi- interstitially; and with filaments 2-2.5 fim thick. ate and the two species could equally well be placed It was recorded by Lendenfeld from the West in cither genus. In one of the specimens of T. Coast, from Port Phillip Heads, and from Port choanoides, furthermore, fibre diameters generally Jackson (here in 20-50 m depth). fil the values given by Bergquist for Thorecta. Ircinia rugosa Lendenfeld (assigned to Psam- Production of slime is a variable trait even within mocinia by Bergquist, 1980b) is here regarded as a species (Thorecta choanoides, see below). De a nomen dubium, pending revision of the compo- Laubenfels' merger of the two genera is therefore site type series in the BNMH (unpublished upheld. Carter/Wilson material from Port Phillip Heads), designation and redescription of a lectotype. Thorecla choanoides (Bowerbank)

Thorecta Lendenfeld Plate 16 figures 1-3, plate 35 figures 1, 2

[Geelongiu\ Carter, 1885c: 306 (nomen oblitum).- Halispongia choanoides Bowerbank, 1872: 123, pi. 6 (F'remantle. WA). -de Laubenfels, 1948: 109, 114 (as syn- Bergquist, 1980b: 468, 470 (as synonym of Thorecta, dis- Thorecta cussion, type species redescribed). onym of bo/eta [recte boletus) (Lamarck)).

I' horectandra choanoides. — Lendenfeld, Thorecta Lendenfeld, 1888: 142. -Lendenfeld, 1889b: 1889b: 373, pi. 34 figs 1, 6-7 (old record). - Bergquist, 1980b: 464, 336. — de Laubenfels, 1936a: 25 (definition, invalid desig- 472, figs lOd, 14c (in review, as good species). nation of type species), —de Laubenfels, 1948: 107 (valid Stelospongus lams (Stelospongus flabelliformis var. designation, but misinterpretation of type species, errone- too). -Carter, 1885c: 306 (part: BMNH 1886.12.15.219 ous revision). — Bergquist, 1965: 128 (discussion).— only; Port Phillip Heads, Vic). Wiedenmayer, 1977a: 70 (erroneous synonymy, clarifi- Stelospongus flabelliformis. - Dendy, 1889a: 325, pis cation of type species, rnisidentification of West-Indian 30-33 (Port Phillip Bay). material). — Bergquist, 1980b: 468 (revision, review).

Antheroplax Lendenfeld, 1889a: 24 (definition as Material examined. Station KG 9, two specimens (F52050, subgenus of Phyllospongia).— Lendenfeld, 1889b: 168, F5205I). Paralectotype of Stelospongus latus Carter 171 (part: type species, Geelongia vusiformis, only, not (BMNH). Hypotype of Stelospongus Jiubellijoimis: 8 other included species). — de Laubenfels, 1948: 123 Dendy, 1889a (NMV). (designation of type species, but neotype designation stalked, invalid). Diagnosis. Often body globular, lobate, Thorectandra lendenfeld, 1889a: 41. -Lendenfeld, or thickly fiabellate. Surface more or less sculp- 1889b: 369. -de Laubenfels, 1936a: 26 (designation of tured, coarsely rugo-reticulate, vermiculate, brain- type species). — de Laubenfels, 1948: 107 (merged in like, or conulose. Oscules conspicuous, often Thorecta). — Bergquist, 1980b: 472 (retained, synonymy, elevated, single and apical on lobes, with smooth, definition, revision, invalid designation of type species). blunt or tabulate rims. Sand armour in ectosome Halispongia. -Bowerbank, 1872: 123 (nee Blainville). variable in thickness, thickest on ridges and oscu- Diagnosis. Thorectidae in which the primary fibres lar rims, then brittle; may be lacking in lower and are cored with a regular axial column of debris and depressed parts, ectosome then often deciduous the secondary fibres are clear. The skeleton forms after collecting. Firmly rubbery to spongy, stem 138 F. WIEDENMAYHR

in stem. F52051 more little compressible; often slimy. Grey with reddish firmer, barely compressible easily broken, stem to orange tinges on surface in life; in unarmoured spongy on top, with brittle skin depressions and choanosome bright yellow, with more compressible than in F52050. Surface of stem dark reddish grey fibres. Choanosome commonly in F52050 more finely conulose than on top, with dense, becoming viscous, flowing out after collect- conules aligned in anastomosing ridges. On top of ing in some specimens. Main skeleton typically F52051, fine rugo-reticulate meshwork superim- regular, but mesh size variable, tighter below sur- posed on coarse folds and grooves, with ridges and face ridges. pits both 0.5-1 mm wide, the latter corresponding to skeletal ostia. Description. F52050 better preserved (pi. 16 fig. 1), Better preserved surface of body of F52050 not more elegant, with terete, slightly bent stem gradu- as coarsely and thickly encrusted with sand as that ally expanding into main body, in size and shape of F52051 (where ectosome is about 1 mm thick). like fist or lion's paw, but conical below, somewhat Low magnification shows evenly speckled pattern, compressed, ending in irregularly fanning cluster like leopard skin, all over depressions, formed by of partly coalescent, slender conical processes with reticulation of fine ridges packed with sand, whit- apical oscules. Stem 6.5 cm high, 1 cm thick, with ish in reflected light, outlining sinuous, often sprawling, rhizomatous base 5 x 8.5 cm wide. vaguely petaloid, darker, slightly recessed inter- Greatest width of body, in upper third, 10.5 cm, stices, in which translucent ectosome, lightly depth 5 cm in middle, total height of specimen 22 armoured with spicule fragments and fine sand cm. grains, overlies vestibules. Dark inhalant fields F52051 poorly preserved, lower half largely about 100-300 (tin wide, fine ridges between them macerated, upper part squashed. Originally like a 50-70 /tm wide. Spaces subdivided by groups of tree, with stocky compressed stem 18-28 mm thick spicule fragments and single sand grains outlining in middle and 5 cm high, rhizomatous base 4 x ostia 50-100 /ini wide. Near primary (coarse) ridges 8 cm wide. Slightly compressed body 10 x 12 cm and around isolated conules, secondary ridges (of wide, distally vaguely divided into 5 broad, largely reticulum) wider, radial around some conule tips, concrescent lobes with apical oscules. Most of sur- with dark interstices smaller. face of body thrown into coarse rugo-reticulate to Concave depressions between primary ridges cerebroid pattern, with blunt ridges, 3-4 mm wide occasionally traversed and raised by underlying out- and deep, contorted pits of similar dimensions, or ermost secondary fibres of main skeleton. narrower meandering grooves. In lower, conical part of body of F52050, now Surface of F52050 more elaborate, also rugo- mostly mecerated, main skeleton very lax below reticulate, but with narrow, sharp ridges often primary depressions, with few secondaries and rare interrupted, polygonal depressions thus confluent. primary fibres in periphery, down about 5 mm. Ridges commonly 1-2 mm wide, up to 4 mm deep. Main skeleton below main ridges of surface much Some areas, particularly near top, have fewer denser, as in deeper choanosome. depressions and wide conulose areas. Some ridges On top of F52050, ectosome packed with mixed near top smooth, conspicuously sandy. Conules detritus, 400-500 /tm thick on primary ridges, below oscules generally 1-2 mm apart, frequently 100-150 fim over inhalant concavities. Choanosome macerated, with tips of primaries protruding. Oscu- dense, contains numerous globular microsymbionts lar chimneys 2-4 cm high, 1.5-2 cm wide at base, with granular surface, 12-17 /tm in diameter. All 1-1.5 cm wide at top, with collared oscules 0.5 to fibres yellow, stratified, primaries 80-160 /tm thick, 1 cm wide. regularly and thickly cored by mixed detritus. Few In F52051, free ends of osculiferous lobes 3-4 sand grains and spicule fragments reach surface, cm wide, with oscules 8-12 mm wide, surrounded occasionally bulging, but always enveloped by by wide, flat collars. F52050 in life light grey with spongin. Spacing of primaries in deeper choano- reddish to orange tinges (10 R 9/2 to 7.5 YR 7/2) some mostly 500-1500 Secondaries always on ridges and oscular collars, pale lemon yellow urn. clear, 30-100 /tm thick, occasionally branched, in depressions (5 Y 9/8). Sandy surface over most irregularly spaced 300-1000 /tm. of body of F52051 with similar greys, but darker; other areas, already partly macerated before preser- Remarks. The specimen of Stelospongus flabel- vation, showed dark reddish grey fibres, and bright liformis: Dendy, 1889a (F52215) (about half of the yellowish orange choanosome (7.5-10 YR 8/12). body cut off and now missing) was obviously Profuse slime of similar colour from this specimen thickly flabellate, and resembles the paralectotype after collecting, no conspicuous exudate in F52050. of Stelospongus latus Carter (here figured for com- Alcohol of F5205O stained brownish orange, that parison, pi. 16 figs 2-3). The primary rugo- brown-red. F52050 of F52051 deeper rubbery, reticulation of the surface on both faces is similar SPONGES FROM BASS STKAIT 139

to that in F52050, but the ridges are more discon- Diagnosis. Globose, softly spongy, slimy, pale tinuous, almost dendritic, with labyrinthic depres- orange-brown in life. Conspicuous oscule single, sions. This pattern becomes finer towards the edge apical, collared. Surface smooth to finely conulose of the body, radially oriented on one face, dissolved and rugo-reticulate. Ectosome thin, mostly lightly into aligned conules on the other. The edge itself, armoured to soft, largely deciduous, as for choano- bearing the oscules, is mostly truncate, speckled some. Main skeleton very regular, spacing of with depressed, sharply rimmed inhalant fields. The primaries chiefly 2 mm, all fibres pale. elevations on the edge, all the ridges on both faces, Description. Originally almost spherical, slightly and the pedicel are encrusted with distinctly coarser compressed on one side, now flattened by own sand than the depressions. The thickness of the weight. Lower end slightly drawn out into short ectosome is intermediate between those in the new neck, where torn from base of attachment. Single specimens. Spacing of primaries is in the general very conspicuous collared oscule on top. Greatest range 0.5-2.5 mm. width of specimen 8.5 cm, height 9 cm. Broken Of the valid species recognised in Thorectu and neck at bottom 1.5 x 2.5 cm. Oscule 1.5 cm wide Thorectandra by Bergquist (1980b) and myself at rim, collar 4-5 mm high. (Wiedenmayer et al., in press, with clarification of Deep brown in life (2.5 YR 4-5/4) with sharply Bergquist's confused revision), several are stalked outlined greenish-grey patches on one side. Much and distally massive or fiabellate, chiefly recorded light brown mucus emitted upon collecting, partly from southern Australia. macerated surface then pale orange-brown (5 YR Thorecla farlovi (Hyatt, 1877; with junior syn- 7/4-6). Softly spongy then as now. onym Stelospongus flabelliformis Carter, see Some smaller, flush oscules, 2-3 mm wide, scat- below, Remarks on Fasciospongia rimosa) is con- tered in lower part of specimen. About half of sur- sistently fiabellate, more or less symmetrical. Its face now partly or totally macerated. Elsewhere, salient diagnostic traits are the coarse rugo- as in life, three types of surface structures: reticulate to — vermiculate pattern on both faces, (1) smooth, (2) fine rugo-reticulation with irregu- the margin with prominent radial atria and distal larly square to polygonal meshes 0.5-1 mm wide, oscules, the pedicel commonly compressed in the and (3) rugo-reticulation combined with conules, plane of the frond, the scantily armoured, and the former occasionally feeble and discontinuous, mostly deciduous ectosome. the latter up to 2 mm high, 1-3 mm apart. Thorecta latus (Carter, 1885c, here restricted to At low magnification, type 1 merely better lectotype 1886.12.15.218; with junior syn- BMNH preservation of type 2, with lightly armoured outer Carter, 1886^, onyms Stelospongus cribrocrusta ectosome masking subjacent tangential reticulation varr. and tenia and Thorecla exemplum secunda of trabeculae packed with debris. Recessed inter- Lendenfeld) is chiefly pedunculate-pyriform, occa- stices in types 2 and 3 now mostly open, often with biconvex-flabellate, with sionally compressed, central round pore, due to contraction of dermal armour, smooth surface, finely clathratc ectosornal membrane. Cribripores occasionally preserved. specimens. It may which is more conspicuous in dry Densely armoured, lower reticulate ectosome prob- pesleonis externally (see resemble Coscinoderma ably discontinuous in life, large areas merely above). Thorecta boletus (Lamarck, with synonym covered by dermal membrane with scattered Lendenfeld) is thickly Thorectandra corticatus detritus. single apical club-shaped, truncate above, with Surface types 2 and 3 with trabeculae of ecto- ectosome oscule. Its surface with thick armoured some packed with ill-sorted mixed detritus, lithic rugo-reticulate on the is regularly and coarsely grains (up to 600 /*m wide) dominating, to depth Thorecta sides, and the main skeleton is coarse. 700 pim. In recessed interstices, ectosome 300 jtm figs 11a, b) typicus (Carter, see Bergquist, 1980b, thick, with crowded detritus better sorted, propor- frond of is symmetrically fiabellate, with elliptical tionately more spicule fragements, smaller lithic both faces even thickness, apical-marginal oscules, grains. Main skeleton very regular at low magnifi- sculp- with vague tuberculate and rugo-vermiculate cation, with primaries spaced 1.4-2.5 mm (2 mm conspicuously ture, armoured ectosome thick and prevailing), 110-200 fim thick, and clear secon- clathrate, coarse main skeleton. daries, 37-120 /mi thick, spaced mostly 1-4 mm. Primaries occasionally branch, thickly cored, rarely Thorecta glomerosus sp. nov. filled, with ill-sorted mixed detritus. Spicule frag- Plate 16 figures 4, 5 ments often disoriented, with one or both ends protruding, always enveloped by spongin. Fibre Holotype: NMV F52052, Station KG 7 (Tasmania, North contours thus commonly knotty or serrated. Secon- Point of East Cove, Deal Island, Kent Group; depth 25-30 branch and anastomose; m, rock face below algal forest). daries clear, frequently 140 F. WIFDFNMAYLiK

junctions with primaries commonly trumpet- laterally, free ends protruding only from taller lobe.

sliaped. All fibres pale. I leight 5 and 7 cm, greatest width of lobes 2.7 and

3.3 cm, their depths 1 and 1.5 cm. Width of whole Remarks. The new species is distinct from most specimen 5.7 cm. "carved" congeners C'Thoreetandra") by having a In life grey with brownish, greenish and purple finer, more regular and weaker surface pattern, a tinges (5-10 R 6/2 to 7.5 YR 7-8/2) on exposed trait shared with T. tuberculatus, from which it parts, beige to dull yellow ( 10 YR 8/6 to 5 YR 2/9) differs by ils large apical oscule, globose shape, in recessed parts. Choanosome brownish orange to relatively weakly armoured ectosomc, and the more peach-coloured. Firmly rubbery. Few round, flush regular reticulation of pale fibres. oscules scattered on top. As in '/'. glomerosus, sur- Thorecta tuberculatus (Carter) face may be smooth, papillate (conulose), or finely rugo-reticulate to unaided eye. Latter surface type Plate 16 figures 6, 7, 10, plate 35 figures 3-5 only locally on top of larger lobes in F5205.3, with Stelospongus? tuberculatus Carter, 1885c: 306 (Port primary pattern of light grey ridges, 0.5 mm wide,

Phillip Heads, Vic), I nubenlels, 1948: 1 17 (as syn- -de connecting papillae (conules) mostly 1 mm high, onym of Thorecta murrayi (Polejaeff)). 1-2.5 mm apart. Secondary, finer rugo-reticulation Thorecta tuberculatus, Lendenfeld, 1889b; 36V, pi, superimposed, with little relief. Numerous ostia 33 fig. 12 pedesenption of hololype). BergqtliSt, l9H()h: commonly crowded in square to polygonal fields 470 (in review, "unrecognisable except as a haploselerid in meshes of secondary reticulum. Individual ostia sponge".) often separated by thin bridges of single sand grains Material examined. Station K(i 5, one specimen (F52053); or spicules, forming tertiary ectosomal reticulum; station K(i 6, one specimen (F52054); station K(i 7, one but not consistenly developed, with many inhalant specimen (F52055). Hololype of Steldspongusl tubercula- fields confluent, petaloid. Latter pattern (secondary tus (BMNII). reticulation only) on almost whole surface of Diagnosis. Irregularly lobate, often with short, F52053, where armoured ectosome best developed irregular pedicel(s). In life, grey with brownish, and continuous. In other specimens, parts of sur- greenish and purple tinges above; beige to dull face now macerated, without armoured ectosome yellow in depressions and below. Choanosome in life, tips of primary fibres protruding from many brownish orange to peach (fibres orange-brown). conule tips. Primary ridges occasionally formed by Firmly rubbery, slimy. Surface smooth to finely alignment of conules. Areas with large, round to papillate, or rugo-reliculate. Few small osculcs on square or contort cd-elongate skeletal ostia (inter- top. Armoured ectosomc variably thick, may be slices of secondary ectosomal reticulum) alternate absent below, thickest on lop (up to I mm), here with clusters of much finer skeletal ostia. with debris often forming compound reticulum, Top of lobe in F52053 has cortex 0.7-1 mm thick less with interstitial ostia. Main skeleton regular for containing almost exclusively lit hie grains (rare spic- genus; primaries knotty, straggling, spaced 1.5 mm. ule fragments and shelly grains). Choanosome dense. Crowded globular microsymbionts confined Description. Irregularly lobate. In F52053 (pi. 16 to linings of canals, inters! ices between sand grains fig. 10), 3 larger aligned lobes and smaller inclined in cortex, and, most conspicuously, just below lobe laterally concrescent, fifth lobe, jutting from cortex. others, free above its constricted base. Smallest, Fibres orange-brown (transmitted reflected concrescent lobeclavate, others fungoid, bulbous and light), often crooked to meandering. Primaries on top. Two short pedicels, one below free lobe, 130-185 jixa thick, with mixed detritus in core of the other, composite and compressed, below lar- variable thickness, occasionally displaced to one gest (wo lobes. Specimen 4 cm high, 3.5 x 6.5 cm side; irregularly knotty with wide. bulging grains and protruding spicule fragments. Average spacing of F52054 comprises (I) thick lamella, with irregu- primaries 1.5 mm. Secondaries clear, 60-80 /tin lar folds inside, 4 cm high, 5.5 cm wide; (2) com- thick, spaced 1 mm. pressed lobe, 5 cm high, 2 x .3.5 cm wide on top, 1.5 cm at base; (3) erect digitate process, 1.5 cm Remarks. Caeospons>ia murrayi Polejaeff ( 1 884b: high, 5 mm wide; all connected proximally by thin 57, pi. 4 fig. 3, pi. 6 fig. 8) shows considerable base. resemblance with the species described above, exter- F52055 comprises two stalked lobes with intert- nally, and might be regarded as a senior synonym wined and concrescent pedicels. Both lobes com- of Thorecta tuberculatus (as was de LaubenleK' pressed in same plane, with digitate portions opinion). Bergquist (1980b: 470) treated C. mur- arranged in candelabra-fashion but largely fused rayi as a good species of Thorecta, but unfor- -

SPONGES FROM BASS STRAIT 141

tunately the specimen figured by her (1980b, fig. Stelospongos as a nomen nudum is incorrect: the 10a) as such is obviously not Polejaeff s holotype genus name is available with the first subsequently (BMNH 1885.8.8.27). The register number quoted included species (in Hyatt, 1877; see ICZN, Arti- by Bergquist actually belongs to the holotype of cles 1 lc(i), 12, 69a). De Laubenfels (1948) used the Ciocalypta penicillus aciculata Carter (1885c), ludicrous stratagem for Stelospongos, Stelospon- which, however is not the figured specimen. The gia and Stelospongus, repeated ad absurd um for latter is presently unidentified. The distinctly larger all genera he wished to sink, of designating the size of the oscules in Polejaeff s figure (p. 4 fig. same specimen as neotype of the respective type 3), in comparison to the holotype of T. tubercular- species, even though original primary types for tus and to the new specimens could be dismissed most, if not all, species concerned were extant. Such as insignificant, but the fine and thorough tertiary designations are, of course all invalid (ICZN Article fibre-reticulation described by Polejaeff make the 75). His subsequent designations of type species for identity of the two species very unlikely. the two unjustified emendations, objective syno-

Bergquist 's misidentification of T. tuberculatus nyms of Stelospongos, are also invalid. All share (as an unrecognisable haplosclerid) can only be the same type species, Stelospongos friabilis explained with another confusion of specimens, as Schmidt (1870).

I cannot imagine that examination of Carter's holo- Stelospongos is a nomen dubium, as Bergquist type (here figured, pi. 16 figs 6, 7) could lead to (1980b: 474) pointed out, since its type species is such a conclusion. unrecognisable. Should any type material of the latter surface, or fresh identifiable material permit Fasciospongia Burton to synonymise Stelospongos with Fasciospongia, then the senior synonym would be a nomen obli- Fasciospongia Burton, 1934a: 602. — de Laubenfels, tum and would have to be suppressed in favour of 1948: 118 (in part).-Vaceld, 1959: 93 (definition, discussion, revision for Mediterranean). — Bergquist, 1965: Fasciospongia (ICZN Article 79c). 135 (discussion). — Van Soest, 1978: 43 (definition, revi- 1980b: 472 sion for the West-Indian Region). — Bergquist, Fasciospongia rimosa (Lamarck) (definition, discussion, revision).

•1 Stelospongos Schmidt, 1870: 29. -Hyatt, 1877: 528 Plate 16 figure 11, text-figure 92 (pan). -Burton, 1934a: 602 (footnote). -de Laubenfels, Spongia rimosa Lamarck, 1814(1813-1814): 377 with 1948: 133 (subsequent designation of type species). - varieties columnaris and subclavata; Australian Seas). Bergquist, 1980b: 474 (discussion, dropped as Stelospongia rimosa. —Lendenfeld', 1889b: 497, pi. 26, unrecognisable). fig. 5 (no locality). [Stelospongia] Schulze, 1879a: 613 (nomen vanum; Tlwrecta rimosa. -Topsenl, 1930a: pi. 3, fig. 5 (2 part).-Lendenfeld, 1888: 163 (part).-Lendenfeld, 29, syntypes redescribed, type of var. subclavata figured). 1889b: 468 (part). -de Laubenfels, 1948: 132 (invalid type Fasciospongia rimosa. —Bergquist, 1980b: 474 (in designation). -Bergquist, 1980b: 474 (revision). review). [Stelospongus\ Ridley, 1884a: 383 (nomen vanum; Stelospongia canalis Lendenfeld, 1888: 164 (south coast neque sensu Ridley). -Carter, 1885c: 303 (part). -de [= ?Tasmania in 1889b]; Sydney Harbour).— Laubenfels, 1948: 133 (invalid type designation). Lendenfeld, 1889b: 495, pi. 24, fig. 10; pi. 29, figs 2, 4,

Diagnosis. "Thorectidae in which the primary fibres 5, 7; pi. 31, fig. 1; pi. 32, fig. 6., (old records?). are cored with detritus and arranged in stout fas- Whitelegge, 1901: 114, pi. 15 fig. 31 (2 syntypes in AM cicles which curve in plumose fashion towards the redescribed; coast of NSW, new record). — Whitelegge, 212 (schizosyntype BMNH). -Bergquist, 1980b: sponge surface. The secondary skeleton is clear of 1902a: ex 472, 474, fig. 14b (as synonym of F. rimosa, lectotype, detritus, and the mesh is branching but regular. Lendenfeld, 1889b, pi. 24 fig. 10, refigured). Sponges belonging to Fasciospongia emphasise a Euryspongia canalis. — Burton, 1934a: 602 (discussion, tubular, cup or fan shape in which globular, redescription of skeleton; Great Barrier Reef; Perth [as either centrally or marked exhalant canals run Swan River|, WA). superficially. In the living sponge the surface has Stelospongia jiubellum Lendenfeld, 1889a: 53 (in key, a shiny, skin-like appearance produced by the con- part). -Lendenfeld, 1889b: 501, pi. 31 fig. 12, pi. 32 fig. siderable collagenous deposition in the ectosomal 8, pi. 39 fig. 6 (part: not Carter-Wilson material (part?); lllawarra, NSW; Port Phillip Heads, region. Conules are multiple because of the fibre Northern Territory; Vic. [Perth, WA, as Swan River, unpublished)). fascicles, and often truncate rather than pointed. I asciospongiaJiabellian. — Bergquisl, 1980b: 472, 474, at the There is only slight differential pigmentation fig. 14a (lectotype, as holotype, = Lendenfeld, 1889b, surface, more frequently the colour is uniform pi. 39 fig. 6), fig. 16d (in review, as good species). sponge." (Bergquist, 1980b: 472.) throughout the Stelospongia reticulata Lendenfeld, 1889a: 53 (in -Lendenfeld, 1889b: 510, pi. 24 fig. pi. 32 fig. Remarks. Burton's (1934a: 602) qualification of key). 1, 142 F. WIEDENMAYER

<*0^m

Figure 92. Fasciospongia rimosa. Perpendicular section showing rugose ectosome (above), canals and skeleton, x 8.2.

5 (Northern Territory). — Bergquist, 1980b: 474 (in review, cous as grooves after collecting and preservation, as synonym of Fasciospongia rimosa). when fleshy dermis collapses. Surface thoroughly Stelospongia flabelliformis. — Whitelegge, 1901: 115 papillate and vaguely rugose in life, distinctly conu- (nee Carter; Lake Illawarra, NSW). lose in preserved specimens. Choanosome dense. Material examined. Station KG 2, one specimen (F52056). Main skeleton typical, primaries and fascicles often Type specimens of Stelospongia canalis, Stelospongia oblique. Fibres stiff, brown. flabellum, Stelospongia reticulata (AM, BMNH). Hypo- types of Stelospongia rimosa: Lendenfeld, 1889b, Description. Stout stalk abruptly expands into Stelospongia canalis: Whitelegge, 1901, Slelospongia thick, complex, folded lamella above. Stalk 6-7 cm flubelliformis: Whitelegge, 1901 (AM, BMNH). Type long, slightly compressed, 1.8 X 2 cm thick, specimens of Stelospongus flabelliformis Carter, 1885c expanding below into base 4 x 4.5 cm wide, (BMNH, for comparison). Hypotypes of Thorecla farlovi: inclined at 45°. Viewing Lendenfeld, 1889b (BMNH, for comparison). specimen with inclined base backwards, main frond expands laterally from Diagnosis. Stalked, or with short, root-like pedicel; top of stalk, about evenly to mid-height, to width main body irregular, lobate, concrescent-columnar 10.5 cm, then higher on left side (11 cm above its or -flabellate. Top firmly rubbery, stalk barely base). Upper margin evenly rounded on the left, compressible. Greyish red-brown on top, cream sagging and sinuous on the right. Frond longitu- below in life; dull brown in alcohol. Oscules 1-8 dinally folded, extends into one secondary frond mm wide, apical on columns, on upper margins of each on front and back, both attached to main fronds. Terete ascending atria at surface conspi- frond through most of its length on both sides of SPONGES FROM BASS STRAIT 143

its axis. Both directed outwards at 45° to plane of with the columnar to subclavate habit of Lamarck's main frond. Secondary frond in front much larger, specimens is demonstrated by F52056, but even projecting up to 5 cm, its inner portion as thick better by the specimens described by Whitelegge as main frond, 1-1.5 cm, its outer half expanding (1901, as Slelospongia canalis). The figured speci- into fluted, somewhat contorted club-shaped struc- men, in particular, shows a main lamella and short ture up to 4 cm wide. Rudimentary lateral frond peduncle very similar to the lectotype of Slelospon- near left margin, on front of main frond, project- gia flubellum, but, in addition, shorter secondary ing up to 1 cm. Secondary frond along back fronds attached vertically and partly folded, like projects up to 2 cm, 1 cm thick, ends 2 cm below in F52056. It should also be noted that two of the upper rim of main frond. Total height of specimen eight paralectotypes of Slelospongia flubellum 19 cm. (BMNH unregistered, dry) are of the rimosa-type. In life greyish red-brown on top (10 R 7-8/4) The erroneous inclusion by Lendenfeld (1889b) grading into cream or off-white below (N 9- 10 (YR of Stelospongus flabellifonnis Carter as synonym

1)); now uniform dull brown (7.5 YR 4-5/4). of his Slelospongia flabellum, which confused Firmly rubbery on top, barely compressible along Whitelegge (1901: 115), needs to be clarified. stalk. Some clear slimy exudate upon collecting. Stelospongus flabellifonnis is a junior synonym of Oscules 1-8 mm wide chiefly on upper edges of Tfioreau farlovi (Hyatt, 1877) on the basis of five all fronds. Many terete atria just below surface, of the six syntypes (one wet) accepted by Wieden- now wholly or partially open, some still roofed mayer et al. (in press). Carter (1885c: 305) men- completely by dermal membrane, as all were in life. tioned only three specimens in his description. In These more or less pronounced grooves in surface his Manuscript Catalogue of Type Specimensfrom generally arranged longitudinally, but meandering, the J.B. Wilson Collection (unpublished, BMNH; occasionally branched downwards. copy in NMV Sponge Archive), however, 7 dry Except these marginal atria, whole surface finely specimens are listed under Stelospongus flabel- papillate and variably rugose in life. Through lifonnis. All the specimens in Carter's Catalogue shrinkage of ectosome, surface now more distinctly are registered (BMNH 1886.12.15.1-508). Apart conulose. Conulcs 0.5-1 mm high, mostly 1-2 mm from inconsistencies in numbers of types and their apart, irregularly distributed, frequently in clusters preservation, also in other species, there are occa- or aligned in ridges. Ridges connecting neighbour- sional inconsistencies between published names, ing conules form vague rugo-reticulation or labyrin- names on Carter's autograph labels, and entries in thic pattern in some areas. Some intervening (he BMNH Register. This may be partly due to depressions bear single or branched groove at negligence by Carter, associated with piecemeal bottom, and all depressions contain irregularly publication of rather hastily written descriptions, clustered, indistinct ostia. Some polygonal depres- and, as with Lendenfeld. changes of names on this sions irregularly open below, like invaginations, occasion, without corresponding corrections on possibly additional oscules. In transverse section labels. But many such discrepancies may have been (text-fig. 92) ectosome about 200 /an thick; dense deliberate: some of Carter's labels bear, after the choanosome traversed by canals mostly 100-1000 species name, page references to publications, while that Carter, in jtm wide, and conspicuous irregular reticulation of others do not. 1 suspect, therefore, indicated in descrip- stiff brown fibres. Primaries and their fascicles fre- the case of larger type series, quently oblique to surface. Primaries 150-200 /an tions only the number of those specimens examined thick, secondaries 30- 120 /mi. Meshsizcs commonly with more care (microscopically). Under "Obs.", (1885c: 305) mentioned in passing the 300-800 urn. Fascicles around I mm wide, irregu- Carter his specimens with the photograph of larly spaced 1-5 mm. Primaries markedly strati- similarity of farlovi in Hyatt (1877, pi. 17 fig. 14). fied, with outer layers often wavy, and thin, Spongel'ia

sufficient to identify ,S'. straggling core of mixed detritus, including spic- This photograph 1 judge it, together with the specimens ule fragments, comparatively fine and well sorted. flubelliformis with correctly identified as Thorecta farlovi by Lenden- pi. 23 fig. refigured by Remarks. The synonymy given above departs from feld (1889b, especially 7, Bergquist, 1980b, fig. lid, with the erroneous cap- that suggested by Bergquist (1980b: 474) in includ- Taonura flabellifonnis). The lectotype of ing Slelospongia flabellum. Though Bergquist did tion Slelospongia flabellum (Bergquist, 1980b, fig. 14a) not elaborate on this, the distinction, judging from the only specimen ot Carter/Wilson in Lenden- the descriptions, and from my cursory, macro- is type series. It is labelled by Carter "'ISponge- scopic examination of Carter's and Lendenfeld's feld's Farlovii Hyatt p. 536, pi. xvii fig. 14, Australia." specimens, relies only on external shape. That the lia It can therefore be surmised that this is one of the flabellate habit may intergrade in some specimens, -

144 F. WIEDENMAYER

seven dry syntypes of Stelospongus flabelliformis Spongelia pallescens Schmidt, 1862: 30, pi. 3 fig. 8 in Carter's Manuscript Catalogue, and that Len- (Sibenik, Yugoslavia; lectotype described and figured by see below). -Schmidt, 1864: 28 (part?, denfeld (1889b) was for the least part right in Burton, 1934a, merged with S. incrustans, which was regarded as a good including S. flabelliformis as synonym of S. species by Vacelet, 1959, and Puliizer-Finali and Pron- flabellum. zato, 1977, 1980). -Burton, 1934a: 591 (part, as synonym of Dysidea fragilis; BMNH 1867.7.26.9 designated Dysideidac Gray, 1867 lectotype [as holotype], described and figured text-fig. 30, Dysidea Johnston pi. 2 fig. 4, pi. 2 fig. 3, Naples). nee Spongelia pallescens. — Schulze, 1878b: 138, pi. 5 Dysidea Johnston, 1842: 251. -Burton, 1934a: 582 figs 1-4; pi. 6 figs 2-3, 5-7 (with sspp. fragilis, elaslica (synonymy with nomina vana and nomina nulla; defini- and varr.; fide Vacelet, 1959: 67).-Polejaeff, 1884b: 42, tion, revision species). — Vacelet, 1959: discussion, of type pi. 3 fig. 1 (Bahia). 66 (synonymy, definition, discussion, revision of Mediter- Dysidea pallescens. — de Laubenfels, 1935: 327 (figured ranean species). — Bergquist, 1965: 142 (discussion).— in de Laubenfels, 1948; Puerto Gatera, Mindoro, Philip- Wiedenmayer, 1977a: 72 (synonymy with further refer- pines). -Boury-Esnault, 1971: 338. ences, definition, discussion of West-Indian species).— Dysidea avara vai. pallescens. — Vacelet, 1959: 69 (Gulf Van Soest, 1978: 51 (definition, discussion, revision of of Marseille).— Riitzler, 1965: 42 (synonymy, descrip- West-Indian species). — Bergquist, 1980b: 480 (synonymy tion, ecology). in part: not SaKOCornea; definition, discussion, revision). Spongelia Nardo, 1847a: 115 (nomen nudum). Material examined. Station KG 4, one specimen (F52057); Schmidt, 1862: 28.- Wiedenmayer, 1977a: 72 (further station KG 6, one specimen (F52058). references). Diagnosis. Commonly small, irregularly lobate, Aulena Lendenfeld, 18851': 308 (part: A. villosu and sprawling, occasionally digitate. In life, mauve, A. flabellum; not A, nigra, a Hippospongia). — de often mixed with grey, or dark purple. In alcohol, Laubenfels, 1948: 33 (part: A. villosa only). violet, brownish nee Aulena sensu Lendenfeld, 1889b: 90 (= or beige. Softly spongy, limp, Echinoclathria). mostly easily torn, some specimens more resistant. Halmopsis Lendenfeld, 1885f: 320.-Bergquist, 1980b: Surface coarsely conulose; conules 1-6 mm high, 454, 480 (as synonym of Dysidea). with similar, irregular spacing; tips may be blunt, Haastia Lendenfeld, 1888: 204. -Bergquist, 1980b: 482 pointed, irregular, composite, crested. Oscules (discussion, of Dysidea). synonym small, irregularly scattered, often in depressions.

Diagnosis. "Dysideidae in which all fibres are filled Choanosome cavernous. Fibrous skeleton lax. with detritus." (Bergquist, 1980b: 472.) irregular. Fibres irregular, may be compressed, fenestrate, commonly loaded with detritus, but with Dysidea avara (Schmidt) spongin still visible. Detritus also interstitial, but not in dermis, which contains a fairly conspicuous, Plate 16 figure 9, plate 35 figure 6, whitish collagenous reticulum. text-figures 93-95 Description. Sprawling depressed lobes, indistinctly Selected synonymy. separated by sinuous incisions, bearing some Spongelia avara Schmidt, 1862: 29, pi. 3 fig. 6 (Zadar moderately deep invaginations above. Whole sur- and Sibenik, Yugoslavia). — Schulze, 1878b: 127, pi. 6 figs face coarsely conulose in life, now, after shrink- 1, 4; pi. 8 figs 1-7, 13-14 (anatomy) (Hvar Island, Yugos- age and partial maceration, frilly, with contorted lavia). -Lendenfeld, 1889b: 667, pi. 43 figs 3-4, 6 (Port conules or lacinules connected by discontinuous Phillip Heads, Vic). ridges, like miniature cockscombs, Dysidea avara. — de Laubenfels, 1948: 142 (part; which frequently intersect, resume, discussion), pi. 23 fig. 43 (specimen from Philip- forming irregular reticulation pines described in 1935 as D. pallescens). — de Lauben- with deep, angular depressions. In life mauve on fels, 1950a: 9 (Hawaii). -de Laubenfels, 1954a: 36, surface, paler on underside, with whitish ridges, text-fig. 18 (Micronesia). -Vacelet, 1959: 68, pi. 1 fig. dull beige internally; now faded to yellowish beige 7 (description, discussion, notes on ecology; off Mar- (10 YR 8-9/4). Softly spongy, limp, easily torn. seille). — Riitzler, 1965: 42 (further references, description, In F52057 (pi. 16 fig. 9, text-fig. 93) conules ecology, dislribution; Northern Adriatic Sea). — Boury- often clustered, with expanded and blunt, compo- Esnault, 1971: 337 (notes on ecology, reproduction, fur- site tips (0.3-1 wide) roughened ther references; western Mediterranean). — Boury-Esnault, mm by protruding 1973a: 288 (Brazil, S of Porto Alegre).- Pulitzer -Finali sand grains. Individual conules 1-3 mm high, and Pronzato, 1977: 87 (Bay of Naples, locality data very irregularly spaced; width and depth of inter- only). — Pulitzer-Finali and Pronzato, 1980: 131 (western stitial depressions about 1-5 mm. Relief of F52058 Mediterranean, locality data, remarks). — Rubio et al., similar, but conules more slender, with fine, taper- 1981: 12, text-fig. 2 (description, distribution, ecology). ing tips. Round oscules, 0.5-1 mm wide, irregu- SPONGES FROM BASS STRAIT

Figures 93-95. Dysidea avara. F52057, details in reflected light, x4,l. Figure 93. View of well preserved portion of surface. Figure 94. Mostly macerated peripheral portion showing tendency of fibres to form fenestrated lamellae. Figure 95. Perpendicular section showing cavernous interior.

lady scattered at bottoms of invaginations and in grains attain 0.5 mm in size. Most fibres have some of shallower depressions between conules. irregularly knotty contours, and many thinner 2-4 Very fine ostia barely visible between vague white fibres abruptly expand to times their diameter. nervures of ectosome in depressions. Peripheral stratification of pale spongin envelop- Choanosome coarsely and thoroughly cavernous ing grains frequently visible. A portion of fibre 700 180 to 90 /*m, free of detri- (text-fig. 95), with many round canals 2-3 mm /im long, tapering from wide, interstices sometimes as narrow as 200 /im. tus, with layered bark and thick pith (140-60 fim) Matrix dense, yellowish-brown in transmitted light, transversely wrinkled. In F52057, fibres often form fenestrate sheets, to 1.3 with elongate, frequently somewhat contorted, sac- tight fascicles and up mm shaped choanocyte chambers often crowded and wide (text-fig. 94). Much detritus also interstitially, aligned. scattered and clustered. thin fibres commonly 60 thick. Fibrous skeleton in F52057 lax and very irregu- In F52058, fim detritus in all fibres generally finer and better lar. Fibres 100-500 /mi thick, commonly almost Mixed but larger lithic or skeletal packed with mixed and ill-sorted detritus. Sand sorted than in F52057, -

]46 F. WIEDENMAYER

grains occur interstitially. Reticulation more regu- Balls Head, Sydney Harbour, probably Lendenfeld's lar than in F52057, without fascicles and fenestrate record). — ?Burton, 1930a: 510, pi. 2 fig. 3 (synonymy in sheets. part: nee Aplysina naevus, nee A. cruor). — ?Burton, 1934a: 595 (record only, Great Barrier Reef). — ?Burton, Remarks. Vacelet (1959) and Rutzler (1965) distin- 1936a: 142 (record only; Oude Kraal, Muizenberg, South guished a typical variety, with coarser conules and Africa). -Bergquist, 1967: 161 (Hawaii). -Levi, 1952:37 deeper purple conserved in alcohol, from a var. (West Africa). -Borojevic et al., 1968: 29 [A. sulfurea merged, local distribution, pallescens with slightly finer conules (see pi. 2 figs ecology, further references; Roscoff). -Pulitzer-Finali and Pronzato, 1977: 98 (local 3, 4, in Burton, 1934a), thinner ectosome and a distribution, A. sulfurea merged, discussion of colour, lighter purple streaked with white, which disappears nomenclature; Bay of Naples). - Pulitzer-Finali and Pron- in alcohol. Rutzler also found that the choanocyte zato, 1980: 152 (Ligurian Sea). chambers in var. pallescens approximate the spher- ? Aplysilla sulfurea Schulze, 1978a: 405, pi. 23 figs 15, ical type. Boury-Esnault (1971: 338) regarded D. 18, 20-27; pi. 24 figs 28-30 (Adriatic). -de Laubenfels pallescens as a good species. The new specimens, 1948: 163 (resume, further reference). — Levi, 1952: 37 with size and density of conules as in the lectotype (West Africa). -de Laubenfels, 1954a: 47 text-fig. 26

of Spongelia pallescens, corresponds with var. (Micronesia). -Vacelet, 1959: 62, pi. 1 figs 1, 2; (with A. rosea pallescens sensu Vacelet also in other external traits. and A. glacialis as synonyms, priority disregarded; discussion of But the almost spherical shape of choanocyte cham- colour; description, ecology, further references).- Rutzler, 1965:41 (description, ecology, bers is not developed. It remains to be seen whether further references). - Bergquist, 1967: 162 (Hawaii).— the equivalent of the typical variety also occurs in Boury-Esnault, 1971: 333~ (purple specimens included; Australian waters. local distribution, ecology, further references). — Vacelet and Vasseur, 1971: 115 (Tulear Reefs, Madagascar). Order Dendroceratida Minchin, 1900 Vacelet etal., 1976: 107 (Tulear Reefs). -Van Soest, 1978: Darwinellidae Merejkowsky, 1879 80 (references to West Indian records). 1 Aplysilla su/phurea. -Lendenfeld, 1889b: 707 (Svdney Synonym: Aplysillidae Lendenfeld, 1883. Harbour). -Thiele, 1905: 488, figs 112, 114 (Strait of Magellan). -Koltun, 1964: 111 (1966a: 112) (further refer- Aplysilla Schulze ences, diagnosis). -Bergquist, 1980b: 484, figs 2d, 20c, d (in review). - Rubio etal., 1981: 40, text-fig. 12 (table Aplysilla Schuhe, 1878a: 404. -Lendenfeld, 188%: 698 of distribution, ecology, further references, western (part). -Topsent, 1905a: CXC (definition in Darwinelli- Mediterranean). -Boury-Esnault and van Beveren, 1982: dae, designation of A. rosea (Barrois) as type species, -de 120 (part: encrusting specimen only, not pi. 20 fig. 78 = Laubenfels, 1936a: 32 (invalid designation of type spe- Dendrilla anlarctica Topsent; Kerguelen). cies). -de Laubenfels, 1948: 163 (part; wrong type spe- 1 Dysideopsis su/phurea Lendenfeld, 1888: 156 (Sydney cies).— Vacelet, 1959: 62 (definition, discussion, revision, Harbour, NSW). - Whitelegge, 1889: 184 (off Shark wrong type species, A. rosea and A. glacialis merged in Island, Sydney Harbour, probably Lendenfeld's A. sulfurca). - Van Soest, 1978: 80 (West Indian records, record). -Bergquist, 1980b: 464 (in review, as synonym discussion). — Bergquist, 1980b: 484 (definition, discus- of Aplysilla sulfurea). sion, revision; A. rosea, A. sulfurea and A. glacialis as distinct species). Material examined. Station KG 4, one specimen (F52061). Simp/icella Merejkowsky, 1878: 259. — Bergquist, 1980b: 484 (as synonym of Aplysilla). Diagnosis. Encrusting, 2-6 mm thick, commonly small, but may cover large areas. Limp, easily Diagnosis. "[Darwinellidae] in which the sponge torn. In life pink, (bright or pale yellow?), occasionally body is always encrusting and the fibres are always dark purple; chiefly dull red clear of detritus and almost unbranched." (Berg- or brownish in alcohol. Surface quist, 1980b: 484.) commonly conulose or bristly, rarely smooth or wrinkled, chiefly lipostomous. Choano- Aplysilla rosea (Barrois) some dense, fleshy. Fibres mostly single, up to 6 high, with basal plate, pith Plate 16 figure 8 mm with timble-shaped accretions, longitudinally striate bark; generally- Selected synonymy. tapering to point in conule axis or above surface, Verongia rosea Barrois. 1876: 57 (Roscoff, English occasionally simply branched. Channel). -de Laubenfels, 1948: 167 (as synonym of Aplysilla glacialis). Description. Aplysilla rosea Schulze, 1878a: 416 (Trieste, Adriatic Crust, 2-4 mm thick, with angular Sea). - ?Lendenfeld, 1888: 26 (part: nee Ap/ysina criwr, outline, 1.5-2 X 3.5 cm wide, deep lateral incision nee A. naevus; Sydney Harbour, 1 hypotype with slide in middle. Pink in life (values not recorded), now- in East Berlin, ZMB 2306).-?Lendenfeld, 1889b: 708, deep red-brown (10 R 4/6-8) with few paler spots pi. 44 fig. 2 (old record). -?Whitelegge, 1889: 180 (off (10 R 6/6). Limp, easily torn. White nudibranch -

SPONGES FROM BASS STRAIT 147 with elongate orange dots grazing on specimen of the extremely variable colour of Pseudoceratina before collecting. crassa in life (Wiedenmayer, 1977a: 75, as Surface rough in one half, irregularly conulose Aiolochroia), though this affects only the ecto- to bristly. Other half rugose with a large smooth some, and the genus is now placed in the order area. Isolated dendritic fibres mostly upright in Verongida. rough half, tips protruding from surface here and there. In other half most of fibres uprooted, with Darwinella Mullet basal discs partly protruding, branches recumbent [Darwinia] Schultze, 1865: 7 (nee Darwinia Bate, 1857). at or just below upper surface. Corresponding con- Darwinella M Ciller, 1865: 344 (nomen novum for Dar- spicuous difference in conformation of lower sur- winia Schullze). — Lcndenfeld, 1889b: 673 (2 species face: (1) Lacunose below rough upper surface, with included). - Dendy, 1905: 203 (definition, discussion).— about half of surface flush, with open canals, Topsent, 1905a: CLXX1II (resume for family, review of adhering to substrate in life; few fibres recumbent, genus with descriptions of 9 species, 4 of which are particularly in glabrous depressions. (2) Undulat- new). -de Laubenfcls, 1948: 168 (part: D. muelleri only, excessive Levi, 1952: 38 (brief review ing, with glabrous surface, fibres all uprooted and with synonymy). — with key). Vacelet, 1959: 63 (resume of Levi, 1952, for recumbent, except small area with large basal spon- — Mediterranean records). — Bergquist, 1961c: 207 (discus- gin plate supporting 2-3 fibres, slightly tilted. sion, new species). -Pronzato, 1975: 5 (resume, review Upper surface mostly glabrous, lipostomous, and revision, with reproductions of original figures, notes clustered grains at or contains scattered and sand on distribution with map, key). — Van Soest, 1978: 79 just below surface. (resume, discussion of West Indian records). — Bergquist Fibres conform to descriptions and figures in 1980b: 486 (synonymy, definition, discussion, revision on literature. Most fibres isolated, including basal the basis of Lcndenfeld, 1889b, only). plate, straggling to angular, branched once or Diagnosis. "[Darwinellidae] in which the slightly twice. Single basal plates about 2-3 mm wide. ramified dendritic fibre skeleton is supplemented Fibres 350-400 thick proximally, longitudinally ^m by diactinal, triactinal, or quadriradiate spongin striate on outside, with pith third to half of spicules. There is no sand in the fibres. The sponges characteristic convex accretion- diameter, showing are encrusting or massive to lobate and [commonly] almost to point, with ary interruptions. Fibres taper small." (Bergquist, 1980b: 486.) bark almost disappearing, commonly reaching length of 7 mm. Darwinella australiensis Carter

(1980b: 484), in keeping sul- Remarks. Bergquist A Plate 16 figure 12, plate 17 figures 1, 2, furea and A. rosea distinct, cited only Vacelet plate 36 figure 1 (1959) as disagreeing with her on this point. Her is a stable specific statement: "It appears that colour Darwinella australiensis Carter, 1885b: 202 (Port Phillip character in this family, and variation from rose Heads, Vic.).-Lendent"eld, 1885a: 22.-Lendenfeld, to yellow would not occur" stands isolated, and she 1889b: 679, pi. 46 figs 1-5, II (rcdescription of holo- does not elaborate on other (axonomic characters. type). -Topsent, 1905a: CLXXVI (resume, old Other workers (French, Italian, and Spanish record). — Hentschel, 1912: 429 (Aru Islands, Arafura Sea, Indonesia). -Dendy, 1916b: 139 (Okhamandal, India). - authors on Mediterranean and Atlantic records) Levi, 1952: 38, fig. 1 (synonymy, description, discussion; have followed Vacelet (1959) in merging the two Senegal). -Pronzato, 1975: 8f., 16, text-figs. 2, 7(2), 8 species in question. According to Pulitzer-Finali (synonymy here expanded, resumes, discussion, distribu- (1977: only Descatoire (1969) has and Pronzato 98) tion; note on colour variants, horny spicules; Ligurian distinct, besides subsequently kept the two species Sea and Gulf of Naples). — Pulitzer-Finali and Pronzato, Bergquist (1967). Of the other workers (Levi, Vace- 1977: 99 (notes on horny spicules, colour variants).—

let, Boury-Esnault, Sara, Pulitzer-Finali), none has Pulitzer-Finali and Pronzato, 1980: 153 (remark on colour

apparently found other differences, besides colour, variants). - Rubio et al., 1981 : 43, text-figs 13, 14 (descrip- distribution and ecology). as in anatomy, ecology, or reproduction, in sym- tion, Darwinella simplex Topsent, 1892c: xxvii (Mediterra- patic populations of the pink and yellow form. Of nean coast of France).— Topsent, 1904: 55, pi. 9 fig. 3 some interest, in this context, is that Pronzato (Azores). -Topsent, 1905a: CLXXVI (resume, discus- (1975: 7, 10, 12), in discussing the taxonomic value sion). -Dendy, 1905:204, pi. 15 figs 1, 2 (Sri Lanka). Darwinella, noted the of colour in Aptysilla and Burton, 1959: 268 (synonymy, record only; south coast presence in the Mediterranean, of the bluish purple of Arabia). phenotype of Darwinella australiensis (see also Darwinella intermedia Topsent, 1894b: xxxvii (Banyuls, Pulitzer-Finali, 1977: 99; Pulitzer-Finali and Pron- Mediterranean coast of France). -Topsent, 1905a: fig. 1 (resume). -Borojcvic et al., 1968: 29 zato, 1980: 153). Another point is my observation CLXXVII, 148 F. WIEDENMAYER

(near Roscoff, English Channel). -Pronzato, 1975: 12, in F52063, comparatively subdued in F52062. Spac- 16, 18 (resumes, synonymy, distribution, key). ing of conules mostly 2-7 mm, but some larger Darwinella duplex Topsent, 1905a: CLXXIX (replace- smooth interstices occur, particularly in depressions for Darwinella simplex: ment name Topsent, 1904). between lobes. Darwinella rosacea Hechtel, 1965: 17 (Jamaica). Oscules 1-4 mm wide, situated mostly on top and Darwinella viscosa Boury-Esnault, 1971: 334, pis 6, 7 on edges of lobes, also on sides, in depressions (Mediterranean coast of France). between conules, some hidden and almost closed. Darwinella muelleri.—de Laubenfels, 1 950b: 38, text- fig. 16 (nee Schultze; Bermudas). Surface in depressions regularly dotted by mouths of inhalant canals, around 1 mm wide, spaced 2-5 Material examined. Station BSS 180, one specimen mm, clearly visible through ectosome. Ectosome (F52062); station BSS 181, two specimens (F52063, contains regular reticulum of collagen fibres, barely F52064). Type specimens of Darwinella australiensis visible with unaided eye, distinct at low magnifi- (BMNH). cation; its meshes square to polygonal in centre of Diagnosis. Thickly encrusting, cushion-shaped, or depressions, 200-300 /an wide, elongate, with radial irregularly compressed-lobate, digitate, tongue- nervures around conule tips. 4 to 10 ostia to each shaped. Surface coarsely conulose, conules angu- mesh. lar, compressed, pyramidal, often connected by From fracture in F52064, a fibre protrudes for ridges. Soft, limp, easily torn, more or less viscid. 15 mm, regularly branched and tapering, branch- Colour in life variable: brownish orange, deep red, ing points 3 to 5 mm apart. Abundant corneous bluish-violet, purple, pink, yellow, orange. Oscules spicules of this specimen pink in reflected light. 1-4 mm wide, mostly apical and marginal, but also Ectosome 45-200 /mi thick, frequently stratified, on flanks and in depressions. Ectosome with two or three layers: (1) Outer, evenly granu- diaphanous, with regular collagenous reticulum. lar layer, with radial or oblique ostia mostly con- Fibres pink, mostly isolated, basally expanded, spicuous; (2) inner, tangentially fibrillate layer with short and sparsely branched in encrusting speci- indistinct radial ostia; and (3) discontinuous layer mens, similar to those in Aplysilla; longer, more of crypts. Stratification and third layer most dis- prolifically dendritic in lobate specimens. Horny tinct in F52062, ectosome thus detachable between spicules of variable abundance, tend to be conules. In other two, stratification may be segregated by size and individuals: commonly pink, indistinct or obscured by interfingering of layers, with crooked rays when large, pale and regular and crypts may occupy part of fibrinous layer. In when small, most frequently triactinal, less often F52064, ectosome contains mulberry-shaped tetractinal, rarely pentactinal and diactinal. microsymbionts, 17-25 urn in diameter, mostly packed in outer layer, crowded to scattered below, Description. F52062 basally attached to fragment also in peripheral choanosome. "Fibrils" of second of shell, compressed-clavate, expanding from width layer possibly filamentous microsymbionts. of 7 mm proximally, to tongue shaped end 18 mm Chaonosome stringy, with subparallel wavy wide and 4 mm thick. Length of specimen 5.5 cm. canals, and elongate, crowded to packed choano- Other specimens (pi. 16 fig. 12, pi. 17 fig. 1) more cyte chambers aligned inbetween. Choanosome massive, with clustering, angular, more or less com- contains much detritus in F52063, few foreign pressed lobes separated by deep, narrow depres- inclusions in other two. F52064 contains ova, sions or invaginations. F52063, 2 x 2.5 x 6 cm; 72-208 /tm in diameter, surrounded by thick layer F52064, 1-2.5 x 2-3 x 4 cm. Base of attachment of follicle cells. small, on narrowest side on both specimens. Fibres comprise typical stratified bark and strag- Brownish orange in life, lighter and duller in gling knotty pith with timble-shaped accretion lines. F52062 (5 YR 6/8), deeper in other two (2.5 YR Fibre diameters commonly 132-272 /im below out- 5/10). Colour now faded to drab, more so in ermost divisions, rarely reaching 373 /on. Pith 7-8/4) than in F52062 (7.5 YR other two (2.5 YR diameters commonly around 20% of thicker fibres, 6/4 to 5 YR 7/4). Soft, limp, slightly viscid. Some 40% of thinner ones. Regularly spaced elongate clear mucus emitted after collecting. cells or pits occasionally seen on surface of pith. Most of surface irregularly conulose; conules Junctions of branches either "grafted", or continu- compressed to pyramidal, occasionally bifid or ous, with confluent pith (cf. Boury-Esnault, 1971: clustering, contorted, connected by coarser or finer 335). variable relief. Ridges ridges of often aligned across Horny spicules rare, pale and small in F52062 several conules, sparingly branched and and F52063, with mostly equal, straight and anastomosed, forming incomplete rugo- pointed rays in one plane, 96-108-1 18 x 10-13 ^m reticulation. Pattern coarser and most pronounced in F52062, 72-53-97 x 9-12 /tm in F52063. Horny SPONGES FROM BASS STRAIT 149

spicules in F52064 much more conspicuous (pi. 36 above synonymy. For the remaining records, vari- fig. all larger, 1), mostly red-brown in transmitted able proportions of diactines and tetractines were light (two very pale), about as frequent as in indicated. diagrammatic figure in Lendenfeld (1889b, pi. 46 It should be noted that there is some overlap, fig. but 2), rays rarely straight, commonly bent, albeit small, in spicule dimensions (length of rays) curved or flexuous, with thin pith of variable dis- between the specimens of Hechtel (1965), of Rubio tinctness. In six sections examined, 62 complete et al. (1981), and F52064. Nevertheless, there is spicules were observed: 34 triactines, 27 tetractines, obviously a strong tendency in this species, as here one pentactine. Of former, some have atrophied conceived, for spicules to be segregated by sizes in fourth ray not exceeding its thickness in length. different specimens. The significance of this is not Rays 187 x 19 ^im to 1347 x 40 /mi. A single pale, understood at present, since, as noted in two of the unequal smaller triactine, longest ray 93 X 7 /im, new specimens, it may occur in strictly sympatric may be foreign. populations. This might be taken as ruling out both biogeographical and ecological (niche) segregation. Remarks. Contrary to Levi's (1952) and Pronzato's But cohabitation of sibling species or subspecies

(1975) opinions, Darwinetla intermedia is here in our area does not preclude spatial isolation of

merged in D. australiensis. I could not detect any the same in other regions, or (particularly for significant difference in structure of ectosome, southern Australia with its complex and poorly choanosome and fibres in the new specimens, of understood biogeographical history and position) which the last two (with identical colour) came in the past. from the same collecting station. Topsent (1894b) The latter point should be viewed in perspective described D. intermedia as golden yellow in life of temporary disjunction (in geological terms, i.e. with small triactines having flexuous rays 100-150 through palaeogeographical evolution) of demes or dines, nm long (67-165 nm in fig. 1 in Topsent, 1905a), and secondary sympatric reunion with or 12-13 nm thick at the base. Darwinetla rosacea, without effect on the panmictic balance, and the merged in D. intermedia by Pronzato (1975: 12), difficulties in justifying subspecies in sponges (see was described by Hechtel (1965: 17) as pink in life, review in Wiedenmayer, 1974). pinkish brown or beige in alcohol, with spicule rays Considering that most descriptions of records of measuring 130-276 x 10-17 /mi. "Venetian Red", this species complex outside the Mediterranean are the colour indicated by Carter (1885b: 202) for the based on single specimens, it seems preferrable, at living D. australiensis, probably means brownish present, not to distinguish a nominotypical subspe- vermilion (10 R 5-6/8-10), while that in alcohol cies from a ssp. intermedia.

is given as "dark grey flesh". Dendy (1905: 204) indicated dark purple in alcohol for his specimen Darwitiella gardineri Topsent simplex. viscosa orange in life (Boury- of D. D. was Plate 17 figures 3, 7, text-figures 96, 97 Esnault, 1971: 334). Pronzato (1975: 10, 12) gave gardineri Topsent, 1905a: dark red, violet, and light bluish purple as colour Darwinetla CLXXIX (Fadifolu Atoll, Maldive Islands, Indian Ocean). — Levi, variants in life for specimens of D. australiensis in 1958: 42 (Abulat Island, Red Sea). -Pronzato, 1975: 13 the Mediterranean. Rubio et al. (1981: 46) described (resume), 16 (distribution), 18 (in key), figs 5 (spicules), the colour of D. australiensis as a strong pink in 9 (in situ, colour, Ligurian Sea, new record). — Pulitzer- in material are pink, com- life. The spicules their Finali and Pronzato, 1980: 152 (brief description includ- paratively small triactines (exclusively), with rays ing colour; Ligurian Sea). measuring 200-750 x 14-32 /im. Segregation of Material examined. Station KG 4, one specimen (F52065); spicules by categories within specimens was station KG 7, one specimen (52066). reported by Topsent (1905a: CLXXIX, for D. duplex), Levi (1952: 38, for D. australiensis), and Diagnosis. Encrusting to irregularly angular-lobate, Boury-Esnault (1971: 334, for D. viscosa). or lacunose-clathrate. Softly to firmly rubbery, Few indications are found in the literature con- slightly viscid; deep yellow in life, with dark red- olivaceous, then cerning the relative frequency of spicules. Contrast- brown fibres, turning dark pur- after collecting and preservation. ing with the high frequency in Lendenfeld' figure plish black Oscules commonly as contorted slits, chiefly mar- (1889b, pi. 46 fig. 2, based on Carter's material) ginal. Surface coarsely conulose, with conules com- and in F52064 are Dendy's statements (1905: 204; pressed, abundant on edges of lobes and 1916b: 139) as to the rather scarce occurrence of more like cocks' combs; otherwise smooth to spicules in his specimens. Exclusive occurrence of lamellae, finely wrinkled. Ectosome 100-200 (im thick, triactines was recorded for D. intermedia and D. very detachable. Choanosome dense, with com- rosacea, but also for some other records in the not 150 F. WIEDENMAYER

«7 !A^**

v ^\ 97

Figures 96, 97. Darwinel/a gardineri. F52065, details in reflected light. Figure 96. Perpendicular section showing thickness of ectosome, canal system, and fully overgrown different sponge (lower right), x8.2. Figure 97. Portion of surface showing inhalant fields, x6.6.

canals, smaller ones often parallel (tiger-skin pat- Description. F52065 (pi. 17 fig. 7, now dissected) tern in sections). Dendritic fibres more profusely angular, lobate, with thick, boxy lobe on one side, branched below edges, tapering little, may be filled and compressed one, transversely folded, on oppo- with filamentous algae. spicules Horny numerous, site side; 1 to 1.8 cm in thickness, 6 cm in width, diactinal, smooth. 1.5-2.5 cm in height. Some folds between lobes SPONGES FROM BASS STRAIT 151

narrow deep, invaginations. Small, multiple base Outer zone of ectosome about 30 /mi thick, more of attachment. deeply pigmented. Peripheral choanosome more Most of surface irregularly and coarsely conu- deeply pigmented, often with palisade of parallel lose. Conules chiefly around tops of lobes, 1-2 mm inhalant canals or vestibules, about 30 fim wide. high, mostly 2 mm apart, often contorted, con- Choanocyte chambers oval, 56-97 /im long. nected by ridges. Some fibres protrude from conule Dendritic fibres more profusely branched just tips. Some conules also on lower sides, where mar- below apical oscules (edges of lamellae in F52066), ginal dendritic fibres bulge below shrunken surface, occasionally with oblique anastomoses, by concres- with often wide, smooth depressions between cence of adjoining branches. Most fibres 200-270 ascending ridges. Along edge of thinner lobe, /mi thick, diameters at base up to 500 /int. Fibres several fibres exposed, already upon collecting. taper little, diameters around 200 /mi at 4-5 mm F52066 (pi. 17 fig. 3) lacunose, clathrate aggre- below conule tips common. Pith, with weak timble- gation of contorted lamellae, with angular outline, shaped accretion lines, commonly 33% to 47% of 3.5 cm high, 6 x 8 cm wide. Lamellae mostly 2-4 fibre diameter, often almost clear, weakly granu- mm thick; lacunae between them, open below and lar. In F52066, filamentous algae common in bark above, some marginal ones also on sides, 0.5-1 .5 and pith. cm wide. Choanosome contains numerous diactinal spic- Surface generally more coarsely conulose than ules in both specimens, some protruding from sur- in F52065, with conules up to 5 mm high; many face along edges of lamellae just above base of aligned on outside and edges of lamellae, like attachment, straight to slightly sinuous, occasion- clustering cockscombs. Inner sides of lamellae ally bent, very pale, with faint contrast in Canada (walls of lacunae) mostly smooth, with few con- balsam, clear except for axial filament, tapering ules. Base of attachment small and multiple, some gradually and symmetrically to sharp points. Few fibres partly free, many protruding from conules. entire ones isolated, 1550-1760 x 15-24 /tm. F52065 has completely overgrown small pinkish drab sponge with sandy ectosome (text-fig. 96). Similar, if not conspecific sponge attached to one Remarks. Specimens of this species hitherto side of F52066. In life, sulfur-yellow in F52065, recorded were encrusting, with low, simple or golden yellow (10 YR to 2.5 Y 8/10) in F52066. sparsely branched fibres. The colour of Topsent's Soon after collecting change to dark olivaceous, holotype in life is unknown, but is stated to be pink purplish black (5 R 2/2) in alcohol, choanosome ("rose") in alcohol. Levi (1958: 42) indicated vivid slightly lighter in F52065 (5 R 3/4-4). Fibres very yellow, turning purple in alcohol. Pulitzer-f inali conspicuous in life, dark red-brown, visible also and Pronzato(1980: 153) gave the index C.C. 316 just below ectosome, colour preserved in alcohol. in Seguy (1936, Code Universe! des Couleurs. Paris: Now compressible, slightly limp in F52065, firmer, Lechevalicr), which is lemon yellow, equivalent to almost leathery in F52066, slightly viscid. 7.5 Y 8-9/8-10 in Munsell. Spicule dimensions Oscules more conspicuous on F52065, contorted given by Topsenl (1905a: CLXXX) arc 1600-2000 slits up to 5 mm long and 0.5-1 mm wide, mostly x 20 /tm (half as much for a few). Pronzato (1975: on upper edges of lobes, some also laterally. With 14, fig. 5) indicated lengths of 804 to 1780 /tm for in Mediterranean material. low magnification (10 x ), surface smooth to very diactines his finely striate or wrinkled in many areas; other The occurrence of massive-lobate specimens in areas, particularly between conules, bear discon- this species is not surprising, given a similar range tinuous inhalant fields with sinuous outlines (text- of shapes in D. australiensis. The lacunose-clalhrate is reminiscent of Dendrilla fig. 97), occasionally connected by meandering habit of F52066 bridges, with reticulation of slightly elevaied col- lucutwsa Hentschel (1912: 431, pi. 15 fig. 3), the lagenous bands, somewhat lighter than remaining general shape of which, however, is terete-ramose, fibre radial, surface in F52065. Bands aboul 100 /tm wide out- with a very strong axial and thinner line rounded depressions of similar width pierced branching fibres. filamentous algae in the fibres by ostia. Occasionally, radial pattern of bands con- The presence of occurrence in Mycale (Arenocha- verging on conule tips. Regularly pitted fields less recalls the similar described above. It is not unique in distinct in F52066. lina) mirabilis found Ectosome 100-200 /*m thick, generally not Darwinella, since Dendy (1905, 1916b) a detachable. Choanosome dense, cheese-like, with similar association (interpreted as "parasitic his material of Dendrilla australien- tiger-striped pattern of subparallel inhalant canals fungus?") in simplex in 1905). (text-fig. 96). Terminal exhalant canals mostly com- sis (as sandy sponge recalls an analogous pressed, section like outline of oscules. The endozoic -

152 F. WIEDENMAYER

association of "Gellius" and Aplysilla described by Phillip Bay, Vic; Adelaide, SA). -Carter, 1886f: 288 Riitzler (1970: 94) from the Adriatic coast. (remarks on histology). -Dendy, 1890a: 79 (Port Phil- lip Bay).- Vacelet, 1958: 143, 144 (in review). -Koltun, Dendrilla Lendenfeld 1971: 42 (Red Sea). -Coleman, 1977: 17 (in colour). Bergquist, 1980b: 488, fig. 21a (in review). — Bergquist Dendrilla Lendenfeld, 1883: 270. -Lendenfeld, 1889b: and Skinner, 1982: 49 (fig. 3.7), 54; pi. 1 fig. 3 (colour). 709 (part: D. rosea and D. aerophoba only). — Topsent, Dendrilla rosea var. typica Lendenfeld, 1888: 28 1 905a: CXC (subsequent designation of type species). — (nomen vanum for nominotypical subspecies; Sydney Hentschel, 1912: 430 f. (part: D. lacunosa, D. mertoni Harbour, NSW). -Lendenfeld, 1889b: 718, pi. 44 figs 4, spp. nov.; not D. lendenfeldi, a Dictyodendri/la). — de 7, 1 pi. 45 figs 3, 4, 7, 9, 11 (old records). -Whitelegge, Laubenfels, 1936a: 32 (invalid designation of type spe- 1; 1889: 180 (off Green Point, Sydney Harbour; probably cies).— de Laubenfels, 1948: 153 (part, wrong type spe- Lendenfeld's records). — Hentschel, 1912: 430 (Aru cies, wrong transfers). — Vacelet, 1958: 143-147 (resume, Islands, Arafura Sea, Indonesia). definition, discussion, erroneous revision, new Mediter- Luffaria digitala Carter, 1885b: 201 (Port Phillip ranean species).— Vacelet, 1959: 65 (definition, discus- Heads, Vic). sion, resume of Mediterranean species). — Van Soest, Dendrilla rosea var. digitala. — Carter, 1886f: 281 1978: 81 (discussion of possible West Indian records).— (Westernport Bay, Vic). -Lendenfeld, 1889b: 718, pi. Bergquist, 1980b: 486 (definition, discussion, revision). 44 fig. 8, pi. 45 fig. 8 (old record). Diagnosis. "Large, erect, branching or complex Aplysilla rosea (sic, nee Barrois). — Rockman, 1974:

lamellate [Darwinellidae] in which the fibres branch 28f. (in i olour, no locality). repeatedly but do not anastomose. The fibres con- Material examined. Station KG 2, one specimen (F52067); tain no foreign coring material." (Bergquist, 1980b: Station KG 7, two specimens (F52068, colour slide only; 486.) F52069, skeleton only).

Dendrilla cactos (Selenka) Diagnosis. Massive, irregularly lobate to bushy and pedunculate, or branching, erect, with isodiamet- Text-figure 98 ric branches. Light red to purplish pink in life, drab

Spongelia cactos Selenka, 1867: 566, pi. 35 fig. 5 (Bass to pale orange and red-brown in alcohol. Soft, Strait). limp, firmer in branching specimens. Flesh tends Aplysilla cactos. — Schulze, 1878a: 417 (holotype re- to collapse and dissolve after collecting, fibre skele- examined, redescribed). ton thus easily macerated. Surface coarsely and Dendrilla cactus. — de Laubenfels, 1948: 152 (part: not regularly conulose, with conules compressed- pi. 26 fig. 47; description composite, partly based on pyramidal, often complex, contorted, with taper- Megalopastas retiaria Dendy, a Dictyodendrilla). ing fibre tips protruding. Surface nee Dendrilla cactus. — Bergquist, 1961c: 215, figs 5a, b. otherwise Megalopastas cactos. — Vacelet, 1958: 144 (in revision, smooth, collagenous reticulum indistinct. Fibrous unrecognisable). skeleton thoroughly dendritic, with basal plate, and Dendrilla rosea Lendenfeld, 1883: 271, pi. 10 figs 3, very thick fibres in peduncle of bushy specimens 4; pi. 12 figs 16, 19-23; pi. 13 figs 24-27, 29-32 (Port and axis of ramose ones.

Figure 98. Dendrilla cactos. F52067, portion of surface showing inhalant fields with elongate ostia, x6.6. SPONGES FROM BASS STRAIT 153

Remarks. This is one of the most common and sioides or D. acantha, the only species presently characteristic species of the southern Australian known from the Mediterranean (see Pulitzer-Finali, shelf fauna, and is likely to be illustrated in other 1983: 613). popular books than the three referred to above. No Bergquist (1980b) recognised only one species complete redescription is therefore needed here. (rosea) with Australian records in Dendrilla. Den- According to my field notes, the first specimen, drilla aerophoba Lendenfeld (1883), which Berg- a bushy one, had a pure rosy pink colour in life quist (1980b: 488) regarded as unrecognisable, may (5 RP 8/8-10). The second specimen, decidedly or may not be a true Dendrilla, depending on ramose, was light red (5 R 7/10-12). The colour whether the peculiar fibre structure (strong longitu- in alcohol of the first specimen is drab to pale dinal folding of the bark) and the regularly orange in depressions (7.5 YR 6-7/2-8), red-brown armoured ectosome are regarded as sufficiently around the conules (10 R 4-5/6-8). aberrant within Dendrilla. The strong colour

The consistency is soft and limp, and the flesh change upon preservation (from brilliant yellow to has a pronounced tendency to collapse, shrink and dark blue or bluish black, reminiscent of Dar- dissolve into tatters in seawater soon after collect- winella gardineri) is also distinctive. It can there- ing. The fine dermal reticulation is not as clear as fore be ruled out that what Selenka and Schulze in the other darwinellids described above, and is described was D. aerophoba, and it is highly usually better developed in irregular fields between unlikely that they had before them a third species conules, as in Darwinella gardineri. Here, the of Dendrilla, or a Darwinella in which they over- meshes with the cribripores are frequently elon- looked the spicules (Schulze mentioned examining gated radially (text-fig. 98), possibly due to the a section). shrinkage, which leaves many of the pale fibres Schulze had borrowed the holotype from the bulging and partly denuded. Zoological Institute in Gottingen (where Selenka Whether single, continuous, thick axial fibres worked in 1867). The Selenka collection is no with short, much thinner lateral branches (as in pi. longer there (teste Prof. P. Kuenzer, II. 45 fig. 8 in Lendenfeld, 1889b, 3-5 mm thick near Zoologisches Institut und Museum der Universitat, the base) are consistently developed in terete- Gottingen, 1983), and nothing is known of its ramose specimens, and whether digitata is justified present whereabouts. The holotype of Dendrilla as a form or subspecies, must be left open to ques- aerophoba Lendenfeld, however, is extant in East tion for now. F52068 (unfortunately lost overboard Berlin (ZMB 2254 with one slide; ZMB 10395, slide). soon after collecting) had this type of skeleton. I second suspect that taxonomic distinction is not justified, Burton (1934a: 595) included Spongelia cactos because Hentschel (1912: 430) described a terete- and Dendrilla rosea in his highly composite syn- ramose specimen having tight fascicles of fibres in onymy of Dendrilla membranosa (Pallas). Spon- the axes, and a transitional specimen (which he gia membranosa Pallas should be dropped as a compared to pi. 44 fig. 7 in Lendenfeld, 1889b) had nomen dubium. According to the original descrip- a similar structure. tion it was whip-shaped, 3.5 feet tall, finger-thick, Lendenfeld (1889b: 718) included Spongelia with a single branch. Whatever it was, it was not cactos Selenka in the synonymy of Dendrilla rosea a Dendrilla. Spongia membranosa: Esper (mem- var. typica, which, if accepted, has to replace rosea branacea in the caption of pi. 34 is an incorrect sub- spelling) is an lotrochota, as as its senior synonym. This synonymy is here sequent Topsent accepted not because of Selenka's inadequate and (1920c: 322) has shown. Dendrilla membranosa misleading description (claiming that the skeletal sensu auctorum is composite. Records from northern Australia (Ridley, 1884a: 398; Lendenfeld, structure is the same as in Spongelia horrens, i.e. to Dic- reticulate), but because of the redescription of the 1889b: 715; Bergquist, 1980b: 488) belong holotype by Schulze (1878a: 417, cited by Lenden- tyodendrilla and will receive a replacement name (Wiedenmayer et al., in press). Antarc- feld, 1888: 28; 1889b: 718). This is sufficiently clear tic/Subantarctic records (Burton, 1929a: to establish the synonymy. The fibres are described 448; Burton 1934b: 46; Burton, 1938: 19; Koltun, 1964: as clear, dendritic, with the structure of those in Koltun, 1976: belong to Dendrilla antarc- Aplysilla. The interstitial choanosome is also said 111; 197) tica. the remaining entries in Burton's to be indistinguishable from that in Aplysilla. Among excessive synonymy are good species of Dendrilla Schulze, five years before Lendenfeld established and Dictyodendrilla. Dendrilla cactus: Bergquist Dendrilla, included massive forms with dendritic- (1961c: 215) is described as having a dendritic skele- fibres in Aplysilla, and cited, in the same context, with occasional anastomoses; but in figure massive specimens from the Adriatic, which today ton 5b anastomoses appear to be rather common, suggest- would probably be identified as either Dendrilla cir- 154 F. WIEDENMAYER

ing that the specimen might belong to Dictyoden- late, partly loose secondary fibres, Fibre- drilla. A special feature is the reinforcement of the reticulation much denser in peduncle. ectosome by spicule fragments. Description. Broadly flabellate, more or less sym- Order Verongiida Bergquist, 1978 metrical, with stocky pedicel 3.5 cm long in front,

Aplysinidae Hyatt, 1875 1 .8 cm thick in middle, its base slightly jutting foreward, torn at back. Flabellum now contorted, 17.5 Aplysina Nardo cm wide, 1 1 .5 cm high in middle, of variable thick- [Aplysia] Nardo, 1833, col. 519 (nee Aplysia Linne, ness (0.5-2 cm). Lateral transition of pedicel into 1767). flabellum elegantly curved. Outline of flabellum Aplysina Nardo, 1834, col. 714 (in part; nomen novum sinuous to angular. Right side, near margin, thicker for Aplysia Nardo). — Wicdenmayer, 1977a: 63 (syn- than left, consisting of several overlapping lobes onymy with further references, definition, discussion). - compressed in plane of flabellum, in two to three Van Soest, 1978: 55, 63 (definition, discussion, revision tiers. of West Indian species). — Bergquist, 1980b: 492 (syn- Pedicel its onymy, definition, discussion, revision). and arched transition into flabellum [Fistularia] Bowerbank, 1844a: 39 (nee Fistulariu Linne, almost smooth, faintly and finely wrinkled longitu- 1758). dinally. Front of frond traversed by some ridges, Vero.tgia Bowerbank, 1845: 403. - Vacelel, 1959: 86 mostly radial, 3-5 mm wide and deep, locally (definition, discussion, review of Mediterranean species). (lower middle) up to 15 mm deep. Most of surface Luffaria Duchassaing and Michelotli, 1864: 59. rough, combination of low conules and wrinkles nee Luffaria sensu Polejaeff, 1884b: 69. - Lendenfeld, of variable profile, never forming distinct rugo- 1889b: 382 (= Luffariella in part). reticulation. Conules and wrinkles rarely exceed 1 Diagnosis. "Aplysinidae characterised by posses- mm in width and relief, with interstices commonly sion of fibres of only one kind with no foreign 1-5 mm wide. Depressed interstices smooth or very detritus and having a thick pith component. The finely wrinkled. Back of frond with some larger fibres form a regular reticulum with large hex- smooth depressions. Margin commonly more agonal meshes and have no specialised surface decidedly conulose. arrangement. Most, if not all, species display a In life, red-brown on elevated parts, golden marked colour change at death or on exposure to yellow in depressions. Rugo-reticulation much air, quickly changing from the normal yellow or more regular and pronounced in life, accentuated yellow-green through blue to dark brown or by colour differences. Now brownish to purplish purple." (Bergquist, 1980b: 492.). Pedunculate spe- black, superficially dark brown in choanosome. cies with a secondary reticulation of finer fibres are Leathery to firmly rubbery, elastic along margin; here provisionally included. peduncle hard. Oscules inconspicuous, probably Aplysina ianthelliformis (Lendenfeld) as deep invaginations extending from bottom of some folds Plate 17 figure 10, plate 36 figure 2 between marginal lobes. Some round, sharp-

rimmed pits about 1 cm wide on both faces Dendrilia ianlhelliformis Lendenfeld, 1888: 29 (Sydney of frond. With low magnification Harbour). -Lendenfeld, 1889b: 719 (old record). (8-15 x ) surface of Aplysina ianthelliformis. -Bergquist, 1980b: 488 ectosome frequently like elephant hide covered with (transfer in review). metallic paint. Ostia occasionally visible as clustering faint papillae in sharply defined round fields, Material examined. Station BSS 187, one specimen which may be slightly raised. (F52070). Holotype of Dendrilia ianthelliformis (AM). In ruptured or partly macerated spots, peculiar Diagnosis. Stalked, flabellate. Frond up to 2 cm main skeleton visible in flabellum: tangled, often thick. Leathery to firmly rubbery, peduncle hard. dense, with chiefly dendritic primary fibres, and In life, red-brown on elevations, golden yellow in irregular reticulation of distinctly thinner secon- depressions; turning brownish to purplish black daries. In reflected light, all fibres externally brown, after collecting and preservation (choanosome then with much lighter pith shining through. dark brown). Surface indistinctly conulose and Fleshy ectosome 225-640 pm thick, partly rugose, with vague rugo-reticulation. Oscules detachable, frequently covers extensive thin crypts. inconspicuous, in marginal invaginations. Ostia in Straggling primary fibres have diameters 100-185 indistinct areoles. Fleshy ectosome about 0.5 mm lim, 63% to 86% pith, spaced around 800 pm. thick, partly detachable over thin crypts. Choano- Secondaries knotty, 14-50 /mi thick, pith variable some dense. Fibre skeleton dense, tangled, with (27%-48%). Anastomosis prevails, with irregular chiefly dendritic primaries and irregularly reticu- meshsizes, from equal to diameters of secondaries SPONGES FROM BASS STRAIT 155

to order of spacing of primaries. Occasional short about 1 mm deep, mostly elongate axially, 1-2 mm fenestrate fascicles. Junctions with primaries wide, 2-5 mm long, with tabulate interstices 1-2 "grafted", pith truncated by continuous surface mm wide. Depressions on pedicel reduced to slits. (bark) of primaries. Many secondaries dendritic, Relief now partly disappeared through shrinkage, some short and stubby. leaving scar-like, shiny dots sharply set off from matte interstitial surface. Remarks. See next species. Upper half of pedicel and base of body now show rugo-vermiculate to rugo- Aplysina lendenfeldi Bergquist reticulate pattern of peripheral fibres bulging through shrunken ectosome. Plate 17 figures 4, 5, plate 36 figures 3-6, Single apical oscule 3x6 mm wide in life, plate 37 figure 1 slightly recessed within truncate apex, with fleshy

contractile membrane and slightly upturned lip. Thorecta Jre ij a.— Lendenf eld, 1889b: 366 (part: F52072 much larger (pi. 17 fig. paralectotypes from Wollongong and Broughton Island, 5), also clavate, NSW, in AM only; not figured lectotype from Torres to cypressiform, straight in life, now bent above like Strait in BMNH; fide Bergquist, 1980b). pedicel, F52071 . Pedicel torn off above its base, Aplysina lendenfeldi Bergquist, 1980b: 470, 488, fig. 9 cm long, 12 mm wide, terete, partly enveloped, 22e (for misidentified paralectotypes of Thorecta freija in lower half, by compound ascidian. Main body and fresh material; new locality (localities?) not indicated). 13 cm long, 4.5 X 5 cm wide in middle, 1.5 x 2.5

Verongia sp. — Bergquist and Skinner, pi. 1 fig. 1982, cm wide at truncate apex bearing central single 2 (colour, no locality). oscule.

Material examined. Station KG 5, one specimen (F52071); Upon collecting, lower half of body showed faint station KG 7, one specimen (F52072). rugo-reticulation, accentuated by differences in colour, with meshes around 7 x 20 wide, elon- Diagnosis. Stalked, body commonly cigar-shaped, mm gated axially. More isodictyal, fainter and tighter with apical plateau bearing single collared oscule; rugo-reticulation partly on the upper portion, with occasionally compound, concrescent. Light orange other parts weakly verrucose. Surface now gener- to dull yellow, with red and purplish tinges in life; ally smooth, except irregular, weak rugae and folds, in alcohol, brownish black externally, dark orange dendritic along and diagonally across body. Some brown internally. Surface on sides, mostly includ- interstitial areas finely verrucose or granular. Fine ing upper part of stalk, faintly to coarsely rugo- rugo-reticulate pattern on upper half of pedicel, like reticulate, with more or less depressed meshes in F52071. usually elongate axially; or areolate, with areoles F52071 upon collecting dull yellow with purplish depressed in life, flush or raised, shiny in alcohol. tinges. F52072 basically light orange (2.5 YR 7/10) Ostia cribriporal in areoles and depressions. Ecto- with light red ridges on lower body (5 R 7/10) and some fleshy, thicker (up to 600 /mi) and stratified some purplish drab in depressions (2.5 R 7/4). Sur- on elevated areas. Regular chone-like inhalant face now brownish black (5 R 2.5-3/2), choano- canals below areoles, palisade-like above, converg- some dark orange-brown (5 YR 4-5/4). Body in ing in lower ectosome to join subdermal crypts. both firmly rubbery, pedicels hard. Choanosome dense around terete axial atrium. In F52071, depressions are inhalant fields Primary fibres chiefly dendritic and longitudinal, (cribriporal groups). Individual ostia now minute, with few anastomes in deeper choanosome, but barely visible in reflected light at 50 x magnifica- with numerous radial branches in periphery, ending tion. No such well defined fields in large specimen, below or within ectosome. Secondaries scarce, where ostia arranged in vaguely outlined, sinuous crooked, mostly as anastomoses between primaries, and confluent fields. occasionally dendritic. Fibre-reticulation much In dissected F52071, atrium axial in middle of denser in peduncle. body, width about equal to oscule, with lining simi- Description. F52071 smaller (pi. 17 fig. 4), a club lar in structure and thickness to ectosome. Choano- bent in middle. Lower part (pedicel) 5 cm long, some dense, with canals rarely exceeding 100 /tin somewhat compressed, 6 x 8 mm thick. Main in diameter. Longitudinal primary fibres prevail in body compressed in plane diagonal to that of deeper choanosome, spaced 500-800 ^m in pedicel, expands symmetrically to 3 cm above top expanded body with thick, light pith and compara- of pedicel, reaching 14-22 mm in width; then bent tively thin bark, chiefly dendritic, occasionally obli- at 30° and tapering slightly to truncate top, 8 x quely anastomosing, moderately straggling. Radial 15 mm wide. Compression less pronounced in life. branches common in periphery, with tips ending Upon collecting, whole surface below apex, and indside or halfway through ectosome. Diameters over much of the pedicel, with regularly spaced pits, of primaries (in F52071) 90-127 ^m, pith 64-74%. 156 I-. WIEDENMAYER

Secondaries, connecting primaries with "grafted" differs externally only by its expanding, trumpet- junctions, scarce, very crooked, occasionally den- to mushroom-shaped top, internally by thicker dritic, 25-40 /im in diameter, with pith of highly fibres (primaries 300-400 nm, secondaries 40-200 variable diameter. /mi) with relatively thinner pith. Structure of stratified ectosome not clear in larger specimen, mostly obscured by masses of Druinellidae Lendenfeld, 1889 mulberry-shaped microsymbionts, some scattered Synonym. Aplysinellidae Bergquist, 1980. in peripheral choanosome. Intracortical crypts common in F52072. In smaller specimen, with less Remarks. Lendenfeld's name (established in 1889a: crowded microsymbionts, ectosome 520-590 /tin 46, as Druinellinae) has priority. The acceptance, thick. In interstitial, non-inhalant areas, stratifi- here, of Druinella as valid senior synonym of Psam- cation most pronounced: (1) outer hyaline layer maplystlla has no influence on this priority. The about 12 /tin thick; (2) middle layer, 100-120 ^m latter would also apply if Druinella were interpreted thick, finely granular, almost devoid of microsym- as an invalid synonym of Psammaplysilla, as Berg- bionts; (3) inner layer, about 450 f*m thick, sub- quist did (1980b: 494), because of ICZN Article 40. tangentially fibrous, with numerous microsymbionts. In cribriporal fields, hyaline layer Pseudoceratina Carter thinner, also middle layer, here often packed with [Dendrospongia] Hyatt, 1875: 400 (nee Dendrospon- microsymbionts, inner layer comparatively thicker. gia Rdmer, 1864). Layers 1 + 2 traversed perpendicularly by regu- Pseudoceratina Carter, 1885b: 204. -Bergquist, 1980b: larly spaced very fine canals, 3-4 /im wide, spaced 494, figs 23a-c (synonymy, definition, discussion, revi- 12-25 /an; many continue below, but progressively sion).— Pulitzer-Finali, 1982b: 138 (new species, merge in wider canals (up to about 15 /

bodies, with thin spongin envelope of 2-3 laminae Diagnosis. "[Druinellidae] with sparse fibre skele- of faint radial structure (pi. 37 fig. 1). Interior ton organised on the dendritic plan typical of the finely granular, contains clearer core (occasionally family. Pith elements are present in the fibres, in eccentric) and more opaque outer zone. Total some examples almost to the exclusion of the bark,

diameter 1 12-176 fim, diameter of cores 37-55 /mi which [may be] evident only as occasional (31^/0-40%), width of the outer zone 28-43 /jm, peripheral patches. The matrix of the sponge is width of envelope 6-29 /*m. extremely dense and heavily infiltrated by collagen:

the texture is Remarks. The present species has in common with hence firm, and indeed often the preceding Aplysina ianthelliformis the hard, extremely hard and incompressible." (Bergquist, densely fibrous peduncle, expanded body with 1980b: 494.) Surface commonly coarsely conulose or papillate, or widely spaced, chiefly dendritic primary fibres, and rugose/pitted. irregular secondary reticulation of distinctly thin- Pseudoceratina durissima Carter ner, partly dendritic fibres (less developed in A. len-

denfeldi). Especially the latter trait does not fit Plate 17 figures 6, 8, 9, 11, Bergquist's diagnosis, and a new genus might be plate 37 figures 2-6 needed for such species. In many other species

assigned to Aplysina, particularly Indo-Pacific Aplysina purpurea. -Carter, 1881b: 103 (part, pi. 9 fig,

ones, with dry types only, the main skeleton is little 2 only, SW Australia; fide Carter, 1885b: 205, confirmed from my examination of the figured known (Bergquist, 1980b: 492). It is therefore not hypotype in question, BMNH 1877.5.21.1883). unlikely that a secondary fibre reticulation, even Pseudoceratina durissima Carter, 1885b: 204 (Port Phil- if scarce, occurs in some massive species, and has lip Heads, Vie.). -de Laubenfels, 1948: 131 (invalid neo- hitherto remained undetected. A new genus is here type designation). — Bergquist, 1980b: 494, fig. 23a judged premature at this stage, and the diagnosis (doubtful lectotype, as "holotype"). — Bergquist et al., of Aplysina is modified accordingly. 1980: 424, 425, 428 (biochemistry; Great Barrier Reef). The "ianthelliformis-group" comprises a third Aplysina laevis Carter, 1885b: 204 (Port Phillip Heads). species: Aplysina pedunculata (Levi, 1969: 969, pi. Material examined. Station BSS 187, four specimens 3 figs 3-5, pi. 4 figs 1-4; the last four also in Brien (F52O73-F52076). Type specimens of Pseudoceratina et al., 1973, fig. 431), from the south-eastern Atlan- durissima and Aplysina laevis (BMNH). Misidentified tic. It is closest to A. lendenfeldi, from which it hypotype of Aplysina purpurea: Carter, 1881b (BMNH). SPONGES FROM BASS STRAIT 157

Diagnosis. Massive, commonly with blunt crest, or (1) F52073, F52074, with conulose surface, thin thickly lamellate. Surface papillate or conulose, ectosome, columns and round enclaves of packed, with conules frequently aligned in radial or mean- agglutinated sand grains in periphery, rare pithed dering pattern on and below apical crest. Firmly fibres; (2) F52075, F52076, with papillate to rugo- rubbery to almost incompressible. Dark brownish vermiculate surface, thick ectosome, regularly scat- red, occasionally with yellowish spots, in life; dark tered to crowded detritus, prolific pithed fibres. purple or purplish black soon after collecting and In first phenotype, surface decidedly conulose, in alcohol. Alcohol and labels stained purplish. with blunt or truncate, often composite conules, Oscules inconspicuous, apical and lateral. Surface apices 0.5-1 mm wide. Conules tend to be aligned in depressions smooth, often with fine detritus on radial ridges towards crest, where usually more forming indistinct tangential reticulum. Choano- prominent. Ridges spaced 3-5 mm. Many conule some very dense, with much detritus. Tendency to tips (all in F52073) much lighter, with sand grains

segregation of two morphotypes: (1) conulose, with bulging through ectosome, often piercing it. thin ectosome, detritus often in ascending columns Few small oscules on both faces of F52073; and round enclaves agglutinated by spongin, rare F52074, in addition, has numerous oscules of same pithed fibres; (2) papillate and rugo-vermiculate, size range (0.5-1 mm) along edge. Magnified sur- with thick ectosome, scattered to crowded detri- face of F52074 with reticulum of nervures, radial tus, prolific pithed fibres. towards conule tips, single ostia in interstices. Only traces of such reticulum seen in F52073, where uni- Description. F52073 (pi. 17 fig. 6) thickly lamel- formly fine detritus at surface (in all four speci- late, tongue-shaped. Base of attachment on whole mens) is mostly aggregated in vague reticulation of lower width of erect lamella, 5 cm. Height 6 cm, its own here and there. thickness below 1-1.5 cm, 5-7 mm at margin. In basal part of lamella of both specimens, F52074 (pi. 17 fig. 9) long thick lamella, like choanosome almost packed with sand, thus virtu- asymmetrical wedge, with broad truncate margin. ally incompressible. In periphery, conules cored by Base of attachment interrupted by some lacunae, tips of straggling, dendritic columns of packed sand one wide. Specimen 12 cm long and 1-3.5 cm wide. grains, agglutinated here and there by little spon- Lamella slightly folded transversely on both sides, gin, becoming indistinct 1-2 cm below surface. rises gradually from height of 2.5 cm on one side, Also large round enclaves of lumped debris. Inter- to 6 cm above base on other. Width of truncate stices always contain scattered to crowded ill-sorted margin 10 to 17 mm. debris. F52075 (pi. 17 fig. 11) thick, blunt wedge, like Only one pithed fibre seen piercing surface in outer half of waisted stone-ax. Base has outline of F52073 (not thoroughly sectioned). F52074 had asymmetrical and rounded lozenge. Central crest three isolated, fairly straight to moderately strag- fairly symmetrical, rising outward to height 3-4 cm gling fibres sticking out from both sides. Neither on sides, then gently arched to median apex 6 cm base area revealed any fibres in thoroughly exposed above base. Margin rounded, 1-1.5 cm wide. choanosome. Of mounted brittle fibre fragments, Median portions of lower flanks bulging to height some terete, one knotty, with several constrictions. 2 cm above base, much more on one side, as nose- Diameters 379-514 /un, pith 69%-94%, bark finely like, slightly upturned lobe. Width of base 7.5 x stratified. Only one fibre in three hand-sections, 7 cm, maximum length of wedge 9.5 cm, height in composite, with convoluted brown spongin layers middle 6 cm. surrounding 2-3 pithed lacunae. Ectosome 50-90 F52076 irregular crescent, like recumbent /tim thick and contains uniformly fine detritus, chie- depressed croissant cracked open along convex side. fly spicule fragments.

Maximum length 1 1 .5 cm, width in middle 4.5 cm, In second phenotype, surface uniformly papil- height above base of attachment 2.2-3 cm. late to rugo-vermiculate, with papillae about 1mm Dark brownish red in life (5 R 4-5/10), occasion- wide, 2-4 mm apart. Surface, with low magnifica- ally with yellowish spots (in F52076), soon changed tion, finely dotted throughout, with reticulum by to dark purple (7.5 RP 3/2) or purplish black upon slightly raised bands of fine detritus enclosing collecting, conserved in preserved specimens. Alco- round or elongate meshes (depressions) bearing hol and labels stained purplish red-brown, similar single ostia. Meshes generally wider than bands, residue on inside of jar not easily removed after but reversed in recessed area along convex side of draining first alcohol. Firmly rubbery, moderately F52076. to very little compressible, depending on amount Ectosome 200-400 /tm thick, fleshy, with fine of incorporated sand. Phenotypic variability is crypts in inner portion, contains few scattered for- amenable to distinguishing two forms: eign inclusions of medium dimensions, and much 158 F. WIEDENMAYER

finer detritus, chiefly in outer portion. No relation- Druinella Lendenfeld ship between papillae of surface and tips of fibres Druinella Lendenfeld, 1889a: 46 (monotypic definilion generally ending just below ectosome, no regular in Spongiidae, published 28th February). — Lendenfeld, crowding of peripheral sand grains below papillae. 1889b: 425.-Thiele, 1899: 24. -Wilson, 1925: 491 (dis- Ill-sorted detritus in choanosome, including cussion). -Burton, 1934a: 594 (part, fide Bergquist, 1965: coarse, angular grains, scattered in F52075, 139). — de Laubenfels, 1948: 97 (part, resume for the type crowded in F52076. Dendritic fibres numerous, species, not Cacospongia camera, see Bergquist, 1965: visible in both base areas and in all sections, irregu- 139,-vanSoest, 1978: 81). -Bergquist, 1965: 138f. (dis- larly straggling, knotty, with meandering, even cussion, unrecognisable). — Bergquist, 1980b: 494 (as syn- knottier pith. Fibres light in reflected light. In trans- onym of Psammaplysilta). nee Druinella. — Burton, 1959: 269 (fide Bergquist, mitted light, finely and irregularly stratified bark 1965: 139). brilliant yellow, pith grey to faintly yellow, occa- Psammaplysilla Keller, 1889: 358 (published 15th sionally opaque, almost black. In longitudinal sec- November). —de Laubenfels, 1948: 172 (discussion, tions, pith commonly irregular, ex-axial, with resume). — Bergquist, 1965: 135f. (synonymy, definition, accretions hemispherical to almost spherical discussion). — Bergquist, 1980b: 494 (ditto). (reminiscent of some sphinctozoans), with inside [Korotnewia] Polejaeff, 1889: 366 (nomen oblitum).— of bark reaching into constrictions, occasionally Topsent, 1905a: CLXXIV (discussion, genus inquiren- thinly enveloping pith dissepiments. Lobed accre- dum possibly related to Psammaplysilla). — de Lauben- tions of pith occasionally bud sideways. fels, 1936a: 32 (genus inquirendum). — Bergquist, 1980b: 494 (as synonym of Psammaplysilla). In F52075, fibre diameters 1 18-/59-169 /tm, pith

67-79-83%. In F52076, fibres 79-118 /xm, pith Diagnosis. "[Druinellidae] in which the form of the

48-67%. sponge is encrusting to complex-ramose [or lobate]. Three specimens contain crowded microsymbi- The skeleton, although having the basic dendritic onts in choanosome. In F52073, a branching and pattern, shows frequent fasciculation. Fibres are anastomosing (irregularly reticulate) filamentous present in moderate quantity in relation to the alga. In F52075, irregularly oblong, occasionally matrix, and do not appear sparse and isolated as polygonal bodies with a wrinkled surface, 12-15 in Pseudoceratina. [Their distribution and cross- ^m long. Similar symbionts, but lobulate (like wal- section are irregular, the latter bulbous-lobate, in nuts or mulberries), 10-12 deep red, /tm long, D. rotunda also cavernous.] Pith is present to the F52076, with filamentous in permeate alga long exclusion of bark in the fibres, [is finely reticulate], strands. F52074 contains far fewer, scattered and sand/spicule fragments may be incorporated microsymbionts of mulberry-type, and smaller [in variable amounts]. The texture of the sponge ones, globular or cocoon-shaped (twinned), 4-5 pm is compressible [to firmly] rubbery; the surface is wide; anatomy thus clearly visible. Choanocyte markedly conulose, [or papillate and rugose]." chambers very conspicuous, large and crowded, (Modified from Bergquist, 1980b: 494.) irregularly elongate. Small ones often spherical, Remarks. Druinella is larger ones pear-shaped, flattened or polygonal reinstated here and in Wiedenmayer through crowding. Length 50-93 y.m, width 50-59 et al. (in press) as a valid senior syn- ^m. Extreme measurements 50 x 50 ^m, 54 x 93 onym of Psammaplysilla. AH authors after Len- denfeld, /*m. Larger canals, typically 200-400 /mi wide, including Bergquist (1965, 1980b) have overlooked the existence of the generally have thin septa, spaced 80-170 fim, com- holotype of the type species, D. rotunda monly annulate, also reticulate, occasionally in ves- Lendenfeld, in East Berlin (see below). This is cicular zone along canal walls. a good species, congeneric with, but distinct from Druinella purpurea. Remarks. Contrary to the statement in Bergquist Druinella rotunda Lendenfeld (1980b: 494), Carter's description of this species is not "quite clear", actually misleading. This and view Plate 17 figure 12, plate 38 figures 1-5 (see Wiedenmayer et al., in press, Annotated Bib- Druinella rotunda liography, under Bergquist, 1980b, and Checklist, Lendenfeld, 1889a: 47 (in monotypic new genus). -Lendenfeld, 1889b: pi. under Pseucloceratina durissima) has been con- 427, 34 figs 3, 10 (Sydney Harbour). -Wilson, 1925: 491 (in discussion). - firmed by Pulitzer-Finali (1982b: 139). It is fairly de Laubenfels, 1948: 98 (resume).- Bergquist, 1965: 138, obvious that Carter's description of microscopic 139 (discussion, holotype pronounced lost, unrecognisa- details was based on a specimen of the first form ble from original description). described above, and that he thus overlooked the ? Psammaplysilla purpurea. - Vacelet et al., 1976: 106 pithed fibres. (part: pi. 5 fig. e; Tulear Reef Complex, Madagascar). SPONGES FROM BASS STRAIT 159

Material examined. Station KG 5, one specimen (F52078). ule fragments. Same type of detritus in fibres. Type slide with five stained microtome sections, labelled Fibres extraordinarily lobose and folded in cross- by Lendenfeld (ZMB 10403). section, even more than in Lendenfeld (1889b, pi. Diagnosis. Irregularly lobate, with lobes com- 34 fig. 3). In fibres, pockets from marginal folds pressed, basally concrescent; or erect-cylindrical. and slits reach deep inside and often communicate axial of pith similar to Very firmly rubbery, little compressible. Externally in region. Fine meshwork that described and Bergquist dull yellow to mustard grading to dull red and by Wilson (1925) 4-5 brownish purple; internally bright lemon-yellow; (1965, fig. 6d), with meshsizes around fim. essentially as turning dark blue after collecting; externally pur- Anatomy of aquiferous system Lendenfeld (1889b, pi. 34 fig. 3, plish black, internally ochre in alcohol. Alcohol represented by 10 of same plate stained orange-brown, labels dark purple. Surface obviously diagrammatic). Figure Brien al., 1973: 205, fig. 170) prob- papillate and rugose. Oscules minute, scarce, scat- (redrawn in et ably phantasy, such details not apparent in type tered. Choanosome very dense, collagenous. Ecto- chambers some 1-1.5 mm thick. Thick, sparsely slide. Spherical to ellipsoidal choanocyte (17-31 /tm in greatest diameter) crowded and res- anastomosing, longitudinally fluted, cavernous tricted to meandering and sinuous areas (deeply fibres. Anatomy of choanosome: main canals of stained) between two systems of canals: (1) With two types, one (exhalant) with thick sheaths con- thin deeply stained lining (pinacoderm) and narrow, taining radial apochetes, delimiting meandering and lightly stained sheath, choanocyte chambers often convoluted fields of crowded choanocyte with pinacoderm. Lumina con- chambers. almost in contact tain regularly spaced transverse septa. Inhalant Description. Irregularly lobate, comprising four canals according to Lendenfeld. compressed and con- lobes, basally concrescent, (2) With thick deeply stained lining (prinacoderm torted mostly at random. Two lobes sprawling in and finely fibrous layer) and wide lightly stained opposite directions, lower face of one as base of sheath with radial structure. Walls of lumina much attachment. Width at base 4.5 x 8 cm, height 7 convolute (meandrine) in longitudinal sections. cm. Most of surface regularly papillate and fre- Exhalant canals fide Lendenfeld. Prosochetes, quently rugose, with rugae connecting neighbour- prosopyles, apochetes and apopyles not clearly visi- ing papillae (or blunt conules) in angular course. ble, very fine. Transverse sections through type 2 Papillae and rugae 1-2 mm wide, spacing 2-3 mm. suggest that apochetes get fewer and larger away Some interstices wider, smooth. from choanocyte chambers, virtually disappearing in Choanosome, in base, bright lemon yellow in deeply stained periphery of exhalant canals, life, turned dark blue, minutes after collecting. Sur- perhaps a sphincter. face was dull yellow to mustard, grading over dull Fibres mostly surrounded by sheaths of lighter red to brownish purple (7.5 YR 7/12 and 5 R stained, concentrically structured collagenous 5/6-10); uniformly purplish black in alcohol. Alco- materal; choanocyte chambers sometimes almost hol, even after change, orange-brown; labels inside at contact with fibres. jar and fingers upon handling stained dark purple. Remarks. The deeply lacunose fibres and the canal Very firmly rubbery, little compressible. Minute, are most distinctive for this species. The rather scarce, flush oscules distributed at random. system compressed-lobate shape of the new specimen may Surface, with low magnification, almost smooth, or not be characteristic. Lendenfeld (1889b: more often finely rugose, rugo-vermiculate or regu- may described his specimen (his consistent use of larly pitted. Ostia, presumably in depressions, not 427) his descriptions is probably mislead- detected. the plural in erect, cylindrical". This remains Choanosome in alcohol very dense, collagenous, ing) as "digitate, verified from a re-examination of the holo- ochraceous. Deep purple ectosome generally 1 to to be The papillae, or blunt, low conules of D. 1.5 mm thick. Numerous commensal barnacles type. more diagnostic, in contrast to the lodged below ectosome. Thick dendritic fibres with rotunda may be generally taller, pyramidal conules of D. purpurea. little colour contrast, longitudinal, often twisted specimen fugured in Vacelet et al. (1976, pi. fluting and slits leading to internal folds. The Psammaplysilla purpurea, is very simi- My sections too thick for anatomical details of 5 fig. e), as to the new specimen externally. The presence interstitial choanosome. But fibre section in all lar bristle-like fine radial canals surrounding prin- regards like those in Lendenfeld's slide from holo- of cipal canals is not unique. A similar feature was type, described below. As in type slide, interstially described by Kirkpatrick (1910) in a sponge from scattered mixed detritus, ill-sorted, including spic- .

160 F. WIEDENMAYER

St Helena, misidentified as Chondrosia plebeja, plete transect along the northern part of Bass Strait, possibly identical with Chondrosia colleclrix is fairly representative of this province. The most (Schmidt) (see Wiedenmayer, 1977a: 189). The only conspicuous gaps, in terms of the relative impor- in the other species in Druinella is D. purpurea (Carter) tance of families and genera missed, are (see Bergquist, 1965: 135, figs. 6a, b, for synonymy Suberitidae, Clionidae, Desmacellidae, Halichon- Crellidae, and Renieri- and description). It is encrusting, massive or repent- driidae, Hymeniacidonidae, particularly well ramose. It is distinct from D. rotunda by its colour dae. The horny sponges are (bright green or yellow in life, reddish brown in represented. No doubt the gaps reflect a very alcohol), the mostly pyramidal tall conules, more incomplete sampling of habitats. solid fibres with frequent anastomoses, and lack A discussion of possible causes of this high diver- of the peculiar aquiferous anatomy of D. rotunda. sity would have to consider the fossil record of siliceous sponges worldwide, and recent developments Notes on ecology in historical biogeography, particularly with rela- The overlap in species composition between the tion to other neritic benthic invertebrate groups, stations sampled is small. Dissimilarity of stations and to plate tectonics. This approach is far too in this respect is thought to reflect (a) high taxo- involved in the context of this report, and will be nomic diversity in the whole area (see below), and dealt with in a separate publication. However, some (b) high environmental differentiation. Environ- salient points are outlined below.

ments covered herein can be grouped as: 1 Characteristics of the sedimentary substrates (1) Rock substrate, mostly sloping to overhang- and hydrodynamic conditions on shelves have ing, with or without algal forest, with high diver- always been of the greatest importance to neritic sity overall and vertical zonation. Sponges below sponges. This applies to individuals, local and wider the algal forest (20-30 m) tend to be large. This populations, all phases of growth (life cycles), type of environment characterises the Kent Group assemblages and provincial faunas, and hierarchic and Wilsons Promontory. levels of time from seasonal to a few millions of

(2) Level bottom below 50 m depth, substrate years. The latter aspect, chiefly one of cycles, is coarse and shelly (BSS station 187). Diversity and thought to be particularly relevant throughout the proportion of endemic/stenotopic species are high. Cenozoic.

(3) Level bottom below 50 m depth, substrate 2. The prevalent bottom fades throughout the muddy sand and silt, with few shells, small and deli- shelf of southern Australia, from the south-west cate algae, occasionally chaetopterid worm tubes to eastern Victoria, and throughout the Cenozoic, (BSS stations 179-181). Diversity is low, with sandy has been that of calcarenites and calcirudites: cal- desmacidids (Stylotellinae) and widely dis- careous sand and rubble, consisting chiefly of inver- tributed/eurytopic species dominating. tebrate shells, tests, and their fragments, with various degrees of penecontemporaneous accretion Taxonomic diversily and cementation. In this trait, and in its extension Wiedenmayer et al. (in press) listed c. 430 valid and persistence, this shelf region is virtually unique. species (including Calcarea) described and recorded The dominant group of invertebrates having con- from Bass Strait, and from the coasts of Tasma- tributed debris to such sediments were and are nia and Victoria, i.e. 38% of the known Australian Bryozoa. These are followed in importance by shelf fauna. Extrapolating from percentages of new associated larger benthic foraminifers and coral- species (25%) and new records) (21 %) for the area line algae. in this work, the number of valid species to be 3. The long-term beneficial effects on popula- expected here is in the order of 600. The relatively tions and provincial faunas of sponges, in terms well known fauna of Calcarea of southern Austra- of taxonomic diversity and relatively high biomass, lia comprises c. 90 species. are not seen in the sedimentary facies alone, but The diversity, in terms of recorded species, also in populations of erect and ramose bryozoans. genera and families, equals or exceeds that of most The diversity and great proliferation of such provincial and regional faunas covered by recent bryozoans, forming thickets and biostromes, are reviews (Table 3). Only the numbers for the (chie- well documented throughout the present shelf of fly western) Mediterranean and for the West Indies southern Australia (Wass et al., 1970). There is a are larger, which doubtlessly reflects the more striking correlation between richness of bryozoans advanced knowledge of these faunas. and diversification/proliferation of siliceous Table 4 shows that the selection of material for sponges in the epicontinental Chalk Sea of Europe description in this report, though from an incom- in the Late Cretaceous, and in similar deposits, of SPONGES FROM BASS STRAIT 161

Table 3. Numbers of families, genera, and species recorded or estimated in (on the basis of) reviews and checklists of regional and provincial demosponge faunas (chiefly of shelves).

Region/Province Author/Reference No. of families No. of genera No. of species

Victoria and Tas- This 41 121 c. 340 mania (Maugean report/Wieden- (known), c. Province) mayer et al. (in 500 (expected) press)

Australia (main- Wiedenmayer et al. 54 227 c. 1040 land and Pacific (in press) islands)

West Indian Region Pulitzer-Finali 69*+ (50) 218* + (161) 561* + (410) (1986:193-208)

Mediterranean Pulitzer-Finali 61*+ (57) 197* + (169) 539 + (466) (chiefly western) (1983: 602-613)

Warm-temperate Topsent (1928), 52+ (40) 538+ (209) eastern Atlantic Levi and Vacelet (1958), Borojevic et al. (1968), Boury-Esnault and Lopes (1985)

Soviet Union, Koltun (1959, 28*+ (29) 78*+ (92) 247*+ (246) Arctic and Pacific 1966b) seas

Antarctic region Koltun (1966a: 7); 38 96 255 + and Kerguelen Boury-Esnault and Province van Beveren (1982)

Western Japan Hoshino (1981: 47) 28* (33) 75* (70) 205

New Zealand Bergquist (1968, c. 350 + 1970); Bergquist and Warne (1980); horny sponges and Poecilosclerida not yet published

New Caledonia Levi (1979: 307) 157

Brazil Hechtel (1976: 237, - 156 238)

*Not revised with respect to Wiedenmayer et al. (in press). + Includes archibenthal records neritic components in the case of Numbers in parentheses are adjusted for taxonomic revisions and Union, for Western Japan; Pulitzer-Finali (1983, 1986); for taxonomic revisions only for the Soviet and for neritic components only for the eastern Atlantic. .. .

162 F. WIEDENMAYER

Table 4. Families and genera of Demospongiae with 16. Desmoxyidae records in the Maugean Province, extracted from 35. Higginsia Wiedenmayer et al. (in press), with revisions from 36. Desmoxya this report. Asterisks denote new generic records 17. Trachycladidae for the province (in this report), crosses are for 37. Trachycladus + genera with species covered in this report. 18. Rhabderemiidae 38. Rhabdosigma

1. Plakinidae 19. Desmacellidae

1. Corticium* 39. Biemna 2. Geodiidae 40. Sigmaxine/la 2. Geodia* 41. Sigmaxia 3. Ancorinidae 20. Raspailiidae 3. Ancorina + 42. Raspailia 4. Stelletta + 43. Echinodietyum 5. Rhabdastrella* 44. Clathriodendron + 4. Pachastrellidae 45 Valedictyum 6. Pachamphilla 21. Halichondriidae 5. Coppatiidae 46. Ha/ichondria 1. Jaspis + 47. Ciocalypta 8. Asteropus 22. Hymeniacidonidae 6. Tetillidae 48. Hymeniacidon 9. Tetilla* 49. Stylinos 10. Cinachyra 50. Leucophloeus 11. Amphitethya 23. Mycalidae 7. Spirasigmidae 51. Mycale + 12. Trachygellius 52. Stylotrichophora 8. Tethyidae 53. Tetrapocillon 13. Tethya + 24. Desmacididae 14. Aaptos 54. Strongylacidon + 9. Polymastiidae 55. Pseudo-

1 5 Polymastia + halichondria 10. Suberitidae 56. Psammodoryx 16. Suberites 57. Burtonispongia 17. Prosuberites 58. Stylotella* 18. Rhizaxinella 59. Phoriospongia +

1 1. Spirastrellidae 60. Psammoelema + 19. Spirastrella + 25. Coelosphaeridae 12. Clionidae 61. Coelosphaera 20. Cliona 62. Amphiastrella 13. Chondrillidae 63. Fusifer 21. Chondrosia + 26. Crellidae 22. Chondrilla 64. CV-e/Za 14. Latrunculiidae 65. Pseudoclathria 23. Latrunculia + 27. Myxillinae 24. Sigmosceptrella 66. Myxilla 25. Negombata 67. Forcepia + 15. Axinellidae 68. Iotrochota 26. Axinella 69. Fibulia 27. Acanthella 70. Lissodendoryx + 28. Phakellia 71. Microtylotella 29. Pseudaxinella + 72. Ectyodoryx* 30. Phycopsis 73. Strongylamina

3 1 Ptilocaulis 74. Waldoschmittia 32. Reniochalina* 75. A earn us 33. Axinosia 28. Tedaniinae 34. Rhaphoxya + 76. Tedania + .

SPONGES FROM BASS STRAIT 163

77. Hemitedania 100. Hippospongia 78. Tedaniopsamma 101. Coscinoderma + 29. Anchinoidae 102. Leiosellay 79. Phorbas* 36. Thorectidae 80. Phimohalichondria 103. Ircina + 81. Ha/nigera 104. Hyrtios 82. Anaata 105. Sarcotragus 30. Microcionidae 106. Taonura 83. Clathriay 107. Thorectay 84. Thalysias 108. Psammocinia 85. Echinociathria + 109. Fasciospongia + 86. Echinochalina 110. Fenestraspongia 87. Paradoryx 37. Dysideidae 31. Renieridae 111. Dysidea + 88. Reniera 38. Darwinellidae 89. Haliclona 112. Ap/ysilla% 32. Callyspongiidae 113. Darwinellay

90. Callyspongia I 114. Dendrillay 91. Dactylia 115. Pleraplysilla 33. Niphatidae 116. Chelonaplysilla 92. Amphimedon 39. Dictyodendrillidae 93 Cribrochalina 117. Dictyodendrilla 94. Gelliodes + 40. Aplysinidae 95. Hoplochalina 118. AplysinaVs 96. Microxina 41. Druinellidae

34. Oceanapiidae 1 19. Pseudoceratina + 97. Oceanapia + 120. DruinellaVi 35. Spongiidae 42. lanthellidae 98. Spongia + 121. Bajalus 99. Carteriospongia +

late Eocene age, in the south-eastern United States. 5. The open-marine shelf faunas of southern the There is indirect evidence for analogous dependence Australia to the west of Bass Strait and along in isolated sponge spicules and bryozoans being coast of New South Wales had long independent common in Tertiary deposits of southern Australia. histories before the Late Eocene, when Tasmania 4. The fossil record of siliceous sponges in the began to separate from Antarctica, with the South late Mesozoic and Cenozoic, at intermediate lati- Tasman Rise subsiding. Exchange between the two tudes, shows that advancing seas (transgressions on old provincial faunas is likely to have occurred con- shelves and in epicontinental seas) fostered diver- tinuously since the Late Eocene or Oligocene (see fauna sification and proliferation, particularly when below). The richness of the Maugean sponge associated with latitudinal warming and low to probably reached a peak in the Middle Miocene. moderate sedimentation rates within well aerated The present fauna of Bass Strait, somewhat deci- be regarded as terrigenous-clastic bottom facies of normal salin- mated by subsequent cooling, can largely, if not wholly, relictic from this peak. ity (glauconitic sands and silts), locally also with clastic vulcanites. There are obvious parallels, in such effects, between sponges and other benthic neritic invertebrates, particularly with larger Zoogeography foraminifers. Such conditions were common in southern Australia between Late Eocene and affinity and degree of presumed Middle Miocene, with such clastic facies often adja- Zoogeographic endemism on the species level are apparent from cent to, and alternating with, calcarenites, and Accordingly (discounting doubtful when two successive, protracted warming pulses Table 5. is proportion of endemics allowed warm-water (Tethyan) faunas to migrate records) 58% the to southern Australia, while 39°/o is endemic to the to southern Australia. Mixture with endemic aus- to diver- Maugean Province. Species with Indo-Pacific tral faunas is thought to have contributed records account for 33% (23% if species with sification of the latter. )

164 F. WIEDENMAYER

Table 5. Zoogeographic distribution of the species described in this Report. Question marks are for doubtful identifications. New records in Area D2 (Maugean Province in Knox, 1960) are circled. Following Ekman (1953) and Veron (1974), the boundary between the northern and southern Australian faunas is here taken to coincide with the southern limits to the distribution of hermatypic corals, i.e. at about 29°S on the west coast, and at about 30°-31° on the east coast. The northern fauna in WA thus includes that of Abrolhos Islands (Dendy and Frederick, 1924) and part of the records in Hentschel (1909-1911: Shark Bay to Geraldton). In NSW the Solitary Islands and Lord Howe Island constitute the southern bastion of coral reefs, but their sponge faunas are unknown.

Columns are numbered as follows:

1. West Indian Region 10. Southern Australia

2. North Atlantic 11. W. Pacific (Great Barrier Reef for Area C)

3. East-Central Atlantic 12. Japan

4. South Atlantic 13. New Zealand

5. Mediterranean 14. Central Pacific

6. Red Sea 15. N. Pacific

7. Indian Ocean (plus Natal Coast) 16. E. Pacific

8. Indo-Malayan Region (WA for Area C) 17. Subantarctic

9. Northern Australia 18. Antarctic

7 8 10 11 12 13 14 15 16 17 \i

A. Tethyan distribution (West Indies to Pacific)

Corticium candelabrum X X X X X X 55 X Tedania anhelans X X X X X X X X Lissodendoryx isodictyalis X X X X X X X X X Phorbas tenacior X 7 Callyspongia pergamentacea X X X X Dysidea a vara X X X X X X Aplysilla rosea X X X X X (8 X Darwinella australiensis X X X X X X Darwinella gardineri X X X ®

B . With Indo-Pacific records Jaspis stellifera X X X Tethya ingalli X X X X X Spirastrella papillosa X X

Chondrosia reticulata X 7 Trachycladus laevispirulifer X X X Echinoclathria laminaefavosa X X Strongylacidon stelliderma X X Phoriospongia kirki X X X X

Mycale mirabilis X X X 7 Callyspongia diffusa X X X X X X Callyspongia ramosa X X X X X Callyspongia serpentina X X Gelliodes incrustans X X ® X Oceanapia putridosa 7 7 ? X Spongia hispida 7 X X X (8 X X Carteriospongia caliciformis X X

Carteriospongia (c . f vermicularis 7 . s Leiosella caliculata X X X X Dendrilla cactos X X X SPONGES FROM BASS STRAIT 165

9 10 11 12 13 14 15 16 17

C. With northern Australia records Geodia punctata X 8 Clathria transiens X X X Echinoclathria leporina X X X Stylotella inaequalis X X

Phoriospongia carcinophila X (8 X Psammoclema densum X X X Callyspongia bilamellata X X X

Carteriospongia silicata X <8> X

Fasciospongia rimosa X <8> X Pseudoceratina durissima X X X

D. Southern Australia

D 1 .With records in SA, WA, NSW. Ancorina robusta X Echinoclathria carteri X Echinoclathria globosa X Echinoclathria tubulosa X Forcepia biceps X Ectyodoryx maculata X Coscinoderma pesleonis X Leiosella levis X Ircinia caliculata X Thorecta choanoides X Aplysina ianthelliformis X Aplysina lendenfeldi X Druinella rotunda X

D 2.Vic. -Bass Strait-Tas. coast only Ancorina geodides X Polymastia crassa X Latrunculia conulosa X Rhaphoxya cactiformis X Clathriodendron cacticutis X Echinoclathria favus X Phoriospongia argentea X Psammoclema callosum X Psammoclema nodosum X Psammoclema ramosum X Callyspongia asparagus X Thorecta tuberculatus

D3 .Single locality only (new species) Ancorina repens X Ancorina suina X X Stelletta arenitecta Rhabdastrella cordata X Rhabdastrella intermedia X X Tetilla praecipua X Latrunculia hallmanni X Pseudaxinella decipiens X Reniochalina sectilis X Rhaphoxya felina X Clathria wilsoni X Echinoclathria egena m

166 i-. wihdlnmayi-k

10 11 12 13 14 15 16 17

I'summoilenw hilextum x Psatnmoclejtia fissuratum x Psammoclema goniodes X

Psammoclema radial it X Psammoclema stipitatum X Psammoclema vansoesti X Callyspongla toxifertt X Callyspongia persculpta X Thorecta glomerosus X

Telhyan distribution are not counted). The oxer- This unusual provinciality in sponges is perhaps lap with northern Australia (essentially the Solan- best explained as relictic palaeoaustral, originating derian Province) comprises 20%. This reflects a in southern Australia in the Cretaceous to Early strong Indo-West Pacific affinity of the sponge Tertiary, and shaped further in the Eocene- fauna of our area. The affinity is strongest to the Oligocene by the final breakup of Gondwana (Zins- faunas of the Indian Ocean and Indo-Malayan meister, 1982). It is unlikely that there was one Region. If northern Australia is included in this centre of origin for all these genera. According to group, and the Telhyan element is discounted, the Woodburn and Zinsmeister (1984: 938) rifting overlap is 26%. The overlap with the West Pacific between Australia and Antarctica probably began (without Tethyan records, but including New much earlier in the south-western part than pre-

Zealand) is 13% (10% with New Zealand alone). viously thought, namely in the Late Jurassic, and The Tethyan element (equals "cosmopolitan" of deep-water conditions may have extended close to

many authors) comprises 1 1%. This is probably a Bass Strait by the I, ate Cretaceous (Campanian). fairly constant proportion for all tropical to tem- This arm of the Tethys or Indo-West Pacific Region perate sponge faunas. The corresponding figure (roughly similar to the Red Sea in its initial phase) over all three zoogeographic tables for West Indian is known as the Austral-Indo-Pacific Province species in van Soest (1978, 1980, 1984) is 13%. (Wilson and Allen, 1987: 45). Another likely centre Interestingly the highest overlap of our collection of origin for endemic demosponge genera is the and this group is with the Mediterranean fauna shelf east and north of Bass Strait, as part of the (10% in contrast to only 4% in van Soest). Weddellian Province (Zinsmeister, 1982). Follow- Except for the widely distributed Aplysilla rosea, ing the detachment of Tasmania and the South no species (with the possible exception of Tasman Rise from Antarctica in the Late Eocene, Callyspongia serpentina) has Antarctic/Sub- some mixture of neritic faunal elements between antarctic records. Such an overlap is extremely the two provinces might be expected to have small with all the species from mainland Australia occurred in south-eastern Australia. Several listed by Wiedenrnayer et al. (in press). brachiopod genera common in south-western Aus- There are at least 12 well known genera recorded tralia in the Palaeocene appeared in south-eastern from the southern Australian Region, which can Australia in the Eocene (Woodburn and Zins- be regarded as endemic in a wider sense. These are meister, 1984: 938, quoting McNamara). Such Renioehalina, Rliaphoxya, Trachycludus, Clathrio- exchanges probably continued during the Late and in later dendron, Stylotella , Psammoclema, Oligocene (Janjukian) transgression Echinocfathria, l.eiosella, *Taonura, Thorecta, stages of high sea levels. * Fenestraspongia, and *Aplysinopsis (* not Evidence from other benthic groups with good represented in the collection of this report). There fossil records, particularly molluscs (Darragh, 1985; have been occasional records from New Zealand, Wilson and Allen, 1987), suggests some modifica- from northern Australia, and from the whole Indo- tions to the above view. The degree of generic ende- West Pacific Region. Those from tropical waters, mism in the South-east Australian Province was if confirmed by modern revisions, are too isolated already high in the Late Eocene (24%), and to suggest any other explanation than southern increased substantially (to 34%) by the Pleistocene. Australian origin for these genera. This particularly The South-east Australian fauna was probably applies to Psammoclema, Echinoclathria, always a temperate assemblage, though its Tethyan Leiosella, Taonura and Thorecla, because of their or Indo-West Pacific component reached 30% in diversity and because of their pronounced habit to the mid-Miocene. The Austral-lndo-Pacific incorporate sand. Province was richer in Tethyan/Indo-West Pacific SPONGES FROM BASS STRAIT 167 elements, and decidedly subtropical to tropical, up Cribrochalina Schmidt to early Pleistocene time. Already in the late Early Oceanapia fistulosa (Bowerbank) Miocene, the South-east Australian Province Pellina carbonaria (Lamarck) expanded westward, and throughout the Neogene IGelliodes Ridley (one species recorded in had a sharp boundary with the Austral-lndo- Mediterranean) Pacific Province, just east of St Vincent Gulf. The Amphimedon viridis Duchassaing and Michelotti South-eastern Province expanded further westward ICallyspongia Duchassaing and Michellotti (one in the Pleistocene. endemic species in Mediterranean) The high proportions of lndo-West Pacific Callyspongia pergamentacea (Ridley) genera and species in the Maugean demosponge Hyattella Lendenfeld fauna nevertheless call for some caution in apply- ICoscinoderma (West-Indian records doubtful) ing molluscan biogeography. Unfortunately, apart Group of Iricinia strobilina (Lamarck), includ- from the lacking fossil record, the living ing /. irregularis (Polejaeff) and /. echinaia Keller demosponge faunas of south-western Australia and (see Vacelet et al., 1976: 102) of New South Wales are not sufficiently known to Smenospongia Wiedenmayer resolve this question. Dictyodendrilla Bergquist Phonospongia is another sandy genus of con- Aplysina lendenfeldi Bergquist and A. pedun- siderable diversity and of common occurrence in culata (Levi) southern Australia, which might be added to the Pseudoceratina Carter endemic group of genera. It has a single well- documented record in the Tulear Reef (Phoriospon- This distribution is best explained with the migra- gia lamella Lendenfeld in Vacelet et al., 1976: 63), tion of elements of the palaeoaustral/Indian Ocean and has a curious extension into the West-Indian fauna around South Africa following the opening Region, w ith four endemic species. Three of these of the South Atlantic in the Lower Cretaceous, and were described from Brazil by Boury-Esnault the rifting between southern and central Atlantic (1973a: 283), in Psammochela and Psamnwtoxa, in the Turonian, lasting as long as favourable cli- and the fourth West-Indian species is "Holop- matic and oceanographic conditions persisted, i.e. samma" helwigi de Laubenfels, redescribed by probably until Mid-Miocene time. Reyment (1973) Pulitzer-Finali (1986:147). and Scheibnerova (1978) have argued that the There is a number of genera, species pairs and South Atlantic housed a uniform zoogeographic groups, and species with a similar distribution, i.e. province, which formed part of the Austral Realm with records in Australia (in part only northern up to lower Turonian time, and was until then dis- Australia), the lndo-West Pacific Region, the tinct from the western Tethys/Caribbean Province. southern and central Atlantic (particularly Wesi The strong influence from the southern Indian Indies), but not in the north-eastern Atlantic, Arctic Ocean and Australia then extended to Nigeria and and Mediterranean. These are listed below: north-eastern Brazil (Recife), and still extended to Stelletta debilis Thiele the Sierra Leone Rise (6°N) in the Late Cretaceous. Rhabdastrella Thiele From then on mixing of faunas between central and Asteropus simplex (Carter) South Atlantic occurred. Spirastrella spinispirulifera (Carter) For neritic sponges, the rises and seamounts of Cliona carteri (Ridley) the South Atlantic probably played an important Cliona carpenter! Hancock role in this exchange, as the faunule (28 species) Tethya seychellensis (Wright) from the Vema Seamount described by Levi (1969) Tethya japonica Sollas imples. Although specific endemism is high (54%) Ptilocaulis spiculifer (Lamarck) the affinities of species are clearly mixed between Ectyoplasia Topsent North Atlantic/Mediterranean, South African, and Mycale (Arenochalina) mirabilis Lendenfeld and Indian Ocean/Australian, though without typical elements. There are close ties between M. (A.) laxissima (Duchassaing and Michelotti) West Indian Neofibularia Hechtel four of Levi's new species and southern Australian Desmapsamma anchorata (Carter) ones: Paresperella curvisigma Levi/P. repens Acarnus innominalus Gray Whitelegge, Mycale diastrochela Levi/M. obscura Rhaphidophlus juniperinus (Lamarck) (the genus (Carter), Spongia violacea Levi/ Hippospongia in the anfractuosa (Carter), and Aplysina pedunculate is only represented by R. jolicoeuri Topsent mentioned. Mediterranean) Levi/A. lendenfeldi, as already Only nine genera in the present collection (19%) Strongyiacidon I enden feld Xestospongia testudinaria (Lamarck) and X. are characterised by circumtropical to warm- (chiefly West Indies, Mediter- muta (Schmidt) terpcrate distribution I6S F. WIEDENMAYER

ranean, and Indo-West Pacific Region). These are siderable number of warm-water, neritic inver- Corticium, Ancorina, Geodia, Spirastrella, Chon- tebrate taxa to colonise the entire shelf of southern droma, Gelliodes, Oceanapia, Fasciospongia, and Australia. This is well documented in groups such Darwinella. as molluscs (Wilson and Allen, 1987), larger The last group of genera that can be distin- foraminifcrs, brachiopods, bryozoans, and, among

guished in our collection, is that of 20 (43%) with sponges, by lithistids and inozoans (pharetronids cosmopolitan distribution (Tcthyan with extensions of authors). In terms of density (biomass), into either boreal or antiboreal waters, or both): bryozoans, coralline algae, foraminifers, molluscs Stelletta, Jaspis, Tetilla, Tethya*, Polymastia, (gastropods, bivalves), and brachiopods were most

Latrunculia, Pseudaxinella* , Clathria, Mycale, important among the immigrants, resulting in the Tedania, Forcepia*, Lissodendoryx, Ectyodoryx, wide distribution of shelly-bioclastic (calcarenitic- Phorbas, Spongia, Jrcinia, Dysidea, Aplysilla, Den- ruditic) facies, with its beneficial effect on sponges, drilla, and Aplysitui. as already mentioned. Besides increasing water tem- The three genera marked by asterisks have no peratures, changing oceanographic patterns in the Antarctic/Subantarctic records. Oligocene and Miocene (see below, under 4), par-

The Antarctic fauna is characterised by the rarity ticularly the heavy influx of surface currents from or absence of members of the Ancoronidae, Geodi- the eastern Indian Ocean (indicated by carbonate

idae, and Tethyidae, and of horny sponges. It could sedimentation) must have aided this mixing and be argued that most of the Antarctic records are augmentation of benthos, both by oceanic trans- archibenthal and therefore not suitable for com- port of dissemules, and by neritic range extensions. parison with the shelf-fauna of southern Austra- The latter avenue was probably substantially com- lia. The numerous records from depths less than plemented in the Miocene by the system of ridges 100 m at Kerguelen, Heard and Macquarie Islands and plateaus between India and south-west Aus- (Koltun, 1976; Boury-Esnault and van Beveren, tralia (Cook, 1977: 146, Fig. 13F). Furthermore, 1982) do not support such a view, nor does the the peripheral influence of the Circum-Antarctic shallow-water fauna of Tierra del Fuego described Current and of upwelling deeper water rich in dis- by Sara (1978). solved silica and nutrients is assumed to have Some questions relating to these affinities and influenced the proliferation of the southern Aus- to the high degree of specific endemism of the Mau- tralian demosponge fauna. gean demosponge fauna remain to be answered. 2. Affinity with Mediterranean fauna. It is chiefly The most relevant ones can be formulated as this affinity which has prompted the use of the term follows: (relictic) Tethyan in the context of Table 5. For 1. Why is the affinity with the faunas of the neritic taxa, the term is largely synonymous with Indian Ocean and of the Indo-Malayan Region so "cosmopolitan" of most authors. strong? There are at least 35 such cases of species, or 2. Why is the affinity with the Mediterranean groups of closely related species, to be expected in fauna unexpectedly high, with several species in the Australian neritic demosponge fauna, mostly common or closely related? with records in the south. About two-thirds of these 3. Why is the affinity with the fauna of New have West-Indian records; several also have records Zealand so weak? in New Zealand, and some have apparently widely 4. Why is the affinity with the Antarctic and disjunct distribution, without records in the Indian Subantarctic faunas almost nonexistent? Ocean, or Atlantic only, not Mediterranean. 5. How did the the "sandy sponges", a very Sponges share with some other groups of sessile important sector (in diversity and density) of the invertebrates their very poor larval dispersal endemic genera of southern Australia, originate? capa- bilities, and the seeming paradox of having very Again, attempts to reasonably answer these ques- large geographic ranges in many species, nonethe- tions, with adequate discussions, will have to await less. Scheltema (1986: 311) argued that such separate publication. But preliminary suggestions species are commonly members of the fouling are given below, with a minimum of references to community, have widely disjunct distribution, the pertinent literature. mostly restricted to harbours, and that this distri- /. Affinity with faunas of Indian Ocean and lndo- bution is best explained with shipping. Malayan Region. This question has been partly Several arguments can be advanced against this answered above. Protracted episodes of warming view. Probably the most important one is that at high latitudes in the Palaeocene to Late Eocene, Scheltema did not consider well studied fossils, for and in the Early to Middle Miocene allowed a con- instance the geographic range of some species of SPONGES FROM BASS STRAIT 169

Numnutlites and related genera, the larger lar to coeval ones in the western tropical Pacific foraminifers dominating the Eocene Tethys, from and Indo-Malayan Region. western Europe as far east as the western Pacific 4. Affinity with Antarctic and Subantarctic faunas. (Premoli-Silva and Brusa, 1981). Related to this The Circum- Antarctic Current, and oceanic con- argument is the probability of non-larval trans- ditions between Australia and Antarctica, were oceanic transport of sessile shallow-water benthos fully established in the Late Oligocene, following by rafting on algae, sea grass and mangrove root the subsidence of the South Tasman Rise into fragments throughout the Cenozoic (Brasier, 1975; deeper water, and the opening of Drake Passage. McCoy and Heck, 1976; Eva, 1980), which is a In the Late Eocene there was a continuum in faunas much more viable alternative to the shipping and floras between the subtropical belt along hypothesis. southern Australia and New Zealand, and the By analogy with molluscs and scleractinian warm-temperate zone around Antarctica. The corals, it is very likely that the species and species- Circum-Antarctic Current had the effect of "lock- groups in question are at least as old as Early Mio- ing in" the Antarctic and Subantarctic fauna, fully- cene, if not Oligocene. The closure of the Mediter- exposing it to increasing cooling, and contributing ranean in the Late Miocene would thus have had to its present uniformity. From the Early Oligo- little if any effect on geographic ranges of these cene on, the southern end of Tasmania moved to species. the north, away from the 60" parallel. The bound- 3. Affinity with fauna of New Zealand. The ary between cold surface waters of the Subantarc- microcontinent of New Zealand (with the plateaus tic Zone and the warmer waters over the southern and rises adjoining the present islands) began to Australian shelf, continental slope, and Great Bight separate from south-eastern Australia in the Late Abyssal Plain became sharp in the Middle to Late Cretaceous (Late Campanian). Seafloor-spreading Miocene. was fast, and the Tasman Sea had reached its The separation of biogeographic provinces at this present width at the end of the Palaeocene (c. 56 time was partly due to the steep temperature Mybp). There was no spreading later, except fur- gradients and partly to a threshold related to ther south (Owen, 1983). phytoplankton and thus to different trophic sys- According to Kauffman (1973: Fig. 5, p. 362f.), tems. This is indicated by the asymmetrical the degree of endemism of the bivalve fauna of distribution (with relation to the axis of New Zealand was weak in the Early Cretaceous, seafloor-spreading along the crest of the South-east but increased markedly in the Cenomanian, sup- Indian Ridge) of, and sharp boundaries (vertical posedly in relation to major marine flooding during and horizontal) between, biogenic fine sediments global transgression at this time. This strong ende- on the floor of the Southern Ocean. These were mism was conserved to the end of the Cretaceous supplied by phytoplankton in superficial waters: and seems to have fluctuated in the Tertiary. Zins- siliceous (chiefly diatoms), in the southern portions, meister (in Woodburn and Zinsmeister, 1984: 938) and calcareous (nannoplankton, coccolithophorids) found little similarity, in shallow-water mollusc adjoining the southern Australian shelf. The biosi- faunas of the Late Cretaceous and Palaeocene, liceous pelagic fades is largely coextensive with ice- between south-eastern Australia and New Zealand. rafted detritus. West of Tasmania, the pelagic sedi- Darragh (1985) found that 30% of the molluscan ment in lower bathyal and abyssal areas tend to be genera in the early Tertiary of the South-east Aus- exclusively calcareous throughout the Miocene, tralian Province occurred in the coeval fauna of north of the South-east Indian Ridge. The crest of New Zealand, and that this proportion dropped to this ridge then formed the boundary with the biosi- 17% by the Pliocene and Pleistocene. liceous facies to the south, and only in the Early Though no such data are available for neritic Pliocene did this boundary shift to the northern et al., 1975). The sedimen- sponges, it can be speculated that this sector of the flank of the rise (Kemp New Zealand fauna had a similar evolution. There tary cover of the South Tasman Rise is known to entirely of pelagic carbonates, after the are also signs, however, of strong influx of warm- consist water taxa from the north, throughout New Oligocene. The present Antarctic Convergence (Polar Front) Zealand, in the Late Eocene and Early Oligocene, approximately coincides with the pelagic facies just as in southern Australia: The diverse spicule boundary along the northern flank of the South- assemblage in the Oamaru Diatomitc (south-eastern east Indian Ridge south of Australia, until off the coast of southern New Zealand), described by southern end of the South Tasman Rise. Further Hinde and Holmes (1892), is associated with a rich it turns southward, the again eastward, skirt- assemblage of larger neritic foraminifers very simi- east 170 F. W1EDENMAYKR

is in regulat- ing the Campbell Plateau. This boundary is thought The benefit by uptake of sand seen to have been installed south of the Campbell ing specific gravity to counter mobility of the sub- Plateau already in the earliest Oligocene. The strate and thus to facilitate recruitment and early sedimentary cover of the Campbell Plateau at its growth. Optimal density, once attained, would be southern end (straligraphically continuous only in conserved in adult sponges because it would offer the Palaeogene) consists entirely of calcareous further advantages in propagation. In the case of pelagics. The steepening of temperature gradients larval settling and recruitment, weak and more or of surface waters in the New Zealand region less permanent substrate mobility, at normal wave occurred gradually throughout the Oligocene and base and within the range of tidal currents, would Early Miocene, and the present oceanographic con- be the chief controlling factor (Wiedenmayer, figuration (Subtropical Convergence) developed 1978). In adults of the same sponges, rolling and during the Middle Miocene (Murphy and Kennett, fragmentation by storm-wave action could be 1986; Kennett and von der Borch, 1986). exploited to promote propagation and controlled Recent syntheses in palaeoceanography thus sup- range extension (fades tracking) (Battershill and port the notion that the neritic benethic faunas of Bcrgquist, 1985). the Maugean Province and of New Zealand, Parallels can be seen with the life cycles and sur- though having evolved independently since the Late vival strategies of fast growing, porous, branch- Cretaceous, were subjected to similar climatic con- ing scleractinians in shallow water, particularly ditions. In both provinces, the strong influx of Acroporidae (Jackson and Hughes, 1985); but also warm-water taxa in the Late Eocene was largely with lithistid demosponges (having thrived in shelly- conserved by gradual adaptation to falling temper- calcarenitic fades since the Middle Cambrian), atures, and by regional permanence of suitable where the interlocking main skeleton should not sedimentary substrates. be seen merely as strengthening of the adult sponge body in fixed position. 5. Sandy sponges. A discussion of this question would have to consider incorporation of sand by Acknowledgements sponges in general, and thus chiefly aspects of life cycles and functional morphology. A starting point The research work at the Museum of Victoria

would be I he only paper wholly dedicated to the which led to this account was funded by Marine phenomenon of sand in sponges, that of Shaw Sciences and Technologies Grant 80/0267. Aus-

(1927b). It seems doubtful that much could be tralian Biological Resources Study Grant 82/1643 added without detailed field observations and con- contributed to travel and expenses during my trolled experiments. survey of type collections in London (BMNH), East

Uptake of sand occurs in many species and Berlin (ZMB) and Jena (PMJ) in 1983. I owe it to genera, distributed among many families and Dr Barry R. Wilson, then Director of the National orders of demosponges, occasionally also in Cal- Museum of Victoria, who devised and formulated carea. The proportions of sand to whole sponges, the project, together with Dr Brian J. Smith, for

and distribution with respect to skeleton, ectosome enabling me to participate. 1 deeply appreciate his and choanosome are often variable. In some spe- initial hospitality, constant encouragement, and cies, these proportions seem to be regular, while help in the final stage of my sojourn in Melbourne. in others the uptake of sand is obviously Dr B. J. Smith, Senior Curator and Head of the facultative. Division of Zoology at the Museum of Victoria was We have few data on the biogeographic sig- the Chief Investigator in the MST-Project and sup- nificance of the phenomenon and on the role of ported my participation in all its stages. evolution, since the fossil record tells us nothing Peter Forsyth, Bronwen J. Scott, Marilyn Olliff, in this respect. Natalie Schroeder, Dr Elizabeth Robertson, Ger- As a preliminary hypothesis, it can be speculated aldine James, and Peter Myroniuk did the necess- that the strong and obligatory content of sand in ary laboratory work (microslides). Bronwen J. many species and genera endemic to (southern) Scott did most of the camera lucida drawings (spic- Australia (e.g. Echinoclalhria, Stylotellinae, several ules and skeletal features) for the text-figures. John Thorectidae) may be the result of the prevalence, Coyne was helpful in photomicrography. Frank over long periods, of sandy substrates in this shelf Coffa and his assistants in the Photography region, of associated hydrodynamics, and of adap- Department provided countless enlargements for tive response by particular taxa, presumably since the plates in this work and for the Photofile (Pho- the Palaeocene (the time when the calcarenitic tographs of type specimens in Sydney, London and bryozoan-algal fades appeared in the north-west East Germany). The staff of the Natural History and gradually spread along the southern shelf). Library, particularly Lily Stefani, was instrumen- +

SPONGES FROM BASS STRAIT 171

tal with the many interlibrary loans. Judith Odgers, Battershill, C.N., and Bergquist, P.R., 1985. The Biomedical Library, Monash University, provided influence of storms on asexual reproduction, recruit- a very useful complement to the Annotated Bib- ment and survivorship of sponges. Abstract, Third International on the Biology Sponges, liography in form of references and abstracts of Conference of Woods Hole, Massachusetts (to be published in the recent papers (1970-82) obtained through a com- Proceedings). puter search. Ms Joan Dixon (Department of Mam- Bergquist, P.R., 1961a. A collection of Porifera from mals) graciously lent me her compound microscope northern New Zealand, with descriptions of seven- for most of my stay in Melbourne. A special grant teen new species. Pac. Sci. 15: 33-48. from the NMV Council made ancillary equipment Bergquist, P.R., 1961b. Demospongiae (Porifera) of the for this microscope available. Lisa Giuliani was Chatham Islands and Chatham Rise, collected by the always patient and reliable in typing the Chatham Islands 1954 Expedition. Bull. N.Z. Dep. manuscript. Sci. hid. Res. 139: 169-206. Bergquist, P.R., 1961c. The Keratosa (Porifera) collected During field work in 1981 I received assistance "by the Chatham Islands 1954 Expedition. Bull. N.Z. in collecting from P.E. Bock, S.A. Shepherd, D. Dep. Sci. Ind. Res. 139: 207-219. Staples, R. Kuiter, (Kent Group) and R. J. Carey Bergquist, P.R., 1965. The sponges of Micronesia, Part survey Wilson (R.V. Tangaroa). During my of type 1. The Palau Archipelago. Pac. Sci. 19: 123-204. collections in Sydney, help and hospitality were Bergquist, P.R., 1967. Additions to the sponge fauna of offered by Dr J.W. Pickett and Dr F.W.E. Rowe the Hawaiian Islands. Micronesica 3: 159-174, 2 pis. Zealand: (Curator in charge, AM). For similar favours, 1 am Bergquist, P.R., 1968. The marine fauna of New indebted to Miss Shirley M. Stone and Dr D. Porifera, Demospongiae, Part 1 (Tetractinomorpha George (BMNH), Dr and Mrs D. Kuhlmann (ZMB, and L ithistida). Bull. N.Z. Dep. Sci. Ind. Res. 188 (Mem. N.Z. Oceanogr. Inst. 37): 105 pp., 15 pis. ( East Berlin), and Dr D. von Knorre (PMJ, Jena). fsp. in colour). Additional information on type collections (mostly Bergquist, P.R., 1969 Shallow water Demospongiae from was obtained from Dr C. through correspondence) Heron Island. Univ. Qd. Pap. Ot. Barrier Reef J.A. Devidts and G. Hild- Levi (MNHP, Paris), Comm. Heron Isl. Res. Stn. 1 (4): 63-72, 2 pis. wein (MZUS, Strasburg), Dr E. Kreissl (MJG, Bergquist P.R., 1970. The marine fauna of New Zealand: Graz), Dr M. Dzwillo (ZMH, Hamburg), Dr P. Porifera, Demospongiae, Part 2 (Axinellida and Bull. N.Z. Dep. Sci. Ind. Res. 197 Kuenzer (Gottingen), Dr I. Hasenfuss (Erlangen), Halichondrida). 85 20 pis. + Dr H. Fechter (ZSM, Munich), and Dr M. Grass- (Mem. N.Z. Oceanogr. Inst. 51): pp., ( hoff (SMF, Frankfurt/M). fsp.). Bergquist, P.R., 1978. Sponges. London: Hutchinson. To all persons and institutions mentioned, and 268 pp., 12 pis. to those that I might have forgotten, I express my Bergquist, P.R., 1980b. A revision of the supraspecific deep gratitude. classification of the orders Dictyoceratida, Den- droceratida and Verongida (Class Demospongiae). N.Z. J. Zool. 7: 443-503, 25 figs (pis). References Bergquist, P.R., Hofheinz, W. and Oesterhelt, G., 1980. and the classification of the Some alphabetic suffixes with years of publication may Sterol composition Biochem. Syst. Ecol. 8: 423-435. seem superfluous, and with olhers the sequence is Demospongiae. and Sinclair, M.E., 1973. Seasonal vari- deliberately discontinuous. This is to facilitate corre- Bergquist, P.R. settlement and spiculation of sponge larvae. lation with the more complete annotated bibliography ation in Biol. (Berlin) 20 (1): 35-44. (Wiedenmayer et al., in press). Mar. of Bergquist, P.R. and Skinner, I.G., 1982. Sponges Ali, M.A., 1956a. Addition to the sponge fauna (Phylum Porifera). Chapter 3, pp. 38-72, pis 1-7 in Madras. J. Madras Univ. (B) 26: 289-301. sponge Shepherd, S.A. and Thomas, l.M. (eds), Marine Ali, M.A., 1956b. Development of the monaxonid Invertebrates Southern Australia. D.J. Woolman: Lissodendorvx similis Thiele. J. Madras Univ. (B) of Adelaide. 491 pp., 32 pis. 26: 553-581. P.R. and Warne, K.P., 1980. The marine Annandale, N., 1915a. Indian boring sponges ol the Bergquist, "fauna of New Zealand: Porifera, Demospongiae, family Clionidae. Rec. Indian Mus. 11: 1-24, pi. 1. Part 3 (Haplosclerida and Nepheliospongida). Mem. Annandale, N., 1915c. Some sponges parasitic on Clion- N.Z. Oceanogr. Inst. 87: 77 pp. 17 pis. idae with further notes on that family. Rec. Indian Bocage, J.V. Barboza du, 1869. Eponges siliceuses nou- Mus. 11: 457-478, pi. 34. Kapstadt und vellcs de Portugal et de Hie de Saint-lago (archipel Baer, L„ 1906. Silicispongien von Sansibar, du Cap-vert). J. Acad. Sci. Lisboa 2: 159-161, pis Papeete. Arch. Naturg. 72: 1-32, pis 1-5. of 10-11. Bakus, G.J., 1966. Marine poeciloscleridan sponges Zool. Borojevic, R., Cabioch, L. and Levi, C, 1968. Inven- the San Juan Archipelago, Washington. J. taire de la faune marine de Roscoff. Spongiaires. Edi- Lond. 149: 415-531, 2 pis. de quel- tions de la Station Biologique de Roscoff, nouvelle Barrois, C, 1876. Memoire sur l'embryologie 3(11): serie. Cahicrs de Biologie marine: Paris. 44 pp. ques Eponges de la Manche. Ann. Sci. Nat. (6) Bosraug, E., 1913. Die Tetractinelliden. Pp. 231-251, 4 1-84, pis 12-16. .

172 F. WII'DPNMAYPR

pis in Voclt/kow, A., Reise in Ostafrika in den Jahren Uricn, P., Levi, C, Sara, M., I u/cl, O. and Vaeelet, J.,

}$03 1905 rail Milk-In der Hermann mid Eltse geb. 1973. Spongiaires. Vol. 3(1), 716 pp. in Giassc, P. Heckmann Wenlzel-Stiftung ausgefiihrt von Allied P. (ed.) Traiti de Zoologie. Anatomie, sysiemaiii/ne.

Voclt/kow. Wissenschaftliche Ergebnisse. Vol, 3 (3). biologie, Masson el Cie.: Paris, B, Schweiterbart: Stuttgart l-tn/jiidsled, H.V., 1923. Sponges from Ihe Auckland and

( Island. liomy Psnanll, N , 1971 . Spongiaires lie la /one lochcusc amphell In: T. Mortensen (ed.), Papers from

lie Bnnyiils sin Mei . II. Sv.slcmaliqiic. Vie Milieu (U) Dr. Ih. Morlensen's Pacific Expedition 1914-16. XV. 22 (2): 2X7 350. I'idensk. Mei/ilr. Dctnsk Naturh. linen. Kbit. 75: lioin y-l'.snanll, N., 1971a. Cnmpagnc de la Calypso an 117 167.

large des c6tcs atlantiqnes de I'Amerique du Sud lirr/mdsted, I I.V., 1924. Sponges from New Zealand. Pari

(1961 1962), I. 29. Spongiaires, Risult. Sclent. Camp. I. In: I . Morlensen (ed.). Papers from Dr. Th. Mor- Calypso 10: 263 295. lensen's Pacific Expedition 1914-16. XXIII. Vidensk. Boui v Psnault, N. and van lievcren, M., 1982, Pes Meddr. Dansk Ntilurli. loren. Kbit. 77: 435-483.

deinosponges du plateau continental de Kerguelen- Ui(/>ndsted, 1 1. V., 1926. Sponges from New Zealand. Pari

I leant. ( Nl HA (l oni. Nam. I r. Reck. Annuel.) 52, II. In: T. Morlensen (ed.), Papers from Dr. Th. Mor- 176 pp., 21 pis. lensen's Pacific Expedition 1914-16. XXV. Vidensk. liomy I'.snanll, N. and lopes, M.I., 1985. les Meddr. Dansk Nalinii. loren. Khli 81: 295-331.

demosponges littorales tie 1'arehipel des Aeores. Ann. Ur

Inst. Oceanagr. 61 (2): 149 225. collected by S. Wallin 1924. Ark. Zool. I9A (6): 1-6. Bowerbank, I.S., 1841c. Description of three species of Hrt/'iidskd, II. V., 1929. Neue Schwamnie aus Amoy an sponge, containing some new Forms of organization, tier loimosaStrassc. Zool. Anz. 81: 224 229.

MicroSC, ./. / inul.: 161 162. (also in Ann. Mag. Nat. Uo/mdsled, H.V., 1934. Sponges. In: Sliaelcn, V. van Hist. 8; 393 394 oi 1842) (ed.), Kesnllals scicntifiques dn voyage aux hides

Bowerbank, J.S., 1844a, On the keratose or homy Orientates neeriandaises tie 1. 1.. AA. RR. le Prince sponges of commerce, Trans. Microsc. Sue. Loud. el la Piineesse Leopold tie ISelgiqne. Mem. Mns. R. I: 52 59, pi. 3. I list. Nat. Belg, horssir. 2(15): 1-27,

Bowerbank, J.s., 1844b. < )n three species of sponge con- Hronii, H.G., 1859. Die Klussen nnd Ordnuniien des taining some new forms of organisation. Trans. I luerreielis. wissensehaftlieli dargesteili m Wort nnd Microsc. Soc. I nnd. I: 63 76, pi. 6 pi.7 omitted. llild. i.ister Hand, die Klassen nnd Ordnungen der Bowerbank, J.S., 1845, Observations on the Spongiadae, fornilosen lltiere (Amorpliozou). CI". Winter: Leip- wiili descriptions of some new genera, Ann. Mux. zig and Heidelberg, wi i 149 pp., 12 pis. Nat. Hist. (I) 16: 4(1(1 41(1, pis 13 14. Burton, M. 1924a. ["be genus Chondrilla. Ann. May.. Nat. Bowerbank, J.S,, 1859. On the anatomy and physiology I list. (9) 14: 206-209. of the Spongiadae. Pari I. On Ihe spictila. Philos. Burton, M., 1924b. A revision of [he sponge family

linns. R. SOC. 148: 279 332, pis 2.1 26. Donalidae. I'roe. /ool. Soc. I ond. 1924: 1033-1045,

Bowerbank, J.S., 1861, I isi of British sponges. In MeAn Pi. I.

drew, R., I isi of the British marine invertebrate Burton, M., I926e. Description of South African sponges fauna. Rep. Br. Ass. Adv. Sci. 30: 235 236. Oxford. collected in the South African Marine Survey. Pari Bowerbank, J.s., 1863c. On the anatomy and physiol- I. Myxospongida ami Aslroleliaxonida. Ann. Rep. ogy o\' die Spongiadae. Pan III. On the generic Fish. Mar. liiol. Surv. S. Afr. 5 (1928), Spec. Rep. characters, Ihe specific characters, and on the method I, 29 pp., 6 pis. of examination. I'lnl. Trans, R. Soc. 152 (1862): Urn ton, M., 1927a. A revision o( the genera and species 1087 1135, pis 72 74. contained in Lendenfeld's "Die Chalineen ties ans- Bowerbank, J.S., 1864. A Monograph of the British iraliselien tiebieles." Ann. Mag. Nat. Hist. (9) 20: Spongiadae, Vol, I: \x, 290 pp., 37 pis. Kay Soci- 289 296, 502-512.

ety: I oiulon. Burton, M.. 1927b. III. Reporl on Ihe sponges. In: Zoo-

Bowerbank, J.S., 1866, I Monograph of the British logical results of ihe Sue/ Canal Expedition, Trans.

Spongiadae. Vol. 2: 388 pp, Kay Society: l ondon. /ool. Soe. I. ond. 22 (1926): 71-83. Bowerbank, J.S., 1872. Contributions to a general his Burton, M., 1928. Report on some deep sea sponges from toiy ol ihe Spongiadae. Pan l. i'roe. /ool. Soc, ihe Indian Museum collected by the R.l. M.S. Inves- Lond. 1872: 115 129, pis 5, 6. tigator. Pari II. Tetraxonida (concluded) and I-iicer- J.S., Bowerbank, 1873a, Contributions to a general his- alosa. Ree. Indian Mas. 30: 109 135, pis I, 2. tory Of the Spongiadae. Part IV. I'roe. Zool. Sue. Burton, M.. 1929a. Porifcra. Pari 2. Antarctic sponges. Lond. 1873: 3 25, pis 14. British Antarctic {"Terra Nova") l:\pedition. ivio. J.S., Bowerbank, 1874a. .1 monograph of the British Natural History Report. Zoology 6 (4): 393-458, pis wii, .'67 Spongiadae. Vol. 3; pp., 92 pis. Kay Sod I -5, ety: I ondon. Button, M., 1930a. Norwegian sponges from the Norman Bowerbank, J.s., [877, Description iA' five new species Collection. I'roe. /ool. Soe. Lond. 1930: 487-546. sponges diseoveied A.li. Of by Meyer on Ihe Philip- Burton, M., 1930c. Additions to the sponge fauna e<( the pine Islands and (iiiinea. I'roe. New Zool. Soe. I ami. Gulf of Mannar. Ann. Mag. Nat. /list. (10) 5: 1877: 456 464. 665 676. lirasiei , M.IV, 1975. oinline history An of SeagrasS com- Burton, M., 1932a. Sponges. Discovery Rep. 6: 2.37-392, munities. Palaeontology IS: 681 702. pis 48 57, SPONGES FROM BASS STRAIT 173

Burton, M., 1934a. Sponges. Sci. Rep. Gt. Barrier Reef the Spongida. Pachytragida. Ann. Mag. Nat. Hist. Exped, 4: 513-621, pis 1, 2. (5) 11: 344-369, pis 14, 15. Burton, M., 1935. The family Plocamiidae, with descrip- Carter, H.J., 1883c. Contributions to our knowledge of tions of four new genera of sponges. Ann. Mag. Nat. the Spongida. Ann. Mag. Nal. Hist. (5) 12: 308-329, Hist. (10) 15: 399-404. pis 11, 14. Burton, M., 1936a. Notes on sponges from South Africa, Carter, H.J., 1885a. Descriptions of sponges from the with descriptions of new species. Ann. Mag. Nat. neighbourhood of Port Phillip Heads, South Aus- Hist. (10) 17: 141-147. tralia. Ann. Mag. Nat. Hist. (5) 15: 107-117, pi. 4. Burton, M., 1937. The littoral fauna of krusadai Island Carter, H.J. 1885b. Descriptions of sponges from the in the Gulf of Manaar. Bull. Madras Govt. Mus. neighbourhood of Port Phillip Heads, South Aus- 196-222. (n.s.) Mat. Hist. Sect. 1 (2): 1-58, pis 1-9. tralia. Ann. Mag. Nat. Hist. (5) 15: Burton, M., 1938. Non-calcareous sponges. Sci. Rep. Carter, H.J., 1885c. Descriptions of sponges from the Australia. Antarct. Exped. (C) 9 (5): 1-22. neighbourhood of Port Phillip Heads, South Aus- Burton, M., 1954. Sponges. In: The "Rosaura" Expedi- tralia. Ann. Mag. Nat. Hist. (5) 15: 301-321. the tion, pt. 5. Bull. Brit. Mus. (Sat. Hist.) Zool. 2 (6): Carter, H.J., 1885d. Descriptions of sponges from 215-239, pi. 9. neighbourhood of Port Phillip Heads, South Aus- Burton, M., 1956. The sponges of West Africa. Allan- tralia. Ann. Mag. Nat. Hist. (5) 16: 277-294. tide Report. Copenhagen 4: 111-147. Carter, H.J., 1885e. Descriptions of sponges from the Burton, M., 1959. Sponges. Sci. Rep. John Murray neighbourhood of Port Phillip Heads, South Aus- Exped. 10: 151-281. 41 figs. tralia. Ann. Mag. Nat. Hist. (5) 16: 347-368. Burton, M., 1963. A revision of the classification of the Carter, H.J., 1885f. Mode of circulation in the Spongida. calcareous sponges. British Museum (Natural His- Ann. Mag. Nat. Hist. (5) 15: 117-122, pi. 4 figs 5, 7. Descriptions of sponges from the tory): London. 693 pp. Carter, H.J., 1886a. South Aus- Burton, M. and Rao, H.S., 1932. Report on the shallow- neighbourhood of Port Phillip Heads, Hist. 17: 40-53. water marine sponges in the collections of the Indian tralia. Ann. Mag. Nal. (5) H.J., 1886b. Descriptions of sponges from the Museum, Part 1. Rec. Indian Mus. 34: 299-356, pi. Carter, Port Phillip Heads, South Aus- 18. neighbourhood of Nat. Hist. 17: 112-127. Cabioch, L., 1968. Contribution a la connaissance de la tralia. Ann. Mag. (5) H.J., 1886c. Descriptions of sponges from the faune des spongiaires de la Manche occidentals. Carter. neiehbourhood of Port Phillip Heads, South Aus- Demospongcs de la region de Roscoff. Call. Biol. Mag. Nat. Hist. (5) 17: 431-441, Mar. 9: 211-246. tralia. Ann. Carter, H.J., 1874. Descriptions and figures of deep-sea 502-516. Descriptions of sponges from the sponges and their spicules from the Atlantic Ocean, Carter, H..L, 1886d. Heads, South Aus- dredged up on board H.M.S. "Porcupine", chiefly neighbourhood of Port Phillip tralia. Ann. Mag. Nat. Hist. (5) 18: 34-55. in 1869; with figures and descriptions of some Descriptions of sponges from the remarkable spicules from the Agulhas Shoal and Carter, H..I., 1886c. of Port Phillip Heads, South Aus- Colon, Panama. Ann. Mag. Nat. Hist. (4) 14: neighbourhood Nat. Hist. 18: 126-149. 207-221, 245-257, pis 13-15. tralia. Ann. Mag. (5) Supplement to the descriptions of Carter, H.J., 1877. On a melobesian form of Carter, H.J., 1886f. Bracebridge Wilson's Australian sponges. Foraminifera (Gypsina melobesioides, mini); and fur- Mr. J. Mag. Nat. Hist. (5) 18: 271-290. ther observations on Carpenteria monticutaris. Ann. Ann. Carter, H.J., I886g. Supplement to the descriptions of Mag. Nat. Hist. (4) 20: 172-176. Bracebridge Wilson's Australian sponges. Carter, H.J., 1879b. Contributions to our knowledge of Mr. J. Mag. Nat. Hist. (5) 18: 369-379. the Spongida. Ann. Mag. Nal. Hist. <5) 3: 284-304, Ann. Carter, H..1., 1886h. Supplement to the descriptions of 343-360, pis 25-29. Mr. J. Bracebridge Wilson's Australian sponges. Carter, H.J., 1881a. Supplementary report on specimens Mag. Nat. Hist. (5) 18: 445-466, pi. 10. dredged up from the Gulf of Manaar, together with Ann. Carter, H.J., 1887a. Description of Chondrosia spurca, others from the sea in the vicinity of the Basse Rocks n. sp., from the south coast of Australia. Ann. Mag. and from Bass's Straits respectively, presented to the Nat. Hist. (5) 19: 286-288. Liverpool Free Museum by Capt. H. Cawne Warren. Carter, H.J., 1887b. On the position of the ampullaceous Ann. Mag. Nat. Hist. (5) 7: 361-385, pi. 18. the function of the water canal-system in the Carter, H.J., 1881b. Conributions to our knowledge of sac and 101-1 Spongida. Ann. Mag. Nal. Hist. (5) 19: 203-312. the Spongida. Am, Mag. Nat. Hist. (5) 8: 12, H.J., 1887c. On the reproductive elements of the 118-120, 241-259, pi. 9. Carter, old Spongida. Ann. Mag. Nat. Hist. (5) 19: 350-360. Carter, H.J., 1882a. New sponges, observations on 1977. A field guide to Australian marine ones, and a proposed new group. Ann. Mag. Nal. Colemanf N., life. Rigby: Adelaide. 223 pp. Hist. (5) 10: 106-125. Indies P..L. 1977. Mesozoic-Cenozoic sediments of the Carter, H.J., 1882b. Some sponges from the West Cook, Museum Eastern Indian Ocean. Pp. 119-150 in: J.R. Heirt- and Acapulco in the Liverpool Free remarks. zler, H.M. Belli, et al. (eds), Indian Ocean geology described, with general and classificatory pis and bioslratigraphy. American Geophysical Union: Ann. Mag. Nat. Hist. (5) 9: 266-301, 346-368, Washington, vii + 616 pp. 11, 12. knowledge ol Darraeh, T.A., 1985. Molluscan biostratigraphy of the Carter, H.J., 1883b. Contributions to our 174 F. WIEDENMAYER

Tertiary of southeastern Australia. Alcheringa 9: Natural History Report, Zoology 6: 269-392, pis 83-116. 1-15. Dendy, A., 1887c. The sponge-fauna of Madras. A report Dendy, A. and Frederick, L.M., 1924. On a collection on a collection of sponges obtained in the neighbour- of sponges from the Abrolhos Islands, Western Aus-

hood of Madras by Edgar Thurston, Esq. Ann. Mug. tralia. ./. Linn. Soc, Zool. 35: 477-519, pis 25, 26. Nat. Hist. (5) 20: 153-165, pis 9-12. Descatoire, A., 1969. Peuplements sessiles de I'Archipel

Dendy, A., 1889a. Studies on the comparative anatomy deGlenan. I. Inventaire, Spongiaires. Vie Milieu (B)

of sponges. II. On the anatomy and histology of 20 (1): 177-209. Stelospongus flabe/liforntis Carter; with notes on the Desqueyroux-Faundez, R., 1981. Revision de la collec-

development. Q. ./. Microsc. Sci. 29: 325-358, pis tion d'eponges d'Amboine (Moluques, Jndonesie) 30-33. constitute par Bedot et Pictet et conservee au Dendy, A., 1889d. Report on a second collection of Museum d'histoire naturelle de Geneve. Rev. Suisse sponges from the Gulf of Manaar. Ann. Mug. Nut. Zool. 88 (3): 723-764. HtsL (6) 3: 73-99, pis 3-5. Desqueyroux-Faundez, R., 1984. Description de la faune Dendy, A., 1890a. Zoology: Invertebrata. Pp. 74-96 in: des Haplosclcrida (Porifera) de la Nouvelle-

W.B. Spencer (ed.), Handbook of the Australasian C aledonie. I. Niphatidae-Callyspongiidae. Rev. Association (ANZAAS) for 1890. Melbourne. Suisse Zool. 91 (3): 765-827. (Sponges pp. 76-79). Dragnewitseh, P., 1906. Spongien von Singapore. Zool. Dendy, A., 1895. Catalogue of non-calcareous sponges Jb. (Systj 23: 439-448. collected by .1. Braccbridge Wilson, Esq., M.A., in Duchassaillg de Fonbressin, P. and Michelotti, G., 1864. the neighbourhood of Port Phillip Heads. Part 1. Spongiaires de la mer Caraibe. Natuurk. Verb. Holl.

Proc. R. Soc. Vic. (n.s.) 7: 232-260. Mautsch. Wet. Haarlem (2) 21 (3): 1-124, pis 1-25. Dendy, A., 1896. Catalogue of non-calcareous sponges Fillers, E., 1870. Die Esper'schen Spongien in der collected by .1. Braccbridge Wilson, Esq., M.A., in zoologischen Saminlung der K. Universitat Erlangen. the neighbourhood of Port Phillip Heads. Part 2. E.T. Jacob: Erlangen. 36 pp. Proc. R. Soc. Vict, (n.s.,) 9: 230-259. Ehrenberg, C.G., 1873. Mikrogeologische Studien uber Dendy, A.. 1897. Catalogue of non-calcareous sponges das kleinste Feben der Meeres-Tiefgrunde aller Zonen collected by .1. Bracebridge Wilson, Esq., M.A., in und dessen geologischen Einfluss. Abb. K. Akad. the neighbourhood of Port Phillip Heads. Part 3. Berlin, 1872, Phys. A/.: 131-397(399), pis 1-12. Proc. R. Soc. Vict, (n.s.) 9: 230-259. Fkman, S.P., 1953. Zoogeography of the sea. Sidgwick Dendy, A., 1898. On the sponges described in Dieffen- and Jackson: London, xiv + 417 pp. bach's "New Zealand." Trans. Proc. N.Z. Inst. 30: Esper, E.J.C., 1791-1799. Die Pflanzenthiere in Abbil- 316-320, pis 33, 34. dungen nach der Natur mit Farben erleuchtet, nebst Dendy, A., 1905. Report on the collected sponges by Beschreibungen. Raspe: Nuremberg. Vol. 2, 303 pp., Professor Herdman, at Ceylon, in 1902. In: \\ A. 109 pis. (Sponges: pp. 165-282, 289-294; pis Spon- Herdman, Report to the Government of Ceylon on gia 1-7, 7A.7B. 8, 9, 9A, 10-21, 21A 22, 23A, 23A, the Pearl Oyster Fisheries of the Gulf of Munaar. 24-48, 48A, 49). Issued in parts; for details, see Sher- Royal Society: London. Vol. 3, Suppl. 18: 57-246, born, 1902: xxii. 1-16. pis Eva, A.N., 1980. Pre-Miocene seagrass communities in Dendy, A., 1916a. Report on the Homoscletophora and the Caribbean. Palaeontology 23: 231-236. Astrotetraxonida collected by H.M.S. Sealark in the George, W.C, and Wilson, H.V., 1919. Sponges of Indian Ocean. In: Reports of the Percy S/utlen Trust Beaufort (N.C.) Harbor and vicinity. Bull. Bur. Fish. Expedition to the Indian Ocean in 1905, Vol. 6. 36: 130-179, pis 56-61. Trans. Linn. Soc. Lond., Zool. 17: 225-271, pis Gray, J.E., 1843. Additional radiated animals and 44-48. annelides. Pp. 292-295 in E. Dieffenbach, Travels Dendy, A., 1916b. Report on the non-calcareous sponges in New Zealand, with contributions to the geology, collected by .lames Mr. Hornell at Okhamandal in botany and natural history of that country. Vol. 2. Kattiawar in 1905-6. In: J. Hornell, Report to the John Murray: London. Government of Buroda on the Marine Zoology of Gray. .I.E., 1867. Notes on the arrangement of sponges, in Okhamandal Kattiawar.\o\.2 (3): 93-146, pis 1 -4. with the description of some new genera. Proc. Zool. Dendy, A., 1917. On the occurrence of gelatinous spic- Soc. Lond. 1867: 492-558, pis 27, 28. ules, and their mode of origin, in a new genus of sili- Guiler, E.R., 1950. Notes on Tasmanian marine sponges. ceous sponges. Proc. R. Soc. Lond. (B)89: 315-322, Rec. Queen Vict. Mus. Launceston 3: 5-14. pi. II. Hallmann, E.F., 1912. Report on the sponges obtained Dendy, A., 1921 . The tetraxonid sponge spicule: a study by the F.l.S. Endeavour on the coasts of New South in evolution. Acta Zool. 2: 95-152. Wales, Victoria, South Australia, Queensland, and Dendy, A., 1922. Report on the Sigmatotetraxonida col- Tasmania, 1909-10. Part 1. Zool. Results Fish. Exp. lected by H.M.S. Sealark in the Indian Ocean. In: Endeavour!: 117-300, pis 21-36. Reports the of Percy Sladen Trust Expedition to the Llallmann, E.F., 1914a. A revision of the monaxonid spe- Indian Ocean in 1905. Vol. 7. Trans. Linn. Soc. cies described as new in Lendenfeld's "Catalogue of lond.. Zool. 18: 1-164, pis 1-18. the sponges in the Australian Museum." Part 1. Proc. Dendy, A., 1924. Porifera. Part 1. Non-antarctic sponges. Linn. Soc. N.S.W. 39: 263-315, pis 15-24. British Antarctic (Terra Nova) Expedition. 1910. Hallmann, E.F., 1914b. A revision of the monaxonid spe- SPONGES FROM BASS STRAIT 175

cies described as new in Lendenfeld's "Catalogue of Kukenthal and T. Krumbach (eds.), Handbuch der

the sponges in the Australian Museum." Part 2. Proc. Zoologie. Vol. 1, Protozoa, Porifera, Coelenteraia, Linn. Soc. N.S.W. 39: 327-376. Mesozoa. Walter de Gruyter: Berlin and Leipzig, xv Hallmann, E.F., 1914c. A revision of the monaxonid spe- + 1060 pp. cies described as new in Lendenfeld's "Catalogue of Hinde, G.J., 1910. On the fossil sponge spicules in a rock the sponges in the Australian Museum." Part 3. Proc. from the deep lead (?) at Princess Royal township, Linn. Soc. N.S.W. 39: 398-446. Norseman district. Western Australia. Bull. Geol. Hallmann, E.F., 1916a. A revision of the genera with Surv. W. Aust. 36: 7-24, pis 1-3. microscleres included, or provisionally included, in Hinde, G.J. and Holmes, W.M., 1892. On the sponge the family Axinellidae; with descriptions of some remains in the Lower Tertiary strata near Oamaru,

Australian species. Part I. Proc. Linn. Soc. N.S. W. Otago, New Zealand. J. Linn. Soc, Zool. 24: 41: 453-491, pis 21-29/1-2; 39/6-7. 177-262, pis 7-15. Hallmann, E.F., 1916b. A revision of the genera with Hooper, J.N. A., 1984b. A new genus and two new spe- microscleres included, or provisionally included, in cies of haplosclerid sponges (Porifera: Demospon- the family Axinellidae; with descriptions of some giae) from the Timor Sea, Northwest Australia. Proc. Australian species. Part 2. Proc. Linn. Soc. R. Soc. Vict. 96 (2): 55-60. N.S. W. 41: 495-552, pis 29/4; 30-32; 33/1-5; 34-37; Hoshino, T., 1981. Shallow-water demosponges of 38/1-4. Western Japan, I. J. Sci. Hiroshima Univ. (Bl) 29 Hallmann, E.F., 1917a. A revision of the genera with (1): 47-205. II. Ibidem (Bl) 29 (2): 207-289. microscleres included, or provisionally included, in Hyatt, A., 1875. Revision of the North American the family Axinellidae; with descriptions of some Poriferae; with remarks upon foreign species. Part Australian species. Part 3. Proc. Linn. Soc. N.S.W. 1. Mem. Bost. Soc. Nat. Hist. 2: 399-408, pi. 13. 41: 634-675, pis 29/3, 5-6; 33/6; 38/5-9; 39/1-5; Hyatt, A., 1877. A revision of the North American 40/1-4; 41-44. Poriferae; with remarks upon foreign species. Part Hallmann E.F., 1917b. On the genera Echinaxia and 2. Mem. Bost. Soc. Nat. Hist. 2: 481-554, pis 15-17. Rhabdosigma. Proc. Linn. Soc. N.S, W. 42: 391-405, Jackson, J. B.C., and Hughes, T.P., 1985. Adaptive pis 21, 22. strategies of coral-reef invertebrates. Amer. Sci. Hallmann, E.F., 1920. New genera of monaxonid sponges 73(3): 265-274. related to the genus Clalhria. Proc. Linn. Soc. Johnston, G., 1842. A history of British sponges and S.S.IV. 44: 767-792, pis 36-40. lithophytes. W.H. Lizars: Edinburgh, xii + 264 pp., Hartman, W.D., 1982. Ponfera. In: S.P. Parker (ed.), 25 pis. Synopsis and classification of living organisms. Keller, C, 1889. Die Spongienfauna des rothen Meeres 20-25. McGraw-Hill: New York. Vol. 1, xviii + 1166 pp. (1. Halfte). Z. Hiss. Zool. 48: 311-405, pis Hartman, W.D., Wendt, J.W. mid Wiedenmayer, F., Kclier, C, 1891. Die Spongienfauna des rothen Meeres 1980. Living and fossil sponges (Notes for a short (2. Halite). Z. Hiss. Zool. 52: 294-368, pis 16-20. Bivalvia. 353-383 course). Sedimenla 8, Comp. Serf. Lab. Univ. Miami, Kauffman, E.G., 1973. Cretaceous Pp. in: Hallam, A. (ed.), Atlas of palaeobiogeography. vi +- 274 pp. Amsterdam, xii, 531 Hechtel, G.J., 1965. A systematic study of the Demospon- Elsevier: pp. Frakes, L.A., and Hayes, D.E., 1975. giae of Port Royal, Jamaica. Bull. Peabody Mus. Kemp, EM.. Paleoclimatic significance of diachronous biogenic Nat. Hist. 20, iv + 103 pp., 8 pis. 28, Deep Sea Drilling Project. Pp. Hechtel, G.J., 1969. New species and records of shallow facies, Leg Hayes, L.A. Frakes et al., Initial water Demospongiae from Barbados, West Indies. 909-917 in: D.E. Reports the Deep Sea Drilling Project, vol. 28. Poslilla Peabody Mus. Sat. Hist. 132: 1-38. of Government Printing Office: Washington, xx Hechtel, G.J., 1976. Zoogeography of Brazilian marine U.S. + 1017 Demospongiae. Pp. 237-260 in F.W. Harrison and pp. Kennett, J. P., and von der Borch, C.C., 1986. South- R.R. Covvden, (eds.). Aspects of sponge biology. west Pacific Cenozoic paleoceanogrpahy. Pp. Symposium, Albany, N. Y., U.S.A., May 1975. Aca- 1493-1517 in: J. P., Kennett, C.C. von der Borch et demic Press: New York, xiii + 354 pp. al.. Initial Reports of the Deep Sea Drilling Project, Hechtel, G..L, 1983. New species of marine Demospon- vol. 90. U.S. Government Printing Office: Washing- giae from Brazil. Iheringia (Zool.J 63: 59-89. ton, xxi + v + 1517 pp. (in 2 parts) Henrschel, E., 1909. Tetraxonida. 1. Teil. In: W. Michael- Kennett, J. P., Houtz, R.E., et al., 1975. Initial Reports sen and R. Hartmeyer, (eds.), Die fauna Siidwest- of the Deep Sea Drilling Project, vol. 29. U.S. Austruliens. 2 (21): 347-402, pis 22, 23. Government Printing Office: Washington, xxi + Hentschel, E., 191 1. Tetraxonida. 2. Teil. in: W. Michael- 1197 pp. sen and R. Hartmeyer, (eds.), Die Fauna Sudwest- Kirk, H.B., 1909. Two sponges from Campbell Island. Australiens. 3 (10): 279-393. Pp. 539-541, pi 25 in: C. Chilton (ed.), The Suban- Hentschel, E., 1912. Kiescl-und Hornschwamme der Aru- tarctic Islands of New Zealand. Vol. 2. Philosophi- und Kai-Inseln. Abh. Senckenb. Naturf. Ges. 34: cal Institute of Canterbury: Wellington. 295-448, pis 13-21. Kirk, H.B., 1911. Sponges collected at the Kermadec Hentschel, L., 1913. Ucber einen Fall von Orthogenese Islands by Mr. W.R.B. Oliver. Trans. Proc. N.Z. bei den Spongien. Zool. Am. 42: 255-267. Inst. 43: 574-581, pi. 27. Hentschel, E., 1923. Erste Unterabteilung der Mctazoa: Kirkpatrick, R., 1900a. IV. On the sponges of Christmas Parazoa, Porifera = Schwamme. Pp. 307-418 in: W. 176 F. WIEDENMAYER

Island (In: Andrews, C.W., Smith, E.A., Bernard, Lamarck, J.B.P.A. de Monet, de, 1813-1814. Sur les H.M., Kirkpatrick, R. and Chapman, F.C.: On the polypiers empales. Ann. Mus. Hist. Nat. Paris 20: marine fauna of Christmas Island (Indian Ocean)). 294-312 (1813); 370-386, 432-458 (1814). Proc. Zool. Soc. Loud. 1900: 127-141, pis 12, 13. Lamarck, J.B.P.A. de Monet, de, 1814-1815. Suite des

Kirkpatrick, R., 1903b. Descriptions of South African polypiers empates. Mem. Mus. Hist. Nat. Paris 1:

sponges. Part iii. Mar. Invest. S. Afr. 2: 233-264, 69-80 (1814); 162-168, 331-340 (1815). pis 5, 6. Laubenfels, M.W. de, 1930. The sponges of California. Kirkpatrick, R., 1910. On new species of Hydrozoa and Stanford Univ. Bull. (5) 5 (98): 24-29. Porifera of St. Helena. Proc. Zool. Soc. Lond. 1910; Laubenfels, M.W. de, 1932. The marine and fresh water 127-131, pi. 7. sponges of California. Proc. U.S. Naln. Mus. Knox, G.A., 1960. Littoral ecology and biogeography of 2927/81 (4): 1-140. the southern oceans. Proc. R. Soc. Lond. (B) 152: Laubenfels, M.W. de, 1935. A collection of sponges from 577-624. Puerto Galera, Mindoro, Philippine Is. Philip. J. Sci. Koltun, V.M., 1959. The Cornacuspongida of the north- 56 (3): 327-337. ern and fareastern seas of the U.S.S.R. In: Keys to Laubenfels, M.W. de, 1936a. A discussion of the sponge the fauna of the U.S.S.R. published by the Zoologi- fauna of the Dry Tortugas in particular, and the West cal Institute, Akademia Nauk SSSR. vol. 67. Indies in general, with material for a revision of the Izdatel'stvo "Nauka": Moscow. 236 pp., 43 pis (in families and orders of the Porifera. Publ. Carnegie

Russian). Instn. (Washington) 467: iii + 225 pp., 22 pis, 1 map. Koltun, V.M., 1964. Sponges of the Antarctic. Part I. Laubenfels, M.W. de, 1936b. A comparison of the Tctraxonida and Cornacuspongida. Pp. 6-116, shallow-water sponges near the Pacific end of the 428-433 (systematic index), pis 1-15 in E.P. Pavlov- Panama Canal with those at the Caribbean end. Proc. skiy (Chief ed.), A. P. Andriyashev and P.V. Ushakov U.S. Nutn. Mus. 83 (2993): 441-466. (eds), Academy of Sciences of the U.S.S.R., Zoo- Laubenfels, M.W. de, 1939. Sponges collected on the logical Institute: Exploration of thefauna of the seas presidential cruise of 1938. Smithson. Misc. Coll. 98 II (X). Biological Results of the Soviet Antarctic (15): 1-7. Expedition (1955-1958) 2. Izdatel'stvo "Nauka": Laubenfels. M.W. de, 1948. The order Keratosa of the Moscow, (in Russian) phylum Porifera. A monographic study. Occ. Pap. Koltun, V.M., 1966a. Sponges of the Antarctic. Part I. Allan Hancock Fdn. 3: 1-217, 30 pis. Tetraxonida and Cornacuspongida. Pp. 6-1 IS, Laubenfels, M.W. de, 1950a. The sponges of Kaneohe 443-448 (systematic index), pis 1-15 in E.P. Pavlov- Bay, Oahu. Pacif. Sci. 4 (1): 3-36. skii (Chief ed.), A. P. Andriyashev and P.V. Ushakov Laubenfels, M.W. de, 1950b. The Porifera of the Ber- (eds), Academy of Sciences of the U.S.S.R., Zoo- muda Archipelago. Trans. Zool. Soc. Lond. 27:

logical Institute: Exploration of the fauna ofthe seas 1-154, pis 1, 2. II (X). Biological Results of the Soviet Antarctic Laubenfels, M.W. de, 1951. The sponges of the Island Expedition (1955-1958) 2. Translated from the Rus- of Hawaii. Pacif Sci. 5: 256-271. sian. Israel Program for Scientific Translations: Laubenfels, M.W. de, 1954a. The sponges of the West- Jerusalem. Central Pacific. Ore. St. Monogr. Zool. 7, x + 306 Koltun, V.M., 1966b. The Tetraxonida of the northern pp., 12 pis. and fareastern seas of the U.S.S.R. In: Keys to the Laubenfels, M.W. de, 1955a. Sponges of Onotoa. Pacif. fauna of the U.S.S.R. published by the Zoological Sci. 9 (2): 137-143. Institute, Akademia Nauk SSSR, vol. 90. Izdatel'stvo Laubenfels. M.W. de, 1957. New species and records of "Nauka": Moscow. 112 pp., 38 pis (in Russian). Hawaiian sponges. Pacif Sci. 11 (2): 236-251. Koltun, V.M., 1969. Porifera. Pp. 13. 14, pis 3, 4 in: Lendenfeld, R. von, 1883. Uber Coelcnteraten der Sudsee. Antarctic map folio series. Eolio II, Distribution of Neue Aplysinidae. Z. Wiss. Zool. 38: 234-313, pis selected groups of marine invertebrates in waters in 10-13. south of 35° latitude. American Geographical Soci- Lendenfeld, R. von, 1885a. Notes to the Australian ety: Washington. 44 pp., 29 pis. sponges recently described by Carter. Ann. Mag. Nat. Koltun, V.M., 1970. Sponges of the Arctic and Antarc- Hist. (5) 16: 20-25. (Also Proc. Linn. Soc. N.S.W. tic; a faunistic review. Pp. 286-297 in: W.G. Fry 10: 151-156.) (ed.), The biology of the Porifera. London Academic Lendenfeld, R. von, 1885b. The Phoriospongiae, Mar- Press (Symp. Zool. Soc. Lond. 25). .xxviii + 512 pp. shall. Proc. Linn. Soc. N.S.W. 10: 81-84. Koltun, V.M., 1971. Glava HI. K vidovomu sostavu Lendenfeld, R. von, 1885c. On a sponge destructive to gubok Krasnogo morya. (On species composition of oyster culture in the Clarence River. Proc. Linn. Soc. sponges of the Red Sea.) Pp. 31-43 in: V.A. N.S.W. 10: 326-329. Vodyanitzky (ed.), Bentos Shel'fa Krasnogo morya. Lendenfeld, R. von, 1885f. A monograph of the Aus- (Benthos the of shelf of the Red Sea.) Naukova tralian sponges. Part 5. Proc. Linn. Soc. N.S. W. 10: Dumka: Kiev (Akad. Nauk USSR). 296 pp. (In 282-325, pis 26-35. Russian.) Lendenfeld, R. von, 1885h. A monograph of the Aus- Koltun, V.M., 1976. Porifera. I. Part Antarctic sponges. tralian sponges. Part 6. Proc. Linn. Soc. N.S. W. 10: BANZAntarct. Res. Exp. Rep. (B) 9 (4): 147-198, 481-553, pis 36-38. pis 1-3. Lendenfeld, R. von, 1886a. A monograph of the Aus- SPONGES FROM BASS STRAIT 177

tralian sponges. Addendum 2. Proc. Linn. Soc. I'Atlantique oriental par le "President -Theodore- N.S.W. 10: 845-850. Tissier" (1955-1956). Rev. Trav. Inst. Peches Marit. Lendenfeld, R. von, 1886e. A monograph of the Aus- 22: 225-246. tralian sponges. Part 4. Two cases of mimicry in Fieberkiihn, N., 1859. Neue Beitrage zur Anatomic der sponges. Proc. Linn. Soc. N.S.W. 10: 569-574, pis Spongien. Arch. Anal. Phvsiol. Wiss. Med. (1859): 43, 44. 353-382, 515-529; pis 9-11. Lendenfeld, R. von, 1886f. An alga, forming a pseudo- Linne, C. von, 1759. Systema naturae. Editio decima, morph of a siliceous sponge. Proc. Linn. Soc. reformata, torn. 2, rcgnum vegetabile: 825-1384. L. N.S.W. 10: 726-728, pi. 48 fig. 5. Salvius: Stockholm. Lendenfeld, R. von, 1887a. Die Chalineen des aus- Lindgrcn, N.G., 1897. Beitrag zur Kenntniss der Spon- tralischen Gebietes. Zool. Jb. fSyst.) 2: 723-828, pis gienfauna des Malayischen Archipels und der 18-26, 28. t hmesischen Meere. Zool. Anz. 20: 480-487. Lendenfeld, R. von, 1888. Descriptive catalogue of the Lindgrcn, N.G., 1898. Beitrag zur Kenntniss der Spon- sponges in the Australian Museum, Sydney. Taylor gienfauna des Malayischen Archipels und der and Francis: London, xiv + 260 pp., 12 pis. chinesischen Meere. Zool. Jb. (Syst.) 11: 283-378, Lendenfeld, R. von, 1889a. Die Verwandtschaftsverhalt- pis 17-20 (Also issued separately as "Inaugural- nisse der Hornschwamme. /ool. Jb. (Sysl.) 4: 1-93. Dissertation", Upsala University, 1900, 96 pp.).

Lendenfeld, R. von, 1889b. A Monograph of the Homy I ocblich, A.R., Jr. and Tappan, FL, 1964. Part C, Pro- Sponges. Trubnerand Co.: London. 936 pp., 50 pis. tista 2. Sarcodina, chiefly "Thecamoebians" and For- Lendenfeld, R. von. 1897a. Die Clavulina der Adria. mianiferida. In: R.C. Moore (ed.), Treatise on Nova Acta Acad. Caesar. Leop. Carol. 69: 1-251, Invertebrate Paleontology. Geol. Soc. America:

pis 1-12. Boulder. 2 vols, xxxi I 900 pp. Lendenfeld, R. von, 1903. Tetraxonia. In: F.E. Sehul/e Lundbeck, W., 1905. Porifera (Part II). Desmacidoni- (ed.), Das Tierreich. Friedlandcr: Berlin. If. 19, xv dac (pars). Danish Ingolf-Expedilion. 6 (2): 1-219, 1-20. + 168 pp. pis Lendenfeld, R. von, 1907a. Die Tetraxonia. Pp. iv, Lundbeck, W., 1909. The Porifera of East Greenland. 59-374, pis 9-46 in: C. Chun (ed.), Wissenschaflliche Meddr. 6>nland 29: 423-464, pi. 14. Ergebnisse der Deulschen Tiefsee-Expedition aufdetn Lundbeck, W., 1910. Porifera (Part III). Desmacidoni- 1-124, Dampfer Valdivta I89S-IH99. 11. dac (pars). Danish Ingolf-Expedition. 6 (3): Levi, C, 1952. Spongiaires de la cole du Senegal. Bull. pis 111. liber Dysicleiden und Inst. Er. Afr. Noire 14: 34-59. Marshall, W., 1880. Untersuchungen Phoriospongien. Z. Wiss. Zool. 35: 88-129, pis 6-8. Levi, C, 1956b. Spongiaires des cotes de Madagascar. Fleck, K.L., Jr., 1976. Biogcography Mem. Inst. Scient. Madagascar (A) 10: 1-23. McCoy, E.D., and mangroves: an Levi, C, 1958. Rcsultats scientifiques des campagnes de of corals, seagrasses, and alternative to the center of origin concept. Sysl. Zool. 25: la Calypso. Campagne 1951-1952 en Mer Rouge. Spongiaires de Mer Rouge recueillies par la Calypso 201-210. ('., 1878. Preliminary account on sponges (1951-1952). Ann. Inst. Oceanogr. Monaco (n.s.) 34: Mcrejkowsky, S-Peterb. Obshch. 3-46. of the White Sea. Trudy Imp. 249-270. Levi, C, 1960. Les Demosponges des cotes de France. Estest. 9: Mosiler, FF, 1976. Porifcrcnspiculae der alpincn Trias. I. Les Clathridae. Cah. Biol. Mar. 1: 47-87. Paluont. Mitt. Innsbruck 6 (5): 1-42. Levi, C, 1965a. Spongiaires recoltes par 1'Expedilion Geol. Miiller, F., 1865. LJebcr Darwine/la aurea, cinen Israelienne dans le Slid de la Mer Rouge en 1962. Schwamm mit sternformigen Hornnadcln. Arch. (Israel S. Red Sea E.xpcd. Rep. 13.) Bull. Sea Fish. Mikrosk. Anal. 1: 344-353, pi. 21. Res. Sin. Israel 40: 3-27. M.G., and Kennetl, J. P., 1986. Development Levi, C, 1967a. Spongiaires d'Afrique du Sud, (3) Tctrac- Murphy, latitudinal thermal gradients during the Oligocene: tinellides. Trans. R. Soc. S. Afr. 37: 227-256, pis of southwest Pacific. 27-29. Oxygen-isotope evidence from the 1347-1360 in: J. P. Kenneft, C.C. von der Borch Levi, C, 1969. Spongiaires du Vema Seamounl (Allan- Pp. Reports the Deep Sea Drilling Project. tiqueSud). Bull. Mus. Nam. Hist. Nat. Paris 41 (4): et al.. Initial of 90. U.S. Government Priming Office: Washing- 952-973, pis 1-5. vol.

Ion, xxi -t v t 1517 pp. (in 2 parts). Levi, C, 1973. See Brien et al. (1973). Nardo, G.D., 1833. Auszug aus cincm neuen System der Levi, C, 1979. The demonsponge fauna from the New Spongiarien, wornach bcreits die Aufstellung in der Caledonian area. In: Proceedings of the International Padua gemacht ist. Isis Symposium on Marine Biogcography and Evolution Universitats-Sainmlung zu Oken. coll. 519-523. in the Southern Hemisphere, Auckland, New 1834. Despongiis. IsisOken, coll. 714-716. Zealand, 17-20 July, 1978. Vol. 1.JV.Z DSIR Inf. Nardo, G.D., Nardo, G.D., 1847a. Prospetlo della fauna marina vol- Ser. 137 (1): 307-315. bathyales garc del Vcneto estuario. Pp. 113-156 (pp. 1-45 in Levi, C. and Levi, P., 1984 Demosponges Nouvelle reprint) in: (Estratlo dall'opera) Venezia e le sue recollees par le N/O "Vauban" au sud de la A, lagune. Anlonclli: Venice. Vol. 2, pt I. Caledonie. Bull. Mus. Natn. Hist. Nat. Paris (4 Nardo, G.D., 1847b. Osservazioni anatomiche sopra 1'ani- ZooL) 5 (4): 931-997. recoltees dans malc detto volgarmente rognone di mare. Atti 1st. Levi, C, and Vacelet, .1., 1958. Eponges 178 F. WILDLNMAYLR

Veneta 6: 267-268. Keymcnt, R.A., 1973. Cretaceous history of the South Norman, A.M., 1869. Shclland final dredging report. Part Atlantic Ocean. Pp. 805-X14 in: D.H. Tarling and

II. On the ( rustacea, Tunicata, Polyzoa, Echinoder- S.K. Runcorn teds.), Implications of continental drift m.ii.i, Actinozoa, Hydrozoa, and Porifera. Rep. Br. to the Earth sciences. Academic Press: London, vol. ASS. Advml. Sci. 38 (1X6X): 247-336. 2, pp. i-xiv, 623-11X4 Owen, H.G., 1983. Alius ofcontinental displacement, 2fXJ Ridley, S.O., 1X84. XI. Spongida. In: Account of the zoo- million yews to the present. Cambridge University logical collections made during the survey of H.M.S.

PreSs (( ambridge Earth Science Series), x I 159pp. "Alerl" in the Slraits ol Magellan and on the coast

Pansini, M ., I983j Notes OH some Mediterranean Axinella ol I'alagonia. I'roc. '/.ool. Soc. l.ond. IXXI: 107-137,

wnh description oi two new species. Boll. Mus. 1st. 140, 141, pis 10, 11.

Biol. Univ. Genova 50-51: 79-98, Ridley, S.O., 1884a. Spongiida. (Part 1, Australian Pick, F.K., 1905. DjeGattung Raspailia. Arch. Natwg, sponges). Pp. 366-482, pis 39-43 in: Report on the 71: 1-48, pis 1-4. Zoological Collections Made in the Indo-Pacific ilu: I'oleiaell, N., 1 884b. Report on Kcratosa eollceted Ocean (taring I he Voyage of H.M.S. Alerl,

by 1 1. M.S. ( hallengei during the years 1X73-1X76. I88I-IHH2. British Museum (Natural History): Rep. Sci. Results Voyage Challenger, Zool. II: i-xx, London.

pis I 10. Ridley, S.O., 1884b. Spongiida. (Part 2, Collections from

Pouliqiten, I ., 1972. Les Spongiaires des grottes sons- the western Indian Ocean). Pp. 5X2-630, pis 53, 54

marines de la region de Matseille: I eologie el sys- in: Report on the Zoological Collections Made in the lemaliqiie. Tethys 3 (4) (1971): 717-75X. Indo-I'acific Ocean during the Voyage of H.M.S. Prcmoli Silva, I., and linisa, C, 19X1. Shallow water Alerl, 1X81-1882. British Museum (Natural History):

skeletal debris and larger roraininilers from Deep Sea London. XXV I 684 pp., 54 pis. Drilling Project Site 462, Nauru Basin, western Ridley, S.O., lX84c. Notes on sponges, with descriptions J equatorial Pacific. l p. 439-473 in: R.L. Larson, S.O. ol new species. Ann. Mag. Nal. Hist. (5) 14: 183-187. Seliianget ei al,, Initial Reports of the Deep Sea Dnii Ridley, S.O. and Dendy, A., 1886. Preliminary report ing Project, vol. 61. U.S. (iovernmenl Printing on the Monaxonida collected by H.M.S. Challenger.

Office: Washington, xxvii I XX5 pp. Ann. Mug. Nal. Hist. (5) 18: 325-351, 470-493. I'ron/alo, R., 1975. Nole lassonomielte snl genen Dm Ridley, S.O. and Dendy, A., 1887. Report on the Monax-

winella (Porifera). Boil. Mus. 1st. Biol. Univ. Genova onida collected by H.M.S. Challenger during the

43: 5 20, I eol. pi, years 1873-76. Rep, Sci. Results Voyage Challenger. (I., l'nlil/er linali, 1972. Report on a collection <>l Zool. 20 (59): Ixviii f 275 pp., 51 pis, 1 map. sponges from the Hay of Naples. I. Sdcrospongiac, Ridley, S.O. and Duncan, P.M., 1881. On the genus

Lithistida, I eliaelinellida, Lpipolasida. I'uhhl. Sit/:.. Plocamia, Schmidt, and on some other sponges of ZooJ. Napoli 38 (1970): 32X-354. the order lxhinonemala. With descriptions of two

Pulitzet Finali, (J., 1978. Report on a collection of addilional new species of Dirrhopalum. .1. Linn. Soc.

sponges I torn the Hay of Naples. III. I ladronierida, Zool. 15: 476-497, pis 28, 29. Axincllida, I'oeeiloselerida, llaliehondrida, Roekman, I., 1974. Underwater Australia. Lansdowne

Haploselerida. Holl. Mus. 1st. Biol. Univ. Cenova Press: Melbourne. 112 pp., col. and b/w pis. 45 (1977): 7X9. Row, R.W.H., 191 1. Report on the sponges collected by

I'ulil/er-linali, G., 19X21). Some new or little known Mr. Cyril Crossland, 1904-5. Part II. Non-Calcarea. sponges Irom the Cheat Battler Keel of Australia. In: Reports on the marine biology of the Sudanese

Boll. Mus. 1st. Biol. Univ. Genova 4X 49 Red Sea. XIX. ./. I inn. Soc, Zool. 31: 287-400, pis (19X0 19X2): 87- 141. 35-41.

I'nhl/er I'inali, G., 19X3. A collection c>( Mediterranean Rubio, M., Urix, J. and Bibiloni, A., 19X1. Contribu- Demospongiae (Porifera) with, in appendix, a list of tion a la fauna de esponjas del litoral Catalan.

the Demospongiae hitherto reeorded from the I sponias corneas, fund. Juan March Set: Univ. 164:

Mediterranean Sea. Ann, Mus. Civ. Slot: Nal. I -54. Genova X4: 44.5-621. Riist, D., 18X5. Beitragezur Kenntniss der fossilen Radi- Pulitzer Linali, (i. and I'ton/ato, R., 1977. Report on olarien aus Gcsieinen des Jura. Palueontographicu a collection Of sponges Irom the Bay of Naples. II. 31: 269-322, pis 26-45. Keralosa. I'uhhl. Slur.. Zool. Napoli 40 (1976): Riit/ler, K., 1965. Systematik unci Oekologie der Potife- 83 104. refl aus Litoral-Sehattengebicten der Nordadria. Z.

Pulitzer I inali, (i. and I'ron/ato, R., 19X0. The Keralosa Morph. Oekol. Here 55: I -82. in a collection of Mediterranean sponges mainly Irom Riit/ler, K., 1970. Spatial competition among Porifera: the Italian eoasts. Ann. Mus. Civ. Slor. Nal. Gia solution by epizoism. Oecologia 5: X5-95. foinn Ihiria X3: 127-I5X. Rulzler, K., I97X. Sponges in coral reefs. In: D.R. Slod- R.inlf, II., 1X93. I'alaeospongiologie. Allgemeiner Theil. dart and R.L. Johannes (eds), Coral reefs: research

I'alaeonlogiaplucu 40: 1-232. methods Monogr. Oeeanogr. Meth. (UNESCO) 5:

Ridlev, S.O., IXXI . XI. Spongida. In: Account of the zoo- 299-313.

logical collections made dining the survey of 1 1. M.S. Sara, M., 1972. Ciuida ai Poriferi della fauna ilaliana. "Alert" in the Straits of Magellan and on the coast Quad, Civ. Staz. Idrohiol. Mi/two 3-4 (1971-2):

of I'alagonia. Proc. Zool. Soe. loud. IXXI: 107-137, 52-97, pi. 1 (colour). 140, 141. pis 10, II. Sara, M.. 1978. Demospongie di aequc superficial! della .

SPONGES FROM BASS STRAIT 179

Terra del Fuoco (Spedizioni AMI- Mares-GRSTS e cao and other Caribbean localities. Part II. SAI). Boll. Mits. 1st. Biol. Univ. Genova 46: 7-117. Haplosclerida. Stud. Fauna Curacao 62 (191): 1-173, Scheibnerova, V., 1978. Aptian and Albian benthic 18 pis. foraminit'ers of Leg 40, Sites 363 and 364, Southern Soest, R.W.M. van, 1982. A small collection of sponges Atlantic. Pp. 741-756 in: H.M. Bolli, W.B.F. Ryan (Porifcra) from Hong Kong. Pp. 85-95 in: B.S. et al., Initial Reports of the Deep Seu Drilling Project Morton and C.K. Tseng (eds.), Proceedings of the vol. 40. U.S. Government Printing Office: Washing- first International Marine Biological Workshop: The ton. x\v + 1079 pp. Marine Flora and Fauna of Hong Kong und Southern Scheltema, R.S., 1986. On dispersal and planktonic larvae China, Hong Kong, 1980. Hong Kong University of benthic invertebrates: an eclectic overview and Press. summary of problems. Bull. Mar. Set. 39 (2): Soest, R.W.M. van, 1984. Marine sponges from Cura- 290-322. cao and other Caribbean localities. Part 111. Schmidt, E.O., 1862. Die Spongien des Adriatischen Poecilosclerida. Stud. Fauna Curacao 66 (199): Meeres. Engelmann: Leipzig, viii + 88 pp., 7 pis. 1-167, 10 pis. Schmidt, E.O., 1864. Supplement der Spongien des Adri- Sollas, W..L, 1880. The sponge-fauna of Norway; a report atischen Meeres. Enthaltend die Histiologie und sys- on the Rev. A.M. Norman's collection of sponges temutische Ergdnzungen. Engelmann: Leipzig, vi + from the Norwegian coast. Ann. Mag. Nat. Hist. (5) 48pp., 4 pis. 5: 130-144, 241-259, 396-409, pis 6, 7, 10-12, 17. Schmidt, E.O., 1868. Die Spongien der Kitste von Algier. Sollas, W.J., 1886. Preliminary account of the letractinel- Mil Nachtrdgen zu den Spongien des Adriatischen lid sponges dredged by H.M.S. Challenger, 1872-

Meeres (drittes Supplement). Engelmann: Leipzig, iv 1876. Part 1. The Chorislida. Sci. Proc. R. Dublin + 44 pp., 5 pis. Soc. (n.s.) 5: 177-199. Schmidt, E.O., 1870. Grundzuge enter Spongien- Fauna Sollas, W.J., 1888. Report on the Tetractinellida collected des Atlanttschen Gebietes. Engelmann: Leipzig, iv by H.M.S. Challenger, during the years 1873-1876. + 88 pp., 6 pis. Rep. Sci. Results Voyage Challenger, Zool. 25 (63):

Schmidt, E.O., 1875. Spongien. Pp. 115-120, pi. 1 in. clxvi + 458 pp., 44 pis, 1 map. Jahreshericht der Commission z.ur wtssenschaftlichen Sollas, W..L, 1889. On the geodine genera, Synops,

Vntersuchung der deutschen Meere in Kiel, Jg. 2, 3 Vosm., and Sidonops. A correction. Sci. Proc. R. (1872, 1873). Dublin Soc. (2) 6: 276-277. Schultze, M.J.S., 1865. (Ueber ein Exemplar von Thiele, J., 1899. Studien uber pazifische Spongien. II. Hyalonema Sieboldi aus Japan und einen Schwamm Ueber einige Spongien von Celebes. Zoologica 24 (2): mil Nadeln aus Hornsubstanz.) Verh. Nttturh. Ver. 1-33, pis 1-5. Preuss. Kheinl. H'estf. 22 (3/2): 6-7. Thiele, J., 1900. Kieselschwammc von Ternate. I. Abh. Schulze, F.E., 1878a. Untersuchungen uber den Bau und Senckenb. Naturf. Ges. 25: 19-80, pis 2, 3. die Entwicklung der Spongien. Viertc Mittheilung. Thiele, J., 1903a. Kieselschwamme von Ternate. II. Abh. Die Familie der Aplysinidae. Z. H'iss. Zool. 30: Senckenb. Naturf Ges. 25: 933-968, pi. 28. 379-420, pis 21-24. Thiele, J., 1903b. Beschreibung einigcr unzureichend Schulze, F.E., 1878b. Untersuchungen iiber den Bau und bekannten monaxonen Spongien. Arch. Naturg. 69: die Entwicklung der Spongien. Sechste Mittheilung. 374-398, pi. 21, figs 1-28A. Die Gattung Spongeliu. Z. H'iss. Zool. 32: 117-157, Thiele, J., 1905. Die Kiesel-und Hornschwamme der pis 5-8. Sammlung Plate. Zool. lb. Suppl. 6: 407-496, pis Schulze, F.E., 1879a. Untersuchungen uber den Bau und 27-33. die Entwicklung der Spongien. Siebente Mittheilung. Thomas, P. A., 1970d. Studies on Indian sponges. IV. Die Familie der Spongae. Z. Wiss. Zool. 32: 593-660, Additions to the genus Corlicium Schmidt with notes pis. 34-38. on the distribution of Corlicium candelabrum Selenka, E.. 1897. Ueber einigeneueSehwamme aus der Schmidt. ./. Mar. Biol. Ass. India 10 (2) (1968): Sudsee, Z. Wiss. Zool. 17: 565-571, pi. 35. 260-263. Shaw, M.E., 1927a. On a collection of sponges from Thomas, P. A., 1973b. Marine Demospongiac of Mahc Maria Island, Tasmania. Proc. Zool. Soc. Land, Island in the Seychelles Bank (Indian Ocean). Ann. (Zool.) 203: i-x, 1-96, pis. 1927: 419-439. pi. 1. Mus. R. Afr. Cent. Shaw, M.E., 1927b. Note on the inclusion of sand in Topsent, E., 1892a. Contribution a l'etude des Spongiaiies Prince sponges. Arm. Mag. Nat. Hist. (9) 19: 601-609. de I'Atlanlique Noid. Result. Camp. Sci. 1-S. Sherborn, C.D., 1902. Index animalium Sectio prima. Albert I 2: 165 pp., 11 pis University Press, lix + Topsent, E., 1892c. Diagnoses d'Epongcs nouvelles de la . . . 1758-18W). Cambridge Mcditerranee ct plus particulierement de Banyuls. 1195 pp. Gen. 10, Notes Rev.: Simpson, T.L., 1968a. The structure and function of Archs. Zool. Exp. (2) of xvii-xxviii. sponge cells: new criteria lor the taxonomy Nouvellc serie de diagnoses d'Eponges poecilosclerid sponges (Dcmospongiae). Bull. Pea- Topsent, E., 1894b. Banyuls. Archs. Zool. Exp. Gen. body Mus. Nut. Hist. 25: 1-141, pis 1-17. de Roscoff et de Notes Rev.: xxxiii-xliii. Soest, R.W.M. van, 1978. Marine sponges from Cura- (1893) (3) 1, 1896b. Etude monographique des Spon- cao and other Caribbean localities. Part I. Keratosa. Topsent, E., France. II. Carnosa. Archs. Zool. Exp. Stud. Fauna Curacao 56 (179): 1-94, 15 pis. giaires de Gen. (3) 3: 493-590, pis 21-23. Soest, R.W.M. van, 1980. Marine sponges from Cura- I HO WILDLNMAYLK

•', lopscni, E., 1897. S|)(iii)'i,iiK lie la baie d'Amboine. Vacclcl, I., I960. Sponges de la Mcdilerrancc nord-

(Voyage ili' MM. M. Bedol el < Pictel dans I'archipcl oceitlcnlalc iccollccs par Ic 'Trcsidcnl Theodorc- Malais.) Rev Suisse Zool. 4: 421 487, pis IK 21. Tissier" (1958). Rev, Trav. Inst. Piches Marit. 24:

I' TopMIU, . 1898. Introduction a I'cludc inonogiaphic|iie (2): 257 272.

des Monaxonidcs dc Prance. < ImssiIk.-iI n»n ties Vacclci, I., 1969. Sponges de la Roche dn Large el de

I ladromeriua. Archs, Zool. i\/>. Gin, (3)6:91 113. I'etage bathyal de Miditerranee (recoil es de la Sou

1 lopscni, I S| < ., 1904, gjaircs des Azores. Result, 'amp, coupe plongeante ( ousteau el dragageg). Mern. Mus.

i pis. Sclent. Prince Itbert 25: 280 pp . 18 Num. Hist. Nat. Puns (A) 59 (2): 145-219.

I ('., opsent, E. ( 1905, Etude sui lea Dendrocerattda, Archs. Vacclcl, I., Vasscui, P. and Levi, 1976. Spongiaires

/mil. Exp. Gin (i) i. clxxi cxeii. de la penle cxlernc des reeils coralliens del nlear (snd- lopscni, r •'., 191 'a. Spongiaires de I'expidition an tare onesi de Madagascar). Mim. Mus. Nuin. Hist. Nat.

tique nationals ecossaisc. Tram R.Soc Edinburgh 1'iiris (ii. s A) 49: I 116, pis I 10.

I 49 (3/9): 579 643, pis 6. Vcion, J.E.N. , 1974. Southern geographic limiis to Lhe lopscni, r , 1913b, Spongiaires prove I descampagncs distribution Of Greal Harrier Keel licrmalypic corals.

si icnli lie pics ilr la I 'iiiices.sc Alice dans les Mo s du Pp. 463 473 in: A.M. Cameron el al. (eels), Proceed

Nord(1898 1899, 1906 1907), Result. ( amp. Sclent. ings <>l the Second International Symposium on t oral

t Prince \ibert 45, 67 pp., 5 pis. Reels. I he (neal Banici Reef Committee: Brisbane.

I ["opsent, E,, 1918. pontes de San Thomi. Essai sui les Vol. I, \ I 630 pp. genres Spirastrelia, Donatio, el ChundriUa. Krchs. Vosmaer, G.C.J ., 1887. Porifera. In: I I.e. Bronn(ed.), Zool. Exp. Gin. 57: ^ts mk. Die Klusseu und Ordnungen des Thierreichs, C.F.

i I opsent, . , 1920< Sui quelques Sponges du Cabinet de Winter: Leipzig. 496 pp., f4 pis (5 pans issued I. Hermann decriies el figurfespai Espei en 1794 1882 6). Bull Sac Zool. France 45: 127, !14 Vosmaer, G.C.J. , 1911. The Porifer-a of the Siboga-

Topsent, E,, 1922, I es megascleres polyt-ylotes des Expedition, ll. I lie genus Spirastrelia. Siboga

l Hull. Monaxonidcs el la parentides al cutiincs. Expeditie, Monogr. 6a I: 69 pp., 14 pis.

Inst, Ocianogr. 415: I 8. Monaco Vosmaer, G.C. J. , 1932 1935, rhe sponges of the Bay of lopscni, l '.., 1924. Kevismii des Mycale de ['Europe Naples. Porifera Inealcaria. C.S. Vosinaci Kocll and

occidentale. Inst. (rt.s.) I Ann, Ocianogr. Monaco M. Burton (eds), Vol. I (1932 3) issued in 5 pails: // 118. (3); ( apita Zooiogica 3: I 456, pis I 35. Vol. 2 (1933- 5)

fopsent, I '., [925a, Studs de Spongiaircs du Golfe de issued in 5 paris: Capita Zooiogica 5: 457 857, pis

Naples Archs. Zool. Exp. Gin. 63: 623 725, pi. 8. 16 71 (pis 4.1, 55 omitted). Also published separately [opsent, r., 1928, Spongiaires de I'Ailamique ei de la in 3 \ols. (2 vols, text, 875 pp., and alias, 69 pis.). Mdditerranie provenant des croisieres dn Prince Mariinus Nijboff: lhe Hague. 1933 '5. Alberl la de Monaco, Risull, Camp. Scient. Prince Vosmaer, G.C.J, and Pekelharing, ('A., 1898, Obser-

Albert I 74: 376 pp.. I I pis. vations on sponges. I'erli. K, Akud. Uetenseh.

I opsent, L . L930a. Sponges de I amarck tonservees an Amsterdam (2) 6 (3): I 31, pis I 4. Nat. Museum de Paris. Arch. Mux. Num. Hist. Paris Wass, K.l ., C'onolly, IK. and and Maelnlyrc, K..L, 0.) S: I 56, pis l 4. 1970. Uiyo/oan caibonale sand continuous along fopsent, i ,, 1930b. < 'hondrociadia vatsui n, sp, de la Baie southern Australia. Mur. Geol. 9: 63 73.

de Sar.ami. Annotnes. Zool. Japan. 12: -4 J* i 432 Whitelegge, I. 1889. I is) of the marine and freshwater

lopscni, I ., 1932a, Sponges de I amarck conscivecsnu fauna of Pon Jackson and the neighbourhood, ./. Museum de Paris, Deuxieme panic Arch. Mus. R. SOC N.S. II. 23: 163 523 (sponges: 178 187).

Naln. Hist. Nat. Paris (6) 8: 61 124, pis I 6. Whitelegge, I '., 1897. lhe sponges of Funafuti. Mem.

lopscni, E., I'm I Sponges de amarck conservees an Aust. Mus. 3: 321 332, pi, 18. Museum de Paris. Fin. Arch. Mus. Num. Hist, Nat. Whitelegge, L, 1901. Report on sponges from the coastal

I pis I Paris (6) 10: 60, J. beaches of New South Wales. Rec. Aust. Mus. 4: lopscni, r., 1936a. Sponges obscrvces dans les parages ss I 18, pis 10 15.

de Monaco (deuxieme panic), Bull, Inst, Ocianogr. Whitelegge, I'., 1902a. Supplementary notes to the report Monaco 686: I 10, on sponges from the Coastal beaches Of New South lopscni, r '., t 1938a, onnncniaiics sni quelques genres Wales. Rec. AUSt. Mus. 4: 211 216.

d'Eponges marines. Hull. Inst. Ocianogr, Monaco l'., Whitelegge, 1902b. Notes on I endenfeWs types

744: i 13 described in the Catalogue of Sponges in the Aus- l sni n. una I, M., 1969, Sponges of Red Sea origin on the tralian Museum. Rec. Aust. Mus. 4: 274 288.

Mediterranean coasi of Israel. Israel J. Zool. 18: Whitelegge, T., 1906. Sponges. Pan I. In: Scientific 149 155. results of the trawling expedition of ll.M.c.s. Thetis v.uclci, i., [958. Dendrilla acantha nov. sp. rtouvelle off the coasi of New South Wales in February and Sponge comic meditei raneenne. Remarques sm les March, 1898. Mem. Aust. Mus. 4: 453 484, pis

genres Dendrilla I endcnfeld el Megalopastas Dendy. 43 44.

Reel, l'tuv. Sm. Mar. 23 147. I'., Endoume (14): 143 Whitelegge, 1907. Sponges. Pan I (Addenda), Pari Y.icelci, J., 1959. Repartition generate des Sponges el sys 2. in: Scientific results of the trawling expedition of leinalit|iie des Sponges coi nccs de la icgion de Mai H.M.C.S, Thetis off the coasi of New South Wales seille el de quelques stations midilerraneennes, Reel, in February and March, I89S. Mem. Aust. Mus. 4: iruv. sm. Mar. Endoume 26: !9 101, pis i 3, 4S7 515, pis 45, 46. SPONGES FROM BASS STRAIT ISI

Wiedenmayer, F., 1974. Recent marine shallow-water Fig. 2. Jaspis stellifera (Carter). NMV F51938, partially sponges of the West Indies and the problem of sped- surrounding a group of vcrmctids, a mouth of which is ation. Verh. Nalurf. Ges. Basel 84 (1): 361-375. visible on top. The cushion-shaped, perpendicularly cut Wiedenmayer, F., 1977a. Shallow-water sponges of the small sponge on the left shoulder is Gelliodes inerustans

western Bahamas. Birkhauser: Basel, (t'xperientia Dendy (NMV F52036). Behind it (extreme upper left) is Suppl. 28), 287 pp., 43 pis. Aitlorrhiza procumbens (Lendenfeld, Calcarea, not Wiedenmayer, F., 1978. Modern sponge bioherms of the described). xO.64. Great Bahama Bank. Eclogae Ceol. Helv. 71 (3): Figs 3, 4. Tetilla praecipua sp. nov. Holotype, NMV 699-744. F51943. Fig. 3. Lateral view, xO.81.Hg. 4. Choanosome Wiedenmayer, F., Hooper, J.N. A. and Raeek, A. A. (in from below, x0.7. press). Porifera. Zoological Catalogue of Australia. Figs 5, 6, 9. Telhya ingalli Bowerbank. Fig. 5. NMV Bureau of Flora and Fauna, Australian Government F5I944, xO.7. Fig.6a-d. Telhya phillipensis Lendenfeld, Publishing Service: Canberra. two wet syntypes, AM G9077, x().96. Fig. 6a, b. Top. Wilson, B.R. and Allen, G.R., 1987. Major components Fig. 6c, d. Bottom. Fig. 9. Tethea cliftoni Bowerbank, and distribution of marine fauna. Pp. 43-68 in G.W. dry holotype, BMNH unregistered (Box D.A. 38), lateral

Dyne (ed.) /'anna of Australia . Vol. I A General Arti- view, xO.64. Note the long pedicels rising from a com- cles. Australian Government Publishing Service: posite cemented cobble, and Bowerbank's autograph Canberra. label. Wilson, H.V., 1925. Silicious and horny sponges collected Figs 7, X. Polymastia crassa Carter. Fig. 7. NMV by the U.S. Fisheries steamer "Albatross" during the F51945, before preservation, xO.38. Fig. 8. NMV Philippine Expedition, 1907-10. In: Contributions 1.51946, xO.73. to the biology of the Philippine Archipelago and adja- figs 10, II. Spiraslrel/a papillosa Ridley and Dendy. cent regions. U.S. Natn. Mus. Bull. 100(2/4): i-vii, Fig. 10. NMV F51948, x().32. Fig. II. NMV F51949, 273-532, pis 37-52. xO.38.

Wisniowski, T., 1888. Beitrag /Air Kenntniss der Mikrofauna aus den oberjurassisehen Feuer- Plate 3 steinknollcn der Umgcgcnd von Krakau. Jb. K.K. lies 1-6. Chondrosia reticulata (Carter). Fig. I. NMV Ceol. Reichsanst. Wien 38: 657-679, pis 12, 13. F5I951, x0.76. Figs 2, 3. Chondrosia spurca Carter, wet Woodburne, M.O., and Zinsmcister, W..I., 1984. The first holotype BMNH 1886. 12.15.128, top and sectioned land mammal from Antarctica and its biogeographic bottom, x().4. Figs 4, 5. Two wet syntypes or two halves implications. J. Paleont. 58 (4): 913-948. of one syntype of Renicra co/leclrix Lendenfeld, AM Zinsmeister, W.J., 1982. Late Cretaceous-Early Tertiary G9014. Fig. 4. xO.38. Fig. 5. x().43. Fig. 6a-e. Wet dis- molluscan biogeography of (he southern Circum- sected syntype of C 'hondrosia co/leclrix Lendenfeld, AM Pacific. J. Paleont. 84-102. 56(1): C,906l,"x0.76. Fig. 7. Lalrunculiticonulosa Hallmann, NMV F5I954 Plales 1-38 before preservation, X0.49. Figs 8- 10. Ixtlrunculia hallmanni sp. nov. Fig. 8. Holo- Unless otherwise stated, all scales with specimens (pis type, NMV F51953, xO.43. Fig. 9. Paratype, NMV 1-17) represent 2 cm. All scales with photomicrographs 151956, x0.6. Fig. 10. Paratype, NMV F5I957, x0.76. (pis 18-38) represent 200 ,un. Fig, II. Pseudaxinella decipiens sp. nov. Holotype, NMV F51961. x0.96.

Plate 1 Plate 4 Fig. 1. Corlicium candelabrum Schmidt, NMV F5I927, X0.87. fig. 1. Reniochalina sectilis sp. nov. Holotype, NMV before Fig. 2. Ancorina geodides Carter, NMV F51929 I 51962, x 1.04. preservation, x0.46. Fig. 2a, b. Reniochalina lamella Lendenfeld. Dry Fig. 3. Ancorina repens sp. nov. Holotype, NMV schizosyniypc, BMNH 1925. 12.1.18, Hast Coast, X0.78. F51930, x0.52. Lendenfeld's sketch of the discarded specimen shows the 0.52. the schi/.otype. Fig. 4. Ancorina robusta Carter, NMV F5 1931 , x original position (hatched) of Fig. 5. Ancorina suina sp. nov. Holotype, NMV Fig. 3a, b. Reniochalina stalaginitis Lendenfeld. Dry F51934, xO.87. syntype, BMNH 1887.4.27.122 (with autograph label), Fig. 6. Stelletla arenitecta sp. nov. Paratype, NMV x0.26 Fig. 4. Reniochalina spiculosa Lendenfeld ms. Dry F51942, xO.53. specimen, Port Jackson, BMNH 1887.4.27.119, xO.38. Fig. 7. Rhubdastrella cordatu sp. nov. Holotype, NMV Fig. 5a, b. Reniochalina arborea Lendenfeld ms. Dry Zealand, 1887.4.27.117, xO.46. F51935, (left), and Tedania anhelans (Lieberkiihn), NMV specimen. New BMNH lamella. F520I3 (right), photographed in situ. x0.64. Lendenfeld's sketch as for R. 6-12. cacliformis (Carter). Fig. 6. Fig. 8. Rhubdastrella intermedia sp. nov. Holotype, Figs Rhaphoxyu NMV F51936. x().64. NMV F5 1963, X 0.6. Figs 7, 8. Acanthellina rugolineata Carter, wet syntype, BMNH 1886.12.15.94. Fig. 7. Side, Plate 2 x().49. Fig. 8. Top, x().6. Fig. 9. Acanthellina par viconulata Carter, wet holotype, BMNH 1886.12.15.56, Fig. 1. Geodia {Geodia) punctata Hentschel. NMV Figs 10, II. Acanthe/la hirciniopsis Carter, wet F51937, xO.46. xO.35. 182 F. WIEDENMAYER

schizosyntype (part of base of large specimen, cut sagi- Figs 8, 9. Phoriospongia carcinophila (Lendenfeld). tally and transversally), x0.32. Fig. 10. View of sagittal Fig. 8. NMV F5 1990, x0.7. Fig. 9. NMVF51991, xl.7. section showing mesial condensation. Fig. 12. Acanlhelki Figs 10-12. Phoriospongia kirki (Bowerbank). Fig. 10. hirciniopsis Carter, wet syntype, BMNH 1886.12.15.48, Dysidea kirki Bowerbank, dry syntype?, Bowerbank Col- X0.46. lection, BMNH 1872.5.21.50, Swan River (Perth), W.A.,

x 0.48. Fig. 1 1 . Dysidea favosa Marshall, portion of wet Plate 5 holotype, PMJ Porif. 125, xO.55. Fig. 12. NMV F51993, Figs 1-3. Rhaphoxya cuctiformis (Carter), dry speci- XO.64.

mens. Figs 1, 2. Acanthellina rugoslincata Carter, syn- Fig. 13. Psainmocleina bitexlum sp. nov. Holotype,

types. Fig. 1. BMNH 1886.12.15.444, xO.32. Fig.' 2, NMV F51996, X0.72. BMNH 1886.12.15.365, xO.53 (label, except register Plate 8 number, in Carter's hand). Fig. 3. Acanlhelki cactiformis Carter, paraleclotype, BMNH 1884.10.10.2, x0.15. Note 1 Fig. . Psainmocleina callosum (Marshall). NMV the similarity (as for Fig. I) with Spongia cardials F5I997, xl.l. Lamarck, redescribed as Acanlhella by Topsent (1930: Figs 2-4. Psainmocleina densum (Marshall). Fig. 2. 40), assigned to Phakeliia by Bergquist (1970). NMV F51998. xO.55. Fig. 3. NMV 52000, xl.l. Fig. Fig. 4. Rhaphoxya fe/ina sp. nov. Holotvpe, NMV 4. Psammopemma densum Marshall, wet holotype, ZMB

I 51964, X0.53. 2631, x2.l. Scale 1 cm. Fig. 5 Truchvcladus laevispirulifer Carter. NMV F Fig, 5. Psainmocleina Jissuratum sp. nov. Holotvpe, 51965, xO.67. NMV F52002, xl.04. Figs 6, 7. Clathriodendron cacticulis (Carter). Fig. 6. Figs 6, 7. Psainmocleina goniodes sp. nov. Fig. 6. Para- NMV F51966, x0.43. Fig. 7. Dictyocyllndrus coctkutb type, NMV F52059, x 2. /scale 1 Fig. 7. 1 cm. Holotype, Carter, wet holotvpe, BMNH 1886.12.15.120, xO.58. NMV F52060, xO.81. Fig. 8. Clalhrla transient Kallmann. NMV F51980, I ijis 8- 1 2. Psainmocleina nodosum (Carter). Figs 8-11. x0.46. Sarcocornea nodosa Carter, dry types. Figs 8-10. Per- Fig. 9. Clathria wilsoni sp. nov. Holotvpe. NMV pendicularly cut paraleclotype, BMNH 1886.12.15.297. F51967, xl.4. Fig. 8. Top view, xl.02. Fig. 9. Interior, xO.67. Fig. Fig. 10, I chinoclthria favus Carter. NMV F51972, 10. Side view, xO.52. Fig. II. Lectotype, BNMH X0.53. 1886.12.15.295, xO.38. Fig. 12. NMV F52003, xO.38. Fig. 11. Echinoclulhria laminaefavosa (Carter). NMV Plate 9 1 51971, xO.39.

Figs Plate 6 1, 2. Psammoclema rudiatum sp.nov. Holotvpe,

NMV F52004. x 0.46. Fig. 1 . Side view. Fig. 2. Top view. Fig. 1. Echinoclathria leporina (Lamarck). NMV Figs 3-6. Psammoclema ramosum Marshall. Fig. 3. F51974, detail from central portion, before preservation, NMV F52005, top, x 0.48. Figs 4-6. Fragments from wet xO.35. holotype, PMJ Porif. 134. Fig. 4. Top, X0.78. Fig. 5. Figs 2-5, 7. Ecfiinoclathria globosa (Lendenfeld), Fig. Longitudinal section, x 1.13. Fig. 6. Bottom, x0.78. 2. NMV F51975, x0.87. Fig. 3. NMV F51976, x0.78. Fig. 7. Psammoclema stipitatum sp.nov. Holotype, Fig. 4. Huline globosa Lendenfeld, only extant svntype, NMV F52006, xO.58. wet BMNH 1886.8.27.71, Port Phillip Bay, X 0.81. Fig. Figs 8-10, 13. Mycale (Arenochalina) mirabilis Len- 5. Halme micropora Lendenfeld, dry syntype, AMG882S, denfeld. Fig. 8. NMV F52017, xO.58. Figs 9, 10. NMV lllawarra, Fig. 7. X0.38. Holopsumma crassu Carter, drv F52015. Fig. 9. Preserved specimen, xO.54. Fig. 10. syntype, BMNH 1886.12.15.313, S. coast, x 0.49. Before preservation, x 0.35. Fig. 13. Sectioned dry holo- Fig. 6. Phoriospongia reticulum Marshall, wet holotype, BMNH 1886.8.27.587, xO.67. type, ZMB 2634, xl.4. Figs 11, 12. Psammoclema vansoesti sp.nov. Fig. 11. Fig. 8. Echinoclathria sp. nov. Holotype, egena NMV Holotype, NMV F52008, X0.43. Fig. 12. Paratype, NMV F 5 1978, xO.61. F52007, X0.32. Fig. 9. Echinoclathria tubulosa (Hallmann). NMV F51979, xO.76. Plate 10 Figs 10, 11. Strongylacidon stelliderma (Carter), Fig. Figs 1, 2. Mycale (Arenochalina) mirabilis Lendenfeld. 10. NMV F51981, X 1.6. Fig. II. Halichondna stelliderma NMV F52016. Fig.l. Before preservation, X0.34. Fig. Carter, wet holotype, BMNH 1886.12.15.148, xO.67. 2. Preserved specimen, xO.64. Figs 3-7. Plate 7 Tedania anhelans (Lieberkuhn). Fig. 3. NMV l'52012, X0.56. Figs 4-6. Wet Hypotypes of Tedania Figs Stylotella 1,2. inaequalis (Hentschel). Fig. 1 . NMV digitata: Dendy (1895). Fig. 4. NMV RN796, Sorrento 151983, x2J6. Fig. 2. NMV F51988 xl.04. jetty, x 0.67. Figs 5, 6. NMV RN871 . Fig. 5. Interior with Figs 3-7. Phoriospongia argentea (Marshall). Fig. 3. embryos, xO.57. Fig. 6. Top, xO.56. Fig. 7. Tedania NMV F51989, x 1.13. Fig. 4. Dysidea argentea Marshall, rubicunda Lendenfeld, wet syntype, AM G9146, x0.46. wet holotype, PMJ Porif. 136.X 1.57. Figs 5-7. Chon- Figs 8-11. Eorcepia biceps (Carter). Figs 8, 9. NMV dropsis arenifera Carter, wet syntypes. Figs 5, 6. BMNH F52018. Fig. 8. Top, xO.35. Fig. 9. Transverse fracture, 1886.12.15.149. Fig. 5. Exterior, xO.52. Fig. 6. Interior, x 0.93. Fig. 10. Eorcepia colonensis: Carter (1885a), dry X0.54. Fig. 7. 1886.12.15.54, BMNH xO.64. hypotype with Carter's label, BNMH 1886.12.15.363, .

SPONGES FROM BASS STRAIT 183

x0.46. Fig. 11. Suberites biceps Carter, wet holotype, below, and from one side, respectively, x0.45. Figs 11, BMNH 1886.12.15.52, Port Phillip Heads, x().5. 12. Dry holotype, BMNH 1886.8.27.32, Torres Strait. Fig. Fig. 12a, b. Lissodendoryx isodictyalis (Carter). NMV 11. From above, x0.46. Fig. 12. From below, x0.41. F52019, xl.04. Plate 14

Plate II Figs 1-5. Carteriospongla silicata (Lendenfeld). Dry

Figs 1, 2. Myxilla isodictyalis: Dendy (1896), wet hypo- hypotypes of Carteriospongia elegans: Burton type, NMV RN778, Sorrento jetty. Fig. I. Top, xu.71. (1934a:600). Note that the (correct) identification on the Fig. 2. Transverse section, xO.74. labels departs from the published one. Figs 1, 2. BMNH Figs 3, 4. Ectyodoryx metadata Hentschel. Fig. 3. NMV 1893.3.4.24, Great Barrier Reef, bottom and top, respec- F52021, X0.78. Fig. 4. NMV F52020, xO.93. tively, xO.32. Figs 3-5. BMNH 1893.3.4.63, Bass Strait, Figs 5a, b. Phorbas cf. (enactor (Topsent). NMV side, top and bottom, respectively. Fig. 3. x0.55. Fig. F52022, x0.9. 4. xO.46. Fig. 5. xO.38. Fig. 6. Callvspongia asparagus (Lamarck). NMV Fig. 6. Carteriospongia cf. vermicularis (Lendenfeld). F52024. xO.93. NMV F52041, x 1.13. Fig. 7. Callyspongia bilamellata (Lamarck). Dry hypo- Figs 7, 9. Coscinodermu pesleonis (Lamarck). Fig. 7. type of Carter (1885d, as Cavochulinu bilamellata), NMV F52042, xO.55. Fig. 9. NMV F52044 in situ, BMNH 1886.12.15.184, Westcrnport Bay, Victoria, xO.58. xO.13 (scale = 5 cm). Figs 8, 10. Leiosella caliculata Lendenfeld. Fig. 8. NMV Figs 8-10. Callyspongia diffusa (Ridley). Fig. 8. NMV F52046, xO.46. Fig. 10. Euspongia compacts Carter. Dry xO.46. F52026, xO.32. Fig. 9. Cladochulina diffusa Ridley, dry leclotype, BMNFM877. 5.21. 1899, "S. Australia", paralectotype, BMNH 1882.10.17.6, Singapore, x().33. Plate 15 Fig. 10. Pachvchalina manus Lendenfeld, wet syntype, xb.33. AM G8927, 1 ins 1-5. Leiosella caliculata Lendenfeld. Fig. I. Dry Fig. II. Callyspongia ramosa (Gray). NMV F52028, syntype, BMNH 1955.4.7.1, x().45. Figs 2, 3. Euspon- xO.75. gia infundibuliformis Carter, dry leetotype, BMNH Plate 12 1886.12.15.222, figured in Lendenfeld (1889b, pi. 13, fig. 7) as Leiosella compacta. Fig. 2. Concavity, x 0.28. Fig. Figs 1-3. Callyspongia pergumenlacea (Ridley). Fig. 3. Detail of concavity and inner rim, x 1.04. Figs 4, 5. 2. Chalinopora paucispina 1. NMV F52027, xO.49. Fig. Geelongia vasifonnis: Dendy rns., dry composite speci- (syntype?), Lendenfeld, dry holotype BMNH men from Victoria, BMNH 1925. 11.1.887, x().2l. Fig. Queenscliff, Victoria, x0.46. Fig. 3. 1886.8.27.413, 4. Open side, showing the partition formed by the wall dry syntype, BMNH Chalinopora typica Lendenfeld, of the almost complete primary vase in the background, Victoria, xO.34. 1886.8.27.411, Port Phillip Bay, and the half-formed secondary vase. Pig. 5. Convex side. serpentina (Lamarck). NMV Fig. 4. Callyspongia Figs 6, 7. Leiosella levis (Lendenfeld). Fig. 6. NMV F52029, xO.29. F52047, xO.35. Fig. 7. One of the misidentificd dry Fig. 5. Callyspongia loxifera sp.nov. Holotype, NMV paralectotypes of Euspongia compacta Carter, BMNFI

F52032, x 1 .08. 1 877.5.21 .1900, Port Fairy, Victoria, here assigned to nov. Holotype Fig. 6. Callyspongia persculpta sp. Leiosella levis, xO.32. (part), NMV F52035, xO.57. Figs 8-10. Ircinia caliculata Lendenfeld. Figs 8, 9, NMV 152036, Fig. 7. Gelliodes incrustans Dendy. NMV F52049. Fig. 8. Open concavity, xO.32. Fig. 9. Convex attached to Jaspis stellifera, x 1 . 1 side, xO.29. Fig. 10. Open concavity of dry syntype, NMVF52037, Fig. 8. Oceanapia putridosa (Lamarck). BMNH 1886.8.27.358, with I.endenfeld's label. x().35. X0.38. Figs 9-12. Spongia hispida Lamarck. Fig. 9. NMV Plate 16 F52038, x 0.355. Fig. 10. NMV F52039, x().77. Fig. II. Figs 1-3. thorecta ehoanoides (Bowerbank). Fig. 1. Lendenfeld, wet syn- Euspongia irregularis var . jacksonia F52050, xO.26. Figs 2, 3. Stelospongus flabel- Euspongia irregularis NMV type, AM Z498, x.0.41. Fig. 12. paralectotype, liformis lutus Carter, misidentificd dry silicata Lendenfeld, wet syntype, AM G3962, xO.6. var. BMNH 1886.12.15.219. Fig. 2. Narrow side, with Len- denfeld's identification "Thorecta exemplum var. secunda" Plate 13 on the back of Carter's label, xO.27. Fig. 3. Broadside, Figs 1-7. Carleriospongiu caliciformis Carter. Fig. I. X0.21. 5. Thorecta glomerosus sp. nov. Holotype, NMV F52040b, x0.43. Figs 2, 3, 6, 7. Dry syntypes. Figs Figs 4, Underside, with 152052. Fig. 4. Before preservation, 0.55. Fig. 5. 2. 3. BMNH 1886. 12.15.214. Fig. 2. NMV specimen, X0.48. labels by Carter (right) and Lendenfeld (left), x 0.3. Fig. Preserved 1886.12.15.215, Figs 6, 7, 10. Thorecta lubercu/atus (Carter). Figs 6, 3. Concavity, xO.35. Figs 6, 7. BMNH holotype, BMNH 1886.12.15.446. Fig. 6. xO.46. X0.23. Fig. 6. Underside. Fig. 7. Concavity. Figs 4, 5. 7. Dry Detail of opposite side, X 1.04. Fig. 10. NMV NMV F52040a. Fig. 4. Detail of concavity and inner rim, Fig. 7. xO.68. X0.56. Fig. 5. Lateral view, x().29. F52053, Figs 8. Aplysttla rosea (Barrois). NMV F52061, x 1.02. Figs 8-12. Carteriospongia silicata (Lendmfe\d). Fig. from 9 Dvsidea avara (Schmidt). NMV F52057, xl.02. 8-10. NMV F52048, before preservation, from top, Fig 184 F. WIBDENMAYF.R

Fig. 11. Faseiospongia rintosa (Lamarck)- NMV Plalc 21 F52056, XO.33. Transverse sections, xO.29. Fig. 12. Darwinella australiensis Carter- NMV F52064, Fig. I. Tetilla praecipua sp.nov. Holotype, NMV xl.04. F51943, choanosome. Plalc 17 Fig. 2. Tethya ingalli Bowerbank, NMV F5I944, cortex. xl Figs 1, 2. Darwinella australiensis Carter. Fig. I. NMV Fig. 3-5. Polymastia crassa Carter. Figs 3, 4. Cortex F52063, xO.93. Fig. 2. Four dissected wet syntypes, and peripheral choanosome of main body. Fig. 3. NMV BMNH 1887. 7.11.10, xO.45. F51945. Fig. 4. NMV F5I946. Fig. 5. Wall of fistule in Figs 3, 7. Darwinella gardmeri Topsent. Fig. 3. NMV NMV F51946. F52066, before preservation, xO.47. Fig. 7. NMV tig. 6. Spirastrella papillosa Ridley and Dendy. NMV F52065, xO.98. F51948, peripheral chaonosomc and inner part of cortex

Figs 4, 5. Aplysina lendcnfeldi Bergquisl. Fig. 4. NMV (left). F52071, xO,58. Fig. 5. NMV FS2072, before preserva- Plalc 22 tion, x0.3.

Figs 6, 8, 9, 11. Pseudoceratine durissimaC&clex. Fig. Transverse sections. 6. F52073, x0.9. Fig. 8. Dry syntvpe, NMV BMNFI Fig, 1. Chondrosia reticulata (Carter). NMV F5195I, 1886.12.15.261, 9. x().4l. Fig. NMV F52074, x0.93.Fig. lower cortex (left) with crypt and peripheral choanosome, 11. NMV F52075, x0.56. x87. Note pervasive microsymbionts and digitate alga Fig. 10. Aplvsina ianlhelliformis (Lendenfeld). NMV on left side of lower sand grain. 1 52070, xO.32. Figs 2, 3. Latrunculia conulosa Hallmann. Cortex and Fig. 12. Druinella rotunda Lendenfeld. NMV F52078, peripheral chaonosomc, x29. Fig. 2. NMV F51953. Fig. before preservation, x 0.39. 3. NMV F51952. Plate 18 Figs 4, 5. Latrunculia hallmanni sp.n. Paratype, NMVF51956. Fig. 4. Cortex and peripheral choanosome, Perpendicular sections. x 29. Fig. 5. Peripheral choanosome, chiefly with small,

Fig. I . Corticium candelabrum Schmidt, NMV F51928, irregular sanidasters, x 87. A small portion of the cortex x 29. Surface above. is visible above. Carter, F 5 Figs 2, 3. Ancorina geodtdes NMV 1929, tig. 6. Pseudaxmella decipiens sp.nov. Holotype, NMV x29. Fig. 2. Peripheral choanosome with crypts, part of F51961, choanosome, x29. inner cortex (above). Fig. 3. Inner cortex with chones. Figs 4-6. Ancorina repens sp.nov. Holotype. NMV Plate 23 Fig. 4. Cortex with surface (above), x29. Fig. F51930. Transverse sections, x29. 5. Continuation inward from the bottom of Fig. 4, with Fig. 1. Reniochalinu sectilis sp.nov. Holotype, NMV innermost cortex, zone of crypts, and peripheral F51962. Ectosomal skeleton (upper half) and main xO.29. 6. cortex (surface choanomsome, Fig. Outer skeleton. above) with chone, x 100. Fig. 2. Rhuphoxya cactiformis (Carter). NMV F51963, Plalc 19 mesial skeleton. Figs 3, 4. Rhaphoxya felina sp.nov. Holotype, NMV Perpendicular sections. F51964. Fig. 3. Choanosome. Fig. 4. Erect lamella at sur- Figs 1, 2. Ancorina repens sp. nov. Holotype, NMV face with chiefly penicillata arrangement of megascleres. Fig. 1. of with oxyspherasters. F51930. Wall chone x87. Fig. 5. Clathriodendron cacticutis (Carter). NMV Fig. 2. Choanosome, x87. F51966, mesial choanosome.

Fig. 3. Ancorina robusta Carter. , NMV F51931 cortex Fig. 6. Clathria transiens Hallmann. NMV F51980, and peripheral choanosome, x29. peripheral choanosome. Figs 4-6. Ancorina suina sp.nov. Holotype, NMV F5I034. Cortex and peripheral choanosome, x29. The Plate 24 surface faces left in Fig. 5, upwards in Fig. 4, of which Fig. 1. Clathria wilsoni sp. nov. Holotype, NMV Fig. 6 is the continuation inwards. F51967, peripheralchoanosome, x29. The surface with Plalc 20 the paratangential open penicils is folded over into the plane of the section in the left half. Figs 1-5. Perpendicular sections through surface, cortex Fig. 2. Echinoclathria favus Carter. NMV F51972, and peripheral choanosome, x 29. transverse section through lamella, x29. Fig. 1. Rhabdaslrella cordata, sp. nov. Holotype, NMV Fig. 3. Phoriospongia reticulum Marshall. Holotype, 151935. ZMB 2634. Teased portion of choanosome, x87, show- Fig. 2. Rhabdaslrella intermedia sp. nov. Holotype, ing arrangement of sigmata. NMV F51936. Figs 4, 5. Echinoclathria leporina (Lamarck). Trans- Figs 3, 4. Geodia punctata Hentschel, NMV F51937. verse sections through fronds, x29. Fig. 4. NMV F51974. Fig. 3. Solid cortex. Fig. 4. Lacunose cortex. Fig. 5. NMV F51973, apical edge. figs 5, 6. Juspis stellifera (Carter), NMV F51938. Fig. Fig. 6. Echinoclathria egena sp.nov. Holotype, NMV 5. Surface facing left. Fig. 6. Dissociated megascleres, F51978. Peripheral choanosome with embryos, surface x87. on top. x29. SPONGES FROM BASS STRAIT 185

Plate 25 fibre, and the transverse spicules protruding from the Transverse sections latter, with partial spongin envelopes. Fig. 3. Tapering free end of primary fibre at surface, x 29. Fig. 4. Filamen- Figs 1, 2. Echinoclathria egena sp.nov. Holotype, NMV tous algae in primary fibre, x87. F51978. Fig. 1. Deeper ehoanosome, x29. Fig. 2. Figs 5, 6. Tedania anhelans (Lieberkuhn), perpendic- Peripheral ehoanosome with surface in upper right, x 87. ular sections. Fig. 5. NMV F52010, x 29 Figs 3, 4. Echinoclathria tubulosa (Hallmann). NMV (surface on top). Fig. 6. NMV F52009, x 29 (surface in upper left corner). F51979, x29. Fig. 3. Deeper ehoanosome. Fig. 4. Peripheral ehoanosome. Plate 30 Figs 5, 6. Strongylacidon stelliderma (Carter). NMV F51981, x29. Fig. 5. Deeper ehoanosome. Fig. 6. Fig. 1. Tedania anhelans (Lieberkuhn). NMV F51940, Peripheral ehoanosome with conules. Transverse section with surface, x 29. Fig. 2. Forcepia biceps (Carter). F52018, trans- Plate 26 NMV verse section, x 29. Ectosome on top with megascleres Transverse sections in paratangential arrangement, in oblique view.

Figs 1-4. Stylolelia inaequalis (Hentschel). Fig. 1. NMV Fig. 3. Lissodendoryx isodictyalis (Carter). NMV F51984, mixed skeletal structure, single sand grains, mac- F52019, transverse section with surface, x29. rosymbionts, X29. Fig. 2. NMV F51983, sandy enclaves, Fig. 4. Phorbas cf. tenacior (Topsent). NMV F52022, scarce spicular whisps, x 29. Fig. 3. NMV F51987, dense transverse section with surface, x29. skeletal structure, x 29. Fig. 4. NMV F51987, surface with Fig. 5. Ectyodoryx maculata Hentschel. NMV F52020, paratangential layer of megacleres, x87. deeper ehoanosome, X29. Fig. 5. Phoriospongia argentea (Marshall), NMV F5I989, x 29. Note small single tallophytous alga below Plate 31 surface. Fig. 1. Ectyodoryx maculata Hentschel. NMV F52020, Fig. 6. Phoriospongia carcinophila (Lendenfeld), NMV perpendicular section through peripheral ehoanosome and F51992, x29. ectosome, x29. Plate 27 Figs 2, 3. Callyspongia asparagus (Lamarck), NMV F52024. Fig. 2. Perpendicular section with ectosome. Transverse sections with ectosome, x29. X29. Fig. 3. Tangential section of ectosome, x29. Figs 1, 2. Phoriospongia carcinophila (Lendenfeld). Figs 4, 5. Callyspongia diffusa (Ridley and Dendy), Fig. I. NMV F5199I . Fig. 2. NMV F51990. Note the felt NMV F52026. Fig. 4. Tangential section of ectosome, of proper spicules at and below the surface. x29. Fig. 5. Perpendicular section with ectosome, x29. Fig. 3. Psammoclema bitextum sp. nov. Holotype, Fig. 6. Callyspongia pergamentacea (Ridley). NMV NMV F51996. F52027, perpendicular section with ectosome, x 29. Fig. 4. Psammoclema callosum (Marshall). NMV F51997. Plate 32 Fig. 5. Psammoclema densum (Marshall). NMV

Fig. I . Callyspongia F5I998. Note the crowded foreign spicule fragments at pergamentacea (Ridley). NMV the surface. F52027. Ectosome, Tangential section, x 29. Figs 2-4. Callyspongia serpentina Fig. 6. Psammoclema nodosum (Carter). NMV F52003. (Lamarck). NMV F52029, perpendicular sections Note infolded ectosome (upper right), lined canal (lower with ectosome. Fig. 2. x29. Fig. 3. x82. Fig. 4. x87. center), ascending column with detritus, and algal filaments (lower right). Figs 5, 6. Callyspongia toxifera sp.nov., paratypes, transverse sections with ectosome, X29. Fig. 5. NMV Plate 28 F52030. Fig. 6. NMV F5203I.

Figs 1, 2. Psammoclema goniodes sp.nov. Transverse Plate 33 sections, x29. Fig. I. Paratype, NMV F52059 (surface above). Fig. 2. Holotype, NMV F52060. Surface upper Figs 1-3. Callyspongia persculpta sp.nov. Holotype, right. NMV F52035, x29. Fig. 1. Ectosome, tangential section.

Figs 3, 4. Psammoclema ramosum Marshall. NMV Figs 2, 3. Transverse sections with ectosome. Fig. 3 shows F52005, X29. Fig. 3. Longitudinal section, surface left. the edge of one of the hollow lobes. Fig. 4. Transverse section. Figs 4-6. Gelliodes incrustans Dendy, NMV F52036, Fig. 5. Psammoclema stipitatum sp.nov. Holotype, transverse sections. Figs 4, 5. Ectosome on top, x 29. Fig. NMV F52006, ehoanosome, transverse section, x29. 6. Deeper ehoanosome, x 87. Fig. 6. Psammoclema vansoesli sp.nov. Paratype, Plate 34 NMV F52007, transverse section with surface, x 29. Fig. 1. Gelliodes incrustans Dendy. NMV F52036, Plate 29 deeper ehoanosome, x87. Figs 1-4. Mycate (Arenochalina) mirahilis Lendenfeld. Fig. 2. Spongia hispida Lamarck. NMV F52039, trans-

Fig. 1. NMV F520I4, peripheral skeleton (surface above). verse section with ectosome, x 29. x29. Fig. 2. Junction of secondary and primary fibre, Fig. 3. Carteriospongia caliciformis Carter. NMV x 87. Note stratification, filamentous algae in primary F52040a, transverse section with ectosome, X29. 1X6 F. WIEDENMAYER

inhalant sieve, Fig. Fig. 4. Carteriospongia cf . vermicularis (Lendenfeld). 4. NMV 52071, ectosome wilh x29. NMV F52041, transverse section wilh eclosome. x 29. 5. NMV F5207I, choanosome with transversely and obli- Figs 5, 6. frcinia caliculata Lendenfeld. NMV F52049. quely cut fibres, x29. Fig. 6. NMVF52071, fibre, x 87. Fig. 5. Transverse section with choanosome, x29. Fig. 6. Choanosome with filaments. Inflated ends of the latter Plale 37 are visible near the upper left corner and halfway down Fig. 1. Aplysina lendenfeldi Bergquist. NMV F52072, near the right margin. choanosome with gemmules, x87. Plate 35 Figs 2-6. Pseudoceratina durissima Carter. Fig. 2. NMV F52074, perpendicular section with ectosome Figs l, 2. Thorecta choanoides (Bowerbank). NMV (above) and peripheral choanosome, x29. Note 1-52050, x29. Fig. 1. Transverse section through eclo- reticulate-septate canal. Fig. 3. NMV F52074, choanosome and peripheral choanosome, the former wilh tran- some with transversally cut fibre, x87. Fig. 4. NMV sition from a superficial elevation (above) to an inhalant F'52074, crowded choanocyte chambers, x 87. Figs 5, 6. depression (below). Fig. 2. Sieve of inhalant depression NMV F52075 and F52076, respectively, longitudinal sec- in ectosome. lions of fibres, x 87. Note irregularly lobose, partly com- Figs 3-5, Thorecta tuberculatus (Carter). Transverse posite pith, and, in Fig. 5, density of microsymbionts. sections. Fig. 3, NMV F52054, eclosome and peripheral choanosome, x 29. Figs 4, 5, NMV F52053. Fig. 4. Plate 38 Peripheral choanosome, x29. Fig. 5. Fibrous-stratfied /one between sandy eclosome and choanosome, x87. Figs 1-5. Druinella rotunda Lendenfeld. ZMB 10403, Fig. 6. Dysidea avara (Schmidt). NMV F52057, choano- one of Lendcnfeld's slides with microtome sections from some with fibre, x 29. the holotype. Fig. 1. Oblique section of fibre, x29. A

transversally cut canal of type 1 is situated near the middle Plaie 36 of the upper margin. Fig. 2. Choanosome, x 29, over-

Transverse sections lapping with the upper margin of Fig. 1 . Canals of type

Fig. 1. Darwinella australiensis Carter . NMV F52064. 2 chiefly in left half, canals of type I in right half. Choanosome, x 29. Choanocyte chambers are confined to the darker mean- Fig. 2. Aplysina iantheltiformis (Lendenfeld). NMV dering fields. Figs 3, 4. Transverse sections of fibres, x 87.

1' 52070. Choanosome, x 29. Fig. 5. Choanosome, x29. Figs 3-6, Aplysina lendenfeldi Bergquist. Fig. 3. NMV Fig. 6. Phoriospongia argentea (Marshall). NMV 152072, ectosome and peripheral choanosome, x29. Fig. 1-5 1989, transverse section with ectosome, x 29. SPONGES FROM BASS STRAIT 187

PLATE 1 F. WIEDENMAYER

PLATE 2 SPONGES FROM BASS STRAIT 189

PLATE 3 190 WIEDENMAYER

4m ^ 5b

*%%.«'

PLATE 4 SPONGES FROM BASS STRAIT 191

PLATE 5 .

192 F. WIEDENMAYER

: i- #t.

PLATE 6 SPONGES FROM BASS STRAIT 193

PLATE 7 194 F. W1EDENMAYER

PLATE 8 SPONGES FROM BASS STRAIT 195

PLATE 9 196 F. W1EDENMAYER

PLATE 10 SPONGES FROM BASS STRAIT 197

PLATE 11 198 F. WIEDENMAYER

PLATE 12 SPONGES FROM BASS STRAIT 199

PLATE 13 200 F. WIEDENMAYER SPONGES FROM BASS STRAIT 201

PLATE 15 202 F. WIEDENMAYER

PLATE 16 SPONGES FROM BASS STRAIT 203

PLATE 17 204 F. WIEDENMAYER 1

m& PI

3 i i

PLATE 18 SPONGES FROM BASS STRAIT 205

t t Mli!

•V

PLATE 19 206 F. WIEDENMAYER

PLATE 20 SPONGES FROM BASS STRAIT 207

PLATE 21 K .'OS F. WII'.DI'.NMAYI

PLATE 22 I

si'ONtil'S l'KOM BASS STRAI 109

I ., I

V. \ x/

3 i i

PLATE 23 .

210 F. WIEDENMAYER

4 i

«* « ^" 4*

r .

E I

PLATE 24 SPONGUS FROM BASS STRAIT 211

* •

\|A 'lj MMt 4

PLATE 25 212 F. WIEDKNMAYER

%*;•*• -

1 l i

€

a«*r' jiriitf •

i-5*

PLATE 26 SPONGES 1ROM BASS STRAIT 213

PLATE 27 214 F. WIEDENMAYER

I 4 i i

PLATE 28 SPONGES FROM BASS STRAIT 215

PLATE 29 216 F. WIEDENMAYER

' ( ,.

-» '.

*~ •* 1

PLATE 30 SPONGES FROM BASS STRAIT 217

f- -- "T *

J [

PLATE 31 21K F. wii:ni;NMAYi;R

~9J*r If * -t^fa

k '

'h.

* *•*-

PLATE 32 1

SPONGES FROM BASS STRA1 219

t.. *%

w "

PLATE 33 220 F. W1EDENMAYER

1 L

PLATE 34 SPONGES FROM BASS STRAIT 221

'"'» *:\ 1W "

*7^ V *

PLATE 35 222 F. WIEDENMAYER

PLATE 36 SPONGES FROM BASS STRAIT 223

2 »'•>

PLATE 37 224 F. WIEDENMAYER

i 2 i

PLATE 38 SPONGliS FROM BASS STRAIT 225

Index A. fictitioides 102 Names of the species-and genus-groups in the A. paupertas 96

1 systematic-descriptive part. Italics denote valid A . tenacior 02 names and combinations; roman type is for invalid Ancorina 14, 65 names, misidentifications, genera inquirenda, and A . acervus 17, 18 species inquirendae. Question marks behind names A. alala 18 in italics commonly mean that the validity, iden- A . auslruliensis 18 tification, or combination is doubtful. Genera in A. corticata 18, 19 roman type with question mark are doubtful as syn- A. geodides 14, 18, 19

onyms. New species are in bold type. A . murrayi 16, 18

A . progressa 18

abruptispicula, Cladochalina nuda var. ... 112 A . repcns 16 abyssi, Dendoricella 95 A. robusla 17, 18 abyssi, Desmaeidon 95 A. slulagmoides 18 Acamas 83, 84 A. suina 18, 19 Acamasina 84 angulata, Chalina polychotoma var 115 acantha, Dendrillu 153 anhelans forma assabensis, Tedania 90 acanthastrum, Corlicium 13, 14 anlteluns, Halichondria 87 Acanthella: Carter 50 anhelans, Myxilla 87 A. cactiformis 50 anhelans, Tedania 87, 89, 92, 96, 100 A. hirciniopsis 51 antarclica, Dendrillu 153 A. stipitata 51 Anthastra 19, 20 Acanthellina 50 Antherochalina 56 A. parviconulata 51 A. perforata 62 A. rugolineata 51 A. lenuispina 62 acanthosanidastera, Latrunculia (Negombatu) 40 Antheroplax (see Phyllospongia) 137 acanthosanidastera, Oxylatrunculia 40 Aphrodite 121

Acervochalina finitima: Ridley 108 Aplysia 1 54 acervus, Ancorina 17, 18 Aplysilla 146, 147, 152, 153 acustella, Clionct 39 A. cactos 152

Aegagropila 83, 84 A . glacialis 1 46

aerophoba, Dendrillu ? 152, 153 A . rosea 146, 1 47

affinis, Pachychalina 113 A. rosea: Rockman 1 52 agminatu, Stylotella 71 A. sulfurea 146, 147 Aiolochroia 147, 156 A. sulphurea 146

A . crassa 147 Aplysina 154, 1 56 alala, Ancorina 18 A. ianthelliformis 154, 156 albida, Spiraslrellu 34 A. laevis 156 albidus, Suberites wilsoni var 34 A. lendenfeldi 155, 156 Alcyonium putridosum 120 A. pedunculate 156 Aleclona 38, 39 A. purpurea: Carter 156 Algol 14 Aplysinopsis 134 aliena, Carteriospongia 126 arborea var. ramosa, Chalinopsilla 80 aliena, Phyllospongia 124, 125 arbuscula, Clathriodendron 55, 56 amaknakensis, Jones 94 arbuscula, Irciniu 134, 136 amaknakensis, Lissodendoryx 98 arenacea, Collosclerophora 69, 71 amaknakensis, Myxilla 94 arenaria, Slellelta 20 amaranthus, Phorbas 95 arenifera, Chondropsis 72, 179 ambigua, Reniera 90 arenifera, lichinoclathria 60, 61 amphiacantha, Chrotella 95 arenifera, Echinoclalhria favus var 60 Amphimedon complanata 105, 112 arenifibrosa, Lendenfeldiu'l 127 Amphimedon! crassiloba 105 arenifibrosa, Mcgalopastas 127 anatipes, Spongia 37 arenitccla, Stellctta 20 Anchinoe 95, 102 Arenochalinu (see Mycale) 84

A. coriaceus 103 A . mirabilis 84 A. coriaceus var. tenacior 102, 103 A. mirabilis: Whitelegge 84, 87 1

226 F. WII'DINMAYI.R

arenosa, Irctnia 134, 137 ,i. wellsi f>2 Arenosclera km, ios Axociella carterl 59

iMcj'.iii.iiis s<-> areolata, Eusi">ii|'i;i v;n . .. 122,123 \,favus areolata, Spirastrella 34 Axos spinipoculum 47 argentea, Dysidea 72 bacillifera var. robusta, Psammastra 17 argentea, Phorlospongla 72, is bacillifera var, robusta, Stcllctta 17 armigera, Cladoehalina 109,110 bacillifei var. robustum, Ecionema 17 armigera var, pergamentacea, Cladoehalina . . . bacterium, Spirophora 53

109, I io bacterium, Trachycladus 54 armigera, Tuba 105 Barbozia 37, W

asparagus, < allyspongla 106, i hi Barbozia primitive J7

asparagus, < lhallna [06 Batzella 67, 69

asparagus, < 'ladochalina j oc> it. friabilis 69

l asparagus, Spongia 106 it. frutex (> >

assabensis, Tedania 90 it. Inaequalis <»

assabensis, Tedania anhelans foi ma w it. intermedia 69 atlantlca, Ectyodot vx 96 behringi, Lissodendoryx 98 Aulena 58, 61, 144 biceps, Forcepia 93, 100 A. crassa 64 biceps, Suberites (>3 A. flabellum 144 bicoloi , Polymastia! 33 A. gigantea 60 bicoloi vai . crassa, Polymastia 52 A. gigantea var. intermedia 61 biformis, Gelliodes 113 A, gigantea var, macropora 61 biformis, Latrunculia (Latrunculia) 58 a. gigantea vai micropore 61 bilamellata, < 'altyspongla 106, 107, 1 1 A- laxa 60 bilamellata, < iavochalina 106, 108

A, laxa vai digitate 60 bilamellata, < ladochalina 106 A. laxa var. minima 60 bilamellata, Cribrochalina 106 A, nigra L44 bilamellata, Pachychalina 106 A. villosa 61, 144 bilamellata, Spongia 106, 108 Aulenella s(> bilamellatum, Echinodictyum 106 ss Aulospongus? cacticutls Biminia 105, 120 aurantlaca, A vlnella 48 blstellata, Diplastrella 38 aurantlaca, Bubaris -is bltextum, Psammoclema 75, 7>>, si aurlvlllii, Spirastrella 54,35 boleta, rhorecta 137 Aurora 'i boletus, Fhorecta 137, 139 A providentiae 23 bonneti, Spirastrella u aurora, Rhabdastrella 23 bowerbanki ( 'orticium 13 Aurorella 21 brunnea, Myxilla % australiensis, Ancorina is Bubaris 48 australiensis, Damiria '»<> b. aurantlaca (correctly Axinella) 48

M7, l australiensis, Darwinella [49, 151 buchanani, i issodendoryx 96, >7, 98 australis, Dactylochalina 112,113 bullata, Callyspongia 109 australis, Euplacella 106, 107, 108 Burtonispongia stelliderma (.s australis var. flabellum, Sigmatella 74 Cacochalina truncatella var. laxa 84 australis, Irc'mia 134, 136 c, truncatella var, mollissima w, i\ australis, Pachychalina 106 ( acospongia 134 australis var. tubaria, Sigmatella 74,75 C. camera 158 avara, Dysidea 144, 146 ( ', mollior I2t avara var. pallescens, Dysidea 144, 146 c, mollior: Ridley 121, 122 avara, Spongelia ill ( :, murrayll 140 Axiamon 4<) c. poculum 124 A. I0I111111 4') cacticutls, Aulospongus? 56 so Axinella 47. 'is, cactos, Spongelia 00

1 aurantlaca 48 cactus, (Dendrilla): de 1 aubenfels 00 1. convexa (correctly Pseudaxinella) 48 cactus, Dendrilla: Bergquisl 00

1 tinosia 48, 49 caliciformis, c 'arteriospotigta 00 1 1

SPONGES FROM BASS STRA1

cacticutis, < tathr'todendron 55, 56 carcinophila, Chondropsis 17 cacticutis, Dictyocylindrus ss carcinophila, Phoriospongta 72, m, fS, 16, 11 cacticutis, Raspailia ss carcinophila, Sigmatella /.' cactiformis, Acanthella 50 < armla 83 cactlformis, Rhaphoxya 50, 51, 52 carnosa, Spirastreila »4 cactos, Aplysilla is: carnosus, Geiliodes ..... 119 cactos, Dendrilla 152 carterl, Axocieila 59 cactos, Megalopastas 152 carteri, Coppatias 56 cactos, Spongelia 152, 153 carteri, Echinoclathrta 59, 60 1 cactus, {Dendrilla)'. de Laubenfels is. carteri, Forcepia 94 cactus, Dendrilla: Bergquisl 152,153 carteri, Oeodia (Ceodla) 26 calkiformis, Carteriospongia 124. 126, 127, 126, carteri, Stellettinopsis 56, 28 132 ( 'arteriospongia 12', 124, 128 caliciformis, Phyllospongia (Antheroplax) .124 c. aliens 126 caliculata, Hircinia 135, 136 c caliciformis ... 124, 126, 127, 128, 132 caliculata, Hircinia (Sarcotragus) 135 C. concentrica ,.,...., 131 caliculata, Ircinia 135, I 16 ( 127 ', eiegans: Burton 126, lis caliculata, Ircinia campana ( foliascens 123, 126 caliculata, Leiosella 130, 131, 1 )2 ( ', laciniata 123 catlosa, Dysidea 76 c. lacinulosa ...... 123 catlosum, Psammoclema iu C, lamellosa I2.< Callyspongia .... 103, 104, 105, 109, 111, US C, otahiticai Hyatl I2.t

c 110 '. asparagus 106, C. pennatula auctorum 123, 124 C. bilamellata 106,107,111 C, polyphylla 123

C. bullata 109 < ', radlata 123

< c. communis 105 '. sillcata 124, I2<», 127, 1 !6

c ( ', conica 105 ', vermicularis 127

< '. ( diffusa 108, 109 '. cf. vermicularis 127 ( '. eiegans hw ( ai tei ispongia 123

( ', exigua 114 ( !ai terospongia 123 c fallax forma debilis ios carvernosa, Damiria 94,95 c. irregularis 10s cavernosa, Damiriella 93 ('. pallida I os cavernosa, Dendoricella 93

( ( '. pergamentacea 106, i<> ), 110, ill cavernosa, 1 issodendoi vx ''5, ')/, '>h C. persculpta 106, no, ill, 112 Cavochalina bilamellata 106, 108

', ( pulvinatae 1 oh cellulosa, Spongia 59 C. ramosa 105, no, 112, 114, H7 Ceraochalina 101. 105, 111, is o') C. serpentina 105,109,114,115 < ceylonica 108, ( 1 '. spinosissima "(5 ( euplax

< '. ( ', stellata 105 extreme 1 C. subcylindrica 105 Clevis 112,113, 14

! 1 14, 1 17 microrhaphis I c ', toxlfera C.

( '. 1 in ncaia 1(15 C. multiformis var, digitate 108, 1 !

C. tuberculata W5 C. multiformis var. dura I I calyciformis, Phyllospongia 124 C. Itiformis var. eiegans I ! calyculata, Hircinia 135 C. multiformis var. lamella l l calyculata, H. (Sarcotragus) 133 C. Itiformis var. maeandra I ! calyculata, Leiosella I M C. multiformis var. manaarensis OK camera, Cacospongia 158 C. multifi s vai His I \ campana caliculata, ircinia 135 ' nuda I.'

' campana, Hircinia 135,136 C. nuda var. oxystrongylui I campana, H. (Sarcotragus) 133 ( nuda var. oxyui 12 canalis, Euryspongia 141 C. papillate 10 canalis, Stelospongia 141,142,143 < papillate var, intermedia no, 112, I 1

< vai I candelabrum, Corttcium 13,14 papillate , mai ropore 110,112, !

var. . l ! candelabrum, Dactylochalina in C. papillata micropora , 110,112, ., 1

228 F. wii;i)I',nmayi<;k

( .16 ( . pergamentacea 110, 1 15 '. ehuealla ('. 160 reteplax 113 c '. collectrix 37, 108 eolleelrix 36 37 ( . retiarmata C. ( 160 C. tenclla 113 ', plebeja 36 C. typica 104, 105, 1 15 c ', reniformis forma spurca ( 'eratodendron 121 C. reticulata 36. 37 36. 37 certa, Lissodendoryx 96 c '. spurca ceylonica, ( 'eraochalina 108, 109 Chrotdla 28 ( c 'haetodoryx 97 '. amphiacantha 95 ( 97 .16 '. richardl ehuealla, ( 'hondrosia c lhalina asparagus 106 ( lionanchora? 56 ( 15.1 '. crassiloba 1 1 cirsioides, Dendrilla ( 105 '. macropora 113 Cladochalina 104, ( c '. iiiida 112 '. armigera 109, 110

('. . 110 c. oculata var. novaezealandiae ... 1 14, 115 armigera var. pergamentacea . . 109, ( 108 '. 106 ( . palmata asparagus

( ( 106 '. pergamentacea 110 !, bilamellata ( '. 114, 115 ( . polyeholoma 112, 113, 114 communis ( 111 ( !, polyeholoma var. aii;'iila(a 115 '. dendroides 112,

('. polyeholoma var, moniliformis 115 ( ',. tlendyi 115 ( ( 10.1 '. polyeholoma var. oeulala . 112, 113, 115 '. diffusa ('. polyeholoma var. Irieholoma ... 112, I 13 c. elegans 109, 112, 113, 115 ( ( '. pulvinatae 108 '. elongata 115 ( ( 11.1 '. pulvinatus 108 '. euplax (halinella 104 ('. lamella 108

( ( '. maeropoia 112 '. maeropoia 115 ( 11.1 C. lenella 113 '. mammillata 112,

( halinissa communis 155, 116 ( ',. maims 108 ( ( 11.1, '. 111 '. communis var. digitata , 12, 115 mollis ( ('. '. communis var. flabellum 115 inula 104, 112

(.'. ( !. elegans 115 nncla var. ahi uplispieula 112 ( ( !. elongata 115 '. paueispiua 110 ( C !, maeropoia I 13, 115 '. pergamentacea IK)

( (.'. '. ramosa 109, 115 tenuifibris 115

( i \1 ('. '. rigida 115 tenuirhaphis 1 12, 11.1 ( '. sei pens 115 C. typica 110 ( '. leimilibi is 115 Clathria 56, 57. 59 ( ( 'halinopora 104 '. costifera 58

( eoimlala I 13 '. tenuis 62 ( lamella 108, 109, 112, 111 '. transiens 57 ( paueispiua 111), 112, 1 13, 114 '. wilsoni 57 58

typica . . . 104, 110, 113 ( 'lathriodendron 54. 55

( typica var. megarhaphis I 10 '. arbuscula 55 56 ( lypiea var. leimispina . no '. cacticutis 55 56 ( ( halinopsilla 75, 104, 105 '. rubrum 56 ( . 80 '. arborea var. ramosa , Clathriopsamma 56 ( '. dichotoma 105 (lalhrissa elegans 90 (halinopsis 104 clavatus, Trachycladus digitatus var 54

I . 1.17 choanoides, lalispongia . clavigera, Ectyodoryx 96 choanoides, Thorecta 30, 1.17 elifloni, Irlliva .10 1.17 choanoides, rhorectandra . ( 7/o/ifl acustella 19

Chondropsis . 71 ( 'liana levispira .19 ( ( !, areiiileia . 72 collectrix, 'hondrosia .17, 160

( . 72 eolleelrix, ( 'hondrosia .16 '. carcinophila , 37

'. i . 71 eolleelrix, Reniera .16 ( A irk , 37

( 'hoinirosiu .16 C olloelalhria 71 1

SPONGES t-'ROM BASS STRAIT 229

C. ramosa 71 C. pyriformis 128 Collosclerophora 69 costifera, Clathria 58 C. arenacea 69, 7 coxi, Chalinula 00 colonensis, Forcepia 93 Cruniella disigma 95 colonensis, Forcepia 93 crassa, Aiolochroia 147 columnaris, Spongia rimosa var 141 crassa, Aulena 64 commixta, Tedania 92 crassa, Echinoclathria 63 com in tin is, Callyspongia 105 crassa, Holopsamma 63, 64, 65 communis, Chalinissa 115, 1 16 crassa, Polymastia 32, 34 communis, Cladochalina 1 14, 115 crassa, Pseudoceralina 147 communis var. digilala, Chalinissa 112, 113, 115 crassanchorata, Forcepia 94 communis, Euplacella 109, 115 crassiloba, Amphimedon'! 105, 112 communis var. flabcllum, Chalinissa 115 crassiloba, Chalina Ill communis, Pachychalina 115 crassiloba, Spongia Ill, 112 communis, Tubulodigitus 115 crassum, Psammopemma 63, 64 compacta, Euspongia 131, 132 crassifibra, Siphonochalina 108, 109 compacta, Leiosella 131 Crella schmidti 94, 95, 96 compacta, Spongia 131 Cribrella hamigera 98 compactella, Spongia 131 ( ribrochulina 107, 1 I I complanata, Amphimedon 105, 112 C. bilamellatu (correctly Callyspongia) . . 106 concent rica, Carteriospongia 131 C. punctata 108 concentricum, Coscinoderma 131, 132 cribrocrusta, Stelospongus 139 cunctatrix, Spiras- confoederaia, Phoriospongia 75 irella 34 conica, Callyspongia 105 cunctatrix var. porcata, Spirastrel/a 35 conigera, Polymastia 33 Cydonium 23 conulata, Chalinopora 113 cylindrica, Spirasirella 34 conulata, Dactylochalina 113 Dactylia 75, 104, 105 conulosa, l.ainmculia (Latruncutia) 40, 43, 44, 46, Dactylochalina australis 112, 113 47 D. candelabrum 113 convexa, Axinella 48 D. conulata 113 convexa, Pseudaxinella 48 I), mollis 1 13 Coppatias carteri 26 D. monilata 113 cordata, Rhubdustrellu 21, 23 coriacca, Jaspis 26 Damiria auctorum 96 coriacea, Phyllospongia 124 D. australiensis 96 coriacea, Stelletlinopsis 28 1). cavernosa 94, 95 coriaceus, Anchinoc (correctly Phorbas) . . 103 Damiriana 94, 95, 96 coriaceus var. tenacior, Anchinoc .... 102, 103 D. hawaiiana 94, 96 corlicata, Ancorina 18, 19 D. sc/iimdli 96 corticate, Latruncutia {Negombata) .... 38, 40 Damiriella 94, 95 corticata, Sigmatclla (4 varieties, see Sigmatella eor- I ). cavernosa 95 ticata) 74 Darwinella 147, 151, 1 53 corticatus, Thorectandra 139 D. australiensis 147, 149, 151 Corticium 13, 14 D. duplex 148, 149

I), C. aeanthastrum 13, 14 gardineri 149, 1 53 C. bo werbanki 13 I), intermedia 147, 149 C. candelabrum 13, 14 I), mue/leri 147 C. reduction 13 D. nuielleri: de l.aubenfels 148 ( '. simplex 13 D. rosacea 148, 149 C. topsenti 13 D. simplex 147, 148, 149, 151

Coscinoderma 1 28, 1 30 D. viscosa 148, 149

C. concentricum (correctly Leiosella) 131, 132 Darwinia 1 47

C. lanugo 1 30 Dasychalina 104, 105 C. lanuginosum 128 D. fragilis 104, 105

C. musicalis 1 30 I), validissima |()5

C. pesleonis 128, 1 30, 1 39 decepta, Mvxil/a'l 94 C. pyrit'orme 128 decipiens, Psammascus'l 67, 75 230 F. WIEDENMAYER

Tedania 90 decipiens, Pseudaxinella 47, 48 digitata var. fibrosa, laxa var 60 decumbens, Spirastrella 34 digitata, Halme Tedania 90 Dendoricella 94, 95 digitata forma inermis, 152 D. abyssi 95 digitata, Luffaria

. 113 D. cavernosa 95 digitata, Pachychalina ramulosa var. 112, Tedania 90 D. obesichela 95 digitata forma polytyla, 92 D. rhopalum 94, 95 digitata, Reniera 89, Tedania 90 D. sc/unidti 95, 96 digitata var. rubicunda, 90 Dendrilki 152, 153 digitata var. rubra, Tedania 34 D. acanlha 153 digitata, Spirastrella digitata, Spirophora 53 D.> aerophoba 1 52, 153 53 D. antarctica 153 digitata, Spirophorella 92 D. cactos 152 digitata, Tedania 89, digitata var. verrucosa, Tedania 90 (D.) cactus: de Laubenfels (composite) .. 152 varieties, see D. cactus: Bergquist (recte Diclvodcndrilla) digitatus, Trachycladus (with 3 152, 153 Trachycladus digitatus) 53 38 D. cirsioides 153 Diplastrella bistellata 30 D. ianl/ielliformis (recte Aplysina) 154 diploderma, Donatia

, 32 D. lacunosa 151, 152 diploderma, Tethya 31 23 (D.) membranosa auctorum (composite) 153 discolor, Stelletta D. rosea 152, 153 disigma, Craniella 95 124 D. rosea var. digitata 152, 153 distans, Phyllospongia .'

. . 121 D. rosea var. typica 152, 153 Ditela Dendrocia 56 Ditriaenella 39 dendroides, Cladochalina 112, 113 diversichela, Lissodendoryx 96 dendroides, Euchalinopsis 113 Donatia diploderma 30 Dendrospongia 156 D. ingalli 30 dendyi, Cladochalina 115 D. ingalli sensu Dendy 31 dendyi, Hamigera 100 D. japonica var. globosa 30 dendyi, Lendenfeldial 127 D. japonica var. nucleata 30 dendyi, Lissodendoryx 96, 100, 102 D. parvistella 30 dendyi, Phakellia 51 D. seychellensis 31 densa, Pachychalina 113 Dosilia plumosa 38 densum, Psammoclema .... 65, 67, 73, 76, 77 Dotona pulehe!la 39 densum, Psammopemma 76 Druinella 156, 158, 160 densum var. subfibrosa, Psammopemma ... 76 D. purpurea 158, 159, 160 Desmacidon 66, 67 D. rotunda 158, 159, 160 Desmacidon abyssi 95 duplex, Darwinella 148 D. plicatum 68 dura, Ceraochalina multiformis var 113 D. stelliderma 68 dura, Euspongia irregularis var 121 Diacarnus 40, 47 dura, Placochalina pedunculata var 107 Diastra 21 durissima, Pseudoceratina 156, 158 dicholoma, Chalinopsilla 105 Dysidea 75, 134, 144 Dictyociona 56 D. avara 144, 146 Dictyocylindrus cacticulis 55 D. avara var. pallescens 144, 146 Diclyodendril/a 152, 153, 154 D. pallescens 144 D. retiaria 152 D. villosa 61 Didiscus 38, 39, 40 Dysidea auctorum 75 D. flavus 38 D. argentea 72 difficilis, Toxochalina 113 D. callosa 76 diffusa, Callyspongia 108, 109 D. favosa 73, 75 diffusa, Cladochalina 108 D. kirki 73 digitata, Aulena laxa var 60 D. nodosa 79 digitata, Ceraochalina multiformis var. 108, 113 Dysideopsis sulphurea 146 digitata, Chalinissa communis var. 112, 113, 115 Echinoclathria 57, 58, 59, 144 digitata, Dendrilla rosea var 152, 153 E. arenifera 60, 61 SPONGES FROM BASS STRAIT 231

E. carteri 59, 60 E. elongata 1 15 E. crassa 63 E. frondosa 106, 107, 108 £. egena 64, 66 E. mollissima 106, 108 E. favus 59, 61 euplax, Ceraochalina 113 E. favus var. arenifera 60 euplax, Cladochalina 113 E. glabra 60 Euricinia 1 34 E. globosa 63, 64, 65, 75 Euryspongia canalis 141 E. laminaefavosa 60, 61, 64, 75 Euspongia 121 E. leporina 56, 61 E. compacta 131, 132 131 E. macropora: Whitelegge 59 E. foliacea: Ridley 131 E. tenuis 59, 62 E. infundibuliformis E. tubulosa 66 E. irregularis 121 Echinodictyum 97, 98 E. irregularis var. areolata 122, 123 121 E. bilamellatum 106 E. irregularis var. dura 122 E. mesenterinum 106 E. irregularis var. frondosa 121, Ecionemia 14 E. irregularis var. hispida 122 .. 121, 122, 123 E. bacilliferum var. robustum 17 E. irregularis var. jacksonia 122 Ectoforcepia 92, 93 E. irregularis var. lutea 121, mollior 122 E. (rilabis 93 E. irregularis var. Ectyodoryx 95, 97, 98, 100 E. irregularis var. silicata .... 121, 122, 123 suriganensis 122 E. atlantica 96 E. irregularis var. 121, 122 E. clavigera 96 E. irregularis var. tenuis 121, 122 E. maculata 96, U)0 E. irregularis var. villosa 132 E. maculatus 100 E. levis 130, 131 E. paupertas 95 E. mathewsi Thorecta 139 E. ramilobosa 96 exemplum var. secunda, 139 Ectyomyxilla 97, 98 exemplum var. tertia, Thorecta 114 egena, Echinoclathria 64, 66 exigua, Callyspongia

1 Euchalina 1 14 elegans, Callyspongia 09 exigua, Ceraochalina 113 elegans, Carteriospongia: Burton 126, 127 extrema, Callyspongia 105 elegans, Ceraochalina multiformis var. ... 113 fallax forma debilis, Spongelia 143 elegans, Chalinissa 115 farlovi, 115 Thorecta 139, 142, 143 elegans, Cladochalina 109, 112, 113, farlovi, 90 Fasciospongia 134, 141 elegans, Clathrissa 132 flabellum 141 elegansl, Leiosella 130, 131, F. 3 F. rimosa 139, 141, 142 elegans, Pachychalina 112, 1 • 126 fastigatus, Trachycladus 54 elegans, Phyllospongia 87 favosa, Dysidea 73, 75 elegans, Spongelia: Lendenfeld Axociella 59 elongata, Chalinissa " 5 favus, favus, Echinoclathria 59, 61 elongata, Cladochalina 15 favus var. arenifera, Echinoclathria 60 elongata, Euplacella 115 felina, Rhaphoxya 52, 53 elongata, Pachychalina 84 Ircinia 1 34 Esperella felix, 86, 87 fibrosa, Rhabdastrella 23 E. spongiosa 84, Sigmosceptrella 39, 40, 47 . . . 84 fibrosa, Esperia 83, 104 Tedania digitata var 90 Euchalina fibrosa, 114 fictitioides, Anchinoe 102 E. exigua 113 ficlitioides, Phorbas 102 E. macropora 113 filiformis, Gelliodes 113 E. paucispina 13 finitima, Acervochalina: Ridley 108 E. phillippinensis (phillippensis) ' 104 firma, Lissodendoryx 97 E. typica ' [J*lii fissuratum, Psammoclema 77, 78, 79, 82 Euchalinopsis dendroides Fistularia 154 E. minima ]] fistulata, Mycale 84, 87 Euplacella 104, 105,111, 15 fistulata var. macrochela, Mycale 84 Eaustralis ^ ' 98 1UV ' 1J flabellata, Lissodendoryx E. communis ' 232 WIliDENMAYHR

flabelliformis var. lata, Stelospongus 137 G. flagelliformis 113 flabelliformis, Stelospongia 142 G. setosa'l (correctly selosusl) 87 flabelliformis, Stelospongus 137, 138, 139, 142, Geodia (Geodia) 23 143, 144 G. (G.) carteri 26 flabelliformis, Taonura 143 G. (G.) g/oboste/lifera 26 flabellum, Aulena 144 G. (G.) nigra 26 flabellum, Chalinissa communis var 115 6'. (G. ) punctata 24, 26 flabellum, Fasciospongia 141 geodides, Aneorina 14, 18, 19 flabellum, Sigmatella auslralis var 74 geodides, Psammastra 14 flabellum, Sigmatella eortieala var 74 geodides, Stelletta 14

flabellum, Stelodoryx 96 gigantea, Aulena (3 varieties, see Aulena gigantea) flabellum, Stelospongia .... 141, 142, 143, 144 , 60

flagelliformis, Gelliodes 113 gigantea, Halme (3 varieties, see Halme gigantea) flammula, Mycale (Arenochalina) 84, 87 60 38 flavus, Didiscus gigantea, Ireiniu 1 36 foliaeea, Euspongia: Ridley 131 gigas, Psammastra 17 foliaeea, Leiosella 131 glabra, Echinoclathria 60 foliaeea, Lendenfeldia 131 glaeialis, Aplysilla 146 foliaeea, Spongia 104, 131 globosa, Echinoclathria 63, 64, 65, 75 foliascens, Carteriospongia 123, 126 g/obosa, Halme 63, 64 foliascens, Spongia 123 globosa, Donatia japonica var 30 folium, Axiamon 49 globosiellata, Rhabdastrella 23 Forcepia 92, 93, 98 globostellifera, Geodia (Geodia) 26 /•'. biceps 93, 100 glomerosus, Thorecta 139, 140 /•'. carter! 94 goniodes, Psammoclema 76, 78, 79, 81 /•'. colonensis 93 gracilis, Trachycladus digitatus var 54 F. colonensis 93 granulosa, Polymastia 34, 37 /•'. crassanchorata 94 Grapelia 83, 84 F. michaelseni 93 Grayax 98 /•'. Irilabis 93 guettardi, Phoriospongia 67 fragilis, Dasychalina 104, 105 Haastia 144 fragilis, Gelliodes 119 Halichondria anhelans 87 fragilis, Lissodendoryx 96, 97 H . frequens 94 freija, Thorccta 155 H. isodietyalis 98, 99, 100 frequens, Halichondria 94 H. mammillata 74 frequens, Paramyxilla 94, 95 H . stellidertna 68 friabilis, Batzella 69 Halisarca reticulata 36, 37 friabilis, Stclospongos 141 Halispongia: Bowerbank 137 friabilis, Strongytacidon 69 H. e/ioanoides 137 frondosa, Euplacella 106, 107. 108 hallmanni, Latrunculia (Latruneulia) 38, 41, 46, 47

irregularis var. . 121, 122 frondosa, Euspongia Halme 58, 61 frutex, Batzella 69 H. gigantea 60 frutex, Strongytacidon 68 H. gigantea var. intermedia 60 65 fuliginosum, Psammoclema H. gigantea var. macropora 60 34 fusca, Polymastia 33, H. gigntea var. micropora 60 gardineri, Darwinella 149, 153 H . globosa 63 , 64 137 Geelongia H. irregularis 61 137 G. vasiformis H. irregularis var. lamellosa 61 117, 119 Gelliodes H. irregularis var. micropora 61 119 G. carnosus H. laxa 60 119 G. fragilis H. laxa var. digitata 60 1 119 G. incrustans 26, 18, H. laxa var. minima 60 G. pumilus 119 H. micropora 61 , 63 119 G. tenuirhabdus H. nidusvesparum 60 Gelliodes biformis 113 H . villosa 61 113 G. filiformis halmiformis, Ircinia 137 SPONGES FROM BASS STRAIT 233

Halmopsis 144 infundibulis, Spongia 131 Hamigera 94, 97, 98 ingalli, Donatia 30 H. hamigera 97, 98 ingalli senu Dendy, Donatia 31 H. rubens 97 ingalli, Tethya 29, 31 , 32 Hamigera auctorum 94, 97 insolens, Myxilla 95 H. dendyi 100 intermedia, Aulena gigantea var 61 H. ternatensis 97, 99 intermedia, Batzella 69 hamigera, Cribrella 97, 98 intermedia, Ceraochalina papillata var. 112, 1 13 hamigera, Hamigera 97, 98 intermedia, Darwinella 147, 149 hawaiiana, Damiriana 94, 96 intermedia, Halme gigantea var 61 Hemigellius 105 intermedia, Rhabdastrella 22, 23 Hemitedania 87 intermedia, Siphonochalina 112 Hippospongia 121 intermedia, Strongylacidon 68, 69 H. nigra 144 Iophon 97 Hircinia caliculata 135, 136 lotrocholu 153 H. calyculata 135 Ircinia 134, 136

H. campana 135, 136 /. arbuscula 1 34, 136

H. (Sarcotragus) caliculata 135 /. arenosa 134, 137 H. (S.) calyculata 135 /. austra/is 134, 136

H. (S.) campana 135 /. caliculata 135, 136

hirciniopsis, Acanthella 51 /. campana caliculata 135

hirsuta, Polymastia 33 /. felix 134

hispida, Euspongia irregularis var 122 /. gigantea 136

hispida, Spongia 76, 121 /. halmiformis 137

hixoni, Raphyrus 34 /. irregularis 136

holdsworthi, Phyllospongia 124 /. irregularis 1 34, 136

Holopsamma 58, 71, 75 /. jacksoniana 134, 136

H. crassa 63, 64, 65 /. lendenfeldi 134, 136

H . laevis 77 /. muscarum 134

Homaxinella"! 48 /. rubra 136

Homoeodictya 1 04 /. rugosa 137

horrens, Spongelia 153 /. spinosula 1 34 horridus, Thorecta 87 /. strobilina 134 Hyattella 134 irregularis, Caltyspongia 105 Hymantho 56 irregularis, Euspongia (11 varieties, see Euspongia Hymedesmial paupertas 95 irregularis) 121, 122 Hymeniacidon 67, 69 irregularis, Halme 61 H. paupertas 95 irregularis, Ircinia 136 Hyrtios 134 irregularis, Ircinia 134, 136 ianthelliformis, Aplysina 154, 156 irregularis var. lamellosa, Halme 61 ianthelliformis, Dendrilla 154 irregularis var. micropora, Halme 61 ignis, Tedania 89, 92 irregularis, Spongia 122 ignis, Tedania 90 irregularis, Spongia zimocca 122 ignis pacifica, Tedania 91 isodictyalis, Halichondria 98, 99, 100 imperfecta, Mycale 84 isodictyalis, Lissodendoryx 94, 95, 96, 97, 98, 100 inaequalis, Batzella 69 isodictyalis, Myxilla 98, 99, 180 inaequalis, Strongylacidon 69 isodictyalis jacksoniana, Lissodendoryx ... 100 inaequalis, Stylotella 67, 69, 71 lsops 23 incerta, Spongelia 128 ivanovi, Lissodendoryx 98 inconstans, Spirastrella 34 jacksonia, Euspongia irregularis var. 121, 122, 123 incrustans, Gelliodes 26, 118, 119 jacksoniana, Ircinia 134, 136 indistincta, Lissodendoryx 95, 97, 98 jacksoniana, "Lissodendoryx" 97, 100 inermis, Tedania digitata forma 90 japonica var. globosa, Donatia 30 inflata, Tethya 30 japonica var. nucleata, Donatia 30 infundibuliformis, Euspongia 131 japonica, Tethya 30 infundibuliformis, Spongia 131 Jaspis 26 234 F. WIEDENMAYER

/. coriacea 26 Leiosella 130 J. stellifera 26, 28 L. caliculata 130, 131, 132 J. tuberculata 26 L. calyculata 131 Jones 94, 95 L. compacta 131, 181 juniperina, Thalysias 58 L. elegansl 130, 131, 132 kirki, Chondropsis 73 L. foliacea 131

kirki, Dysidea 73 L. laevis 1 32 kirki, Phoriospongia 72, 73, 75, 77 L. levis 132 kirki, Psammascus 73 lendenfeldi, Aplysina 155, 156 kirki, Spongelia 73 lendenfeldi, Ircinia 134, 136 Korotnewia 158 Lendenfeldial 123, 124, 127 kyma, Lissodendoryx 96, 97 L. areni fibrosa 127 laciniata, Carteriospongia 123 L. dendyi 127 lacinulosa, Carteriospongia 123 L. foliacea (recte Leiosella) 131 lacunosa, Dendrilla 151, 152 L. lamellaris 127

lacunosa, Spirastrella 34 /.. plicata 127 laeviaster, Thoosa 38 L. polyphemus 127

laevis, Aplysina 1 56 L. ridleyi 127 laevis, Holpsamma 77 L. torresia 127 laevis, Leiosella 132 leporina, Echinoelathria 56, 61 laevis, Tethya 30 leporina, Spongia 59, 62 laevispirulifer, Trachycladus .... 53, 54, 61, 97 Leptoclathria 56 lamella, Ceraochalina multiformis var. ... 113 leptoderma, Tedania 100 lamella, Chalinopora 108, 109, 112, 113 levis, Ceraochalina 112, 113, 114 lamella, Cladochalina 108 levis, Euspongia 130, 132

lamella, Pachychalina ramulosa var. . 112, 113 levis, Leiosella 132 lamella, Phoriospongia 67, 75 levispira, Cliona 39 lamella, Psammascus 67 lindgreni, Lissodendorvx 96 lamella, Reniochalina 48, 49, 50 Lissodendoryx 94, 95, 96, 97, 98, 100 lamellaris, Lendenfeldial 127 L. amaknakensis 98 lamellosa, Carteriospongia 123 L. behringi 98 lamellosa, Halme irregularis var 61 L. buchanani 96, 97, 98 lamellosa, Spongia 123, 124 L. cavernosa 95, 97, 98

laminaefavosa, Echinoelathria . . 60, 61, 64, 75 L. certa 96 lanugo, Coscinoderma 130 L. dendyi 96, 100, 102 lanuginosum, Coscinoderma 128 L. diversichela 96 lata, Stelospongus flabelliformis var 137 L. firma 97 Latrunculia 37, 38, 39, 40 L. flabellata 98 Latrunculia (Latrunculia) 40 L. fragilis 96, 97 L. (L.) biannulata 47 L. indistincta 95, 97, 98 L. (L.) biformis 38 L. isodictyalis 95, 96, 97, 98, 100 L. (L.) conulosa 40, 43, 44, 46, 47 L. isodictyalis jacksoniana 100 L. (L.) hallmanni 38, 41, 46, 47 L. ivanovi 98 L. (L.) multirotalis 38 "L. "jacksoniana 97, 100 L. (L.) spinipoeulum 47 L. kyma 96, 97 Latrunculia (Negombata) 40 L. lindgreni 96 L. (N.) acanthosanidastera 38 L. lobosa 96 L. (N.) corticata 38, 40 L. lundbecki 96 L. (N.) purpurea 40 L. mollis 96 L. (N.) tarentina 38, 40 "L. " oxeota 98 latus, Stelospongus 137, 138 L. papillata 100 latus, Thorecta 130, 139 L. papiltosa 98

laxa, Aulena (2 varieties, see Aulena laxa) . 60 L. sigmata 94, 96 laxa, Cacochalina truncatella var 84 L. similis 98, 100

laxa, Halme (2 varieties, see Halme laxa) . . 60 L. sinensis 99, 100 laxissima, Mycale (Arenochalina) 87 L. sophia 96 SPONGES FROM BASS STRAIT 235

L. stipitata 96, 98 M. roosevelti 94, 96 L. strongylata 94 M. tortugasense 94, 95 L. stvloderma 96 mesenterinum, Echinodictyum 106 L. ternatensis 97, 99, 100 michaelseni, Eorcepia 93 Litaspongia 58 Microciona 56, 59 lobosa, Lissodendoryx 96 micropora, Aulena gigantea var 61 loveni, Podospongia 37, 40 micropora, Ceraochalina papillata var. 112, 113

Luffaria 154 micropora, Halme 61 , 63 L. digitata 152 micropora, Halme gigantea var 61 Luffaria: PolejaelT 154 micropora, Halme irregularis var 61 Luffariella 154 microrhaphis, Ceraochalina 113 lunae, Pachychalina 113 microxea, Tetilla 29 iundbecki, Lissodendoryx 96 Microxina 105 lutea, Euspongia irregularis var 121, 122 minima, Aulena laxa var 60 lutea, Stellettinopsis 26, 28 minima, Euchalinopsis 113 luteus, Coppatias 26 minima, Halme laxa var 60 macrochela, Mycale fistulata var 84 mirabilis, Arenochalina 84 macropora, Aulena gigantea var 61 mirabilis, Arenochalina: Whitelegge .... 84, 87 macropora, Ceraochalina papillata var. 112, 113 mirabilis, Mycale (Arenochalina) .. 84, 87, 151 macropora, Chalina 113 mollior, Cucospongia 123 macropora, Chalinella 112 mollior, Cacospongia: Ridley 121, 122 macropora, Chalinissa 113, 115 mollior, Euspongia irregularis var 122 macropora, Cladochalina 115 mollis, Ceraochalina multiformis var 113 macropora, Echinociathria: Whkelegge .... 59 mollis, Cladochalina 113 macropora, Euchalina 113 mollis, Daetylochalina 113 macropora, Halme gigantea var 61 mollis, Lissodendoryx 96 macropora, Plectispa 59 mollis, Placochalina pedunculata var 107 38, 39 macrospina, Pachychalina 1 12, 113 mollis, Thoosa iruncatella var. 69, 71 maculat a, Ectyodoryx 96, 100 mollissima, Cacochalina 107 maculaturn, Merriamium 100 mollissima, Euplacella 106, maculatus, Ectyodoryx 100 monilata, Daetylochalina 113

madagascarensis, Phyllospongia 124 moniliformis, Chalina polyehotoma var. . . 115 maeandra, Ceraochalina multiformis var. 113 monodi, Tetilla 29 mammillaris, Sigmatella corticata var 74 muggiana, Reniera 89, 90, 92 mammillata, Cladochalina 112, 113 muelleri, Darwinella 147 mammillata, Halichondria 74 muelleri, Darwinella: de l.aubenfels 148 Cer- manaarensis, Ceraochalina multiformis var. 108 multiformis, Ceraochalina (8 varieties, see manaarensis, Pachychalina multiformis var. 108 aochalina multiformis) 113 manus, Cladochalina 108 multirotalis, Latrunculia (Latrunculia) 38 manus, Pachychalina 108, 109 multistella, Tethya 30 18 massa, Spirastrella 34 murrayi, Ancorina 16, 140 massalis, Polymastia'? 33 murrayil, Cacospongia 140 mathewsi, Euspongia 131 murrayP., Thorectu! 23 muscarum, Ircinia 1 34 Mauricea ' 1 30 Megalopastas arenifibrosa 127 musicalis, Coscinodcrma

83 , 84 M. cactos l 5 ^ Mycale fistulata 84, 87 M. retiaria I 52 'M. macrochela 84 megarhaphis, Chalinopora typica var 110 M. fistulata var. 84 mcmbranacea, Spongia '53 M. imperfecta 87 membranosa, Dendrilla (antiboreal records) 153 M. spongiosa 84, 87 membranosa, Dendrilla: Burton (1934a) .. 153 M. tylostrongylal 84, 84 membranosa, Dendrilla (Indo-Paeific records) 153 Mycale (Arenochalina) 87 membranosa, Spongia: Esper 153 M. (A.) flammula 84, 87 membranosa, Spongia: Pallas 153 M. (A.) laxissima Merriamium 94, 95, 98 M. (A.) mirabilis 84, 87, 151 00 M. (A.) pluriloba 87 M. maculatum ' 236 F. Wli;i)l'NMAYI-K

Mycalecarmia X4 P. communis 115 Myriastra 19, 20 P. densa 113 Myxilla 94, 96, 97, (>K P. elegans 112, 113 M. brunnea % P. elongata 115 A/.? decepta 94 P. lunae 113 Myxilla amaknakensis 94 P. macrospina 112, 113 M. anhelans 87 P, maims 108, 109 M. insolens ^^ P. multiformis var. manaarensis iox M. isodictyalis 98, 99 P. oculata 112 M. veneta 97 P, paucispina 112, 113 Negombata (see / atrunculia) 40 P. punctata 108 Negombo 40 P. punctata 107

I', 112 nidusvesparum, I [alme 60 ramosa nigra, Aulena 144 l'. ramosa 115, 116 nigra, Geodia {Geodla) 26 P. ramulosa var. digitata 112, 113 nigra, Hippospongia 144 P. ramulosa var. lamella 112, 113 nigrescens, Renieral 89 P. rigida 112, 113 nigrescens, Tedania 89 pacifica, Tedania ignis l)l

( ) 144 nodosa, l )ysidea 7 pallescens, Dysidea

l nodosa, Saicocoinca 75, 7 ) pallescens, Dysidea avara var 144, 146

( nodosa. Spongelia 7 ) pallescens, Spongelia 144, 146 nodosum, I'sanimoilcina 76, 79, 83 pallescens, Spongelia: Scluil/c 144 novaezealandiae, Chalina oculata var. 114, II s pallida, Callyspongia 105 nncleala, Donalia japonic.! var 10 pallida, Khaphoxva 53 nuda var. abruptispicula, Cladochalina ... 112 palmata, Chalina 108

mula, ( leraochalina 112 palmata, Spongia 108 mula, Chalina 112 Pandaros 56, 58, 59

inula, ( lailochalina 104, I 12 panis, Papillina 35 34 inula var, oxystrongylus, Ceraochalina . ... 112 panis, Spirastrella

inula var. o\yns, ( 'eiaoclialiua 112 panis, Subcrilcs 90 obesichela, Dendoricella 95 panis, Tedania 90

( )ceanapia 120 papilla, Ophlitaspongia 59

( ). putridosa 120 papillata, Ceraochalina (with 3 varieties, sec Cer- oculata, Chalina polychotoma var. 112,113,115 aochalina papillata) 112,113

oculata var. novaezealandiae, Chalina 1 14, 115 papillata, Lissodendoryx 100 oculata, Pachychalina 112 Papillina panis 35 l, ( )nychomyxilla 7 papillose/, l issodendoryx 98

Ophlitaspongia 56, 57, 59 papulosa vai . porosa, Spirastrella 35 o. papilla 59 papillosa, Sigmatella corticata var 74, 76 s ( ). seriata '> papillosa, Spirastrella 35, 36

( )phliiaspon)'ia aucioi iiiu 58, 59, 66 papyracea, Phakeliia 62

( >. tenuis 62 papyracea, Phyllospongia 124

( ). tubulosa 66 papyracea, Spongia 104 Orina 105 Paramyxilla 94, <>5

otahitica, ( 'arteriospongia: l Ivan 123 p, frequens 94, 95 otahitica, Spongia 123 Paraspongia 104, 105 oxeata, Tedania 96 Paratedania 87

l oxeata, Tetilla 2 ) parviconulata, Acanthellina 51 oxeota, "l issodendoryx" 98 parvistella, Donatia 30 Owlalruncnlia 40 I'ainloscula 104 40 o. acanthosanidastera paucispina, Chalinopora . . . 1 10, 1 12, 113, 1 14 oxystrongylus, Ceraochalina nuda var 112 paucispina, cladochalina no oxyus, Ceraochalina nuda var 112 paucispina, Euchalina 113 Pachychalina affinis 113 paucispina Pachychalina 112, 113 P. australis 106 paupertas, Anchinoe 96 P, bilamellata 10<> paupertas, Ectyodoryx 95 SPONGES FROM BASS STRAIT 237

paupertas, Hymedesmia 95 P. vasiformis 131 paupertas, Hymeniacidon 95 P. vermicularis 127 paupertus, Phorbasl 96 Phyllospongia (Antheroplax) caliciformis .. 124 pedunculata, Aplysina 156 P. (A.) perforata 113 pedunculata, Placohalina (3 varieties, see Plaeoeha- Phyllospongia (Carteriospongia) silica/a ... 126 lina peduneulata) P. (C) vermicularis 107 1 27 pennatula aucloruin, Carteriospongia 123, 124 Piloehrola 19, 20 perforata, Antherochalina 62 Placochalina 104 perforata, Phyllospongia (Antheroplax) ... 113 P. peduneulata var. dura 107

pergamentacea, Callyspongia 106, 109, 110, 114 P. peduneulata var. mollis |()7 pergamentacea, Ceraoehalina 110, 115 P. peduneulata var. poeuluin 107 pergamentacea, Chalina 110 Platyehalina 104 pergamentacea, Cladochalina 110 plebeja, Chondrosiu: Kirkpatriek 160 pergamentacea, Cladochalina armigera var. 109 Pleelispa maeropora 59 110 plicata, hendenfeldial 127 persculpta, Callyspongia ... 106, 110, 111, 112 plieatuni, Desmacidon 68 pesleonis, Coscinoderma 128, 130, 139 Phtmohalichondria 9g pesleonis, Spongia P. 1 28 plumosa 97 Phakellia 48 plumosa, Dosi/ia 38 P. dendyi 51 plumosa, Plumohalichondiia 97 P. papyraeea Carter 62 pluriloba, Mycale (Arenochalina) 87 P. papyraeea Ridley poeuluin, [ Dendy 62 Cacospongia 124 P. tumida poeuluin, 48 Placochalina peduneulata var. .. 107 Phakettia 48 Podospongia 40 P. wellsi 62 P. loveni 37, 40 phillipensis, Tethya 30 polyeholoma, Chalina (with 4 varieties, see Chalina phillippensis, Euehalina 113 polychotoma) 112, 113, 114, 115 phillippinensis, Euehalina 113 Polyfibrospongia 123 Phloeodictyon Polymastia 1 20 32, 33 37 P. putridosa P.! 1 20 bicolor 33 P. putridosum /'. 1 20 bicolor var. crassa 32 Phlyctaenopora 38 P. conigera 33 Phorbas 95, 102 P. crassa 32, 34 P. amaranthus 95 /' fitsca 33,' 34 P. fictitioides P. 1 02 granulosa 34, 37 P. ? paupertas /'. hirsuta 96 33 P. el", tenacior P. 1 02 ? rnassalis 3} Phoriospongia P. 64, 67, 71, 76, 105 polylylola 37 P. argentea 72, 75 polyphemus, Lendenfetdial 127 P. carcinophila 72, 73, 75, 76, 77 polyphylla, Carteriospongia 123 P. confoederata 75 polytyla, Tedaniu digitata froma 90 P. guettardi polytylota. 67 Polymastia 37 P. kirki 72, 73, 75, 77 poreata, Spirastre/la eunetatrix var 35 P. lamella 67, 75 porosa, Spirastrellu pupillosa var 35 P. reticulum 63, 64, 67 praecipua, Tetilla 28, 29

P. syringiana primitiva, .' 75 Barbozia 37 P. wilsoni 65, 76 primitiva, Rhuhdustrella 23 Phyllospongia progressa, 123, 124 Ancorina 18 P. aliena 124, 125 Pronax 9^ P. calyciformis 124 providentiae, Aurora 23 P. coriacea 124 Psammaplysilla 155 158

P. distans P. ' 1 24 purpurea'! 158 P. elegans 1 26 Psamniascus'l 67 75

P. holdsworthi 1 P. '! decipiens 24 67, 75 P. madagasearensis 124 Psammaseus auetorum 67' 71 P. papyraeea 124 P. kirki '//.... .' 73 1

238 F. WIEDENMAYER

P. lamella 67 pyri forme, Coscinoderma 128 Psammastra 14 pyriformis, Coscinoderma 128 P. bacilli fcra var. robusta 17 quadrilobata, Sigmosceptrella 39, 40, 47 P. gigas 17 Quizciona? 56 Psammochela 67, 71 radiata, Carteriospongiu 123

Psammocinia 134, 1 35 radiatum, Psammoclema 80 Psammoclema 64, 67, 71, 75, 77, 104 ramilobosa, Ectyodoryx 96

/'. bitextum 75, 79, 81 ramosa, Callyspongia . 105, 110, 112, 114, 117 P. callosum 76 ramosa, Chalinopsilla arborea var 80 /'. densum 65, 67, 73, 76, 77 ramosa, Chalinissa 109, 115 P. tissuratiim 77, 78, 79, 82 ramosa, Colloclathria 71 P. fuliginoswn 65 ramosa, Pachychalina 112 P. goniodes 76, 78, 79, 8 ramosa, Pachychalina 115, 116 P. nodosum 76, 79, 83 ramosa, Spongia 112, 113, 114 P. radiatum 80 ramosum, Psammoclema 76, 79, 80

P. ramosum 76, 79, 80 ramulosa var. digitata, Pachychalina . 112, 113

P. stipitalum 78, 81, 82 ramulosa var. lamella, Pachychalina . 112, 113 P. vansoesli 79, 80, 82 Raphioderma 83 Psarnmopcmma 36, 75 Raphiodesma 83 P. crassum 63, 64 Raphyrus hixoni 34 P. densum 76 Raspailia 54, 55 P. densum var. sub fibrosa 76 Raspailia cacticulis 55 Psammotoxa 67, 7 reductitm, Corlieium 13 Pseudanchinoe 56 Renieral nigrescensl 89 Pseuduxinella 47, 48, 49 Reniera ambigua 90 P. eonvexa 48 R. collectrix 36, 37 P. decipiens 47, 48 R. digitata 89, 92 P.? tumida 48 R. muggiana 89, 90, 92 Pseudoeerulina 156, 158 reniformis forma spurca, Chondrosia 36 P. erassa 147 Reniochalina 48, 49 P. durissima 156, 158 R. lamella 48, 49, 50 Pseudoespcria 84 R. seetitlis 49, 50

Ptiloeaulis 49 A', stalagmitis 48, 49, 50 pulchella, Dotona 39 repens, Ancorina 16 pulvinatae, Chalina 108 reteplax, Ceraoehalina 113 pulvinatae, Callyspongia 108 reteporosus, Trachycladus 54 pulvinatus, Chalina 108 retiaria, Dictyodendrilla 152 pumilus, Gel/iodes 119 retiaria, Megalopastas 152 punctata, Cribrochalina 108 retiarmata, Ceraoehalina 108 punctata, Geodia, (Geodia) 24, 26 reticulata, Chondrosia 36, 37 punctata, Pachychalina 108 reticulata, Halisarca 36, 37 punctata, Pachychalina 107 reticulata, Rhabdastrella 23 punctata, Spirastrellu 34 reticulata, Stelospongia 141 , 142 purpurea, Aplysina 156 reticulum, Phoriospongia 63, 64, 67

purpurea, Druinella 158, 159, 160 Rhabdastrella 21 , 26 purpurea, l.atrunculia (Negombata) 40 R. aurora 23

1 purpurea'., Psammaplysilla 158, 159 R. cordata 21 , 23 purpurea, Spirastrella 34 R. fibrosa 23 purpurea, Spirastrella: Vosmaer 35 R. globostellata 23 purpurea, Stellettinopsis 28 R. intermedia 22, 23 puslulosus, Trachycladus 54 R. primitiva 23 putridosa, Oceanapia 120 R. reticulata 23 putridosa, Phloeodietyon 120 R. rowi 23 putridosa, Rhizochalina 120 R. spinosa 23 ptttridosuin, Alcyonium 120 R. tenuis 23 pulridosum, Phloeodietyon 120 R. virgula 23 1

SPONGES FROM BASS STRAIT 239

Rhaphidophlus 71 schmidti, Waldoschmittia 95 Rhaphoxya 50 sectilis, Reniochalina 49, 50 R. cadiforinis semilunaris, 50, 5 1 , 52 Spirastrel/a 34 R- felina 52^ 53 seriata, Ophlitaspongia 59 R. pallida 53 serita, Spongia 59 R. typica 53 serpens, Chalinissa [15 Rhizochalina 1 20 serpentina, Callyspongia ... 105, 109, 114, 115 R. putridosa 1 20 serpentina, Spongia 114, 115, 116 rhopalum, Dendoricella 94, 95 serrata, Sigmatella corticata var 74 richardi, Chaetodoryx 97 setose! , Gelliodes (correctly setosusl) 87 ridleyi, Lendenfeldial 127 seychellensis, Donatia 31 rigida, Chalinissa 112, 115 seychellensis, Tethya 30, 31 , 32 rigida, Pachychalina 112, 113 Sidonops 24 rimosa, Fasciospongia 139, 141, 142 sigmata, Lissodendoryx 94, 96 rimosa, Spongia (2 varieties, see Spongia rimosa) Sigmatella 71 141 S. australis var. flabellum 74 rimosa, Stelospongia 141, 142 S. australis var. tubaria 75 rimosa, Thorecta 141 S. carcinophila 72 robusta, Ancorina 17, 18 S. corticata 74 robusta, Ancorina (Eeionemia) 17 S. corticata var. I'labellum 74 robusta, Psammastra bacillifera var 17 S. corticata var. mammillaris 74 robusta, Stelletta bacillifera var 17 s. corticata var. papillosa 74, 76 robusta, Tethya 30, 3 S. corticata var. serrata 74 robustum, Ecionema bacilli ferum var 17 Sigmosceptrella 37, 39, 40 roosevelti, Merriamium 94, 96 S. fibrosa 39, 40, 47 rosacea, Darwinella 148, 149 S. quadrilobata 39, 40, 47 rosea, Aplysilla 146, 147 silicala, Carteriospongia ... 124, 126, 127, 136 rosea, Aplysilla: Rockman 152 silicata, Euspongia irregularis var. 121, 122, 123

rosea, Dendrilla 1 52, 153 silicata, Phyllospongia (Carteriospongia) .. 126 rosea var. digitata, Dendrilla 152, 153 similis, Lissodendoryx 98, 100 rosea var. ( typica, Dendrilla 152, 153 simplex, 'or/icium 13 rosea, Verongia 146 simplex, Darwinella 147, 148, 149, 151 rotunda, Druinella 158, 159, 160 Simplicella |4(, row'/, Rhabdastrella 23 sinensis, Lissodendoryx 99, 100 rubens, Hamigera 97 Siphonella 104, 105 rubicunda, Tedania 90, 91 Siphonochalina auctorum 104, 105 rubicunda, Tedania digitata var 90 S. crassifibra 108, 109 rubra, Ircinia 136 S. intermedia 112 rubra, Tedania 90 Smenospongia 134 rubra, Tedania digitata var 90 solida, Spirastrella 34 rubrum, Clathriodendron 56 sophia, Lissodendoryx 96 rugolineata, Acanthellina 51 spiculifera, Spirastrella 34 rugosa, Ircinia 137 spinipoculum, Axos 47 Samus 39 spinipoculum, Latrunculia (l.atrunculiu) ... 47 Sanidastrella 14 spinispiru/ifera, Spirastrella 34, 39 sansibarensis, Stongylacidon 68 spinosa, Rhabdastrella .' 23 Sarcocornea 75, 144 spinosa, Spongelia 128 S. nodosa 75, 79 Spinosissima, Callyspongia 105 Sarcotragus 134, 135 Spinosella I04 j 105 scabrosus, Trachycladus 54 spinosula, Ircinia ] 34 scariola, Stylinos Spirastrella 66 34, 38, 39 Sceptrella 40 .V. albida 34 Sceptrintus 38, 39 S. areolata 34 schmidti, Crella 94, 95, 96 S. aurivillii 34 35 Schmidti, Damiriana 96 S. bonneti 34 schmidti, Dendoricella 95, 96 S. camosa 34 240 F. W1EDENMAYER

S. cunctatrix 34 S. infundibulis 131 S. cunctatrix var. porcata 35 S. lamellosa 123, 124 S. cylindrica 34 S. leporina 59, 62 S. decumbens 34 S. membranacea 153 S. digitata 34 S. membranosa: Esper 153 S. inconstant 34 S. membranosa: Pallas 153 S. lacunosa 34 S. otahitica 123 S. massa 34 S. palmata 108 S. panis 34 S. papyracea 104 S. papillosa 35 36 S. pesleonis 128 S. papillosa var. porosa 35 S. ramosa 112, 113, 114 S. punctata 34 S. rimosa 141 S. purpurea 34 S. rimosa var. columnaris 141 S. purpurea: Vosmaer 35 S. rimosa var. subclavata 141 S. semilunaris 34 S. seriala 59 S. solida 34 S. serpentina 114, 115, 116 5. spiculifera 34 spongiosa. Esperella 84, 86, 87 S. spinispirulifera 34 39 spongiosa, Mycale 84, 87 S. spirastrelloides 34 spurca, Chondrosia 36, 37 S. tentorioides 34 spurca, Chondrosia reniformis forma 36 S. transitoria 34 stalagmitis, Reniochalina 48, 49, 50 S. vagabunda 34 stalagmoides, Ancorina 18 S. wilsoni 34 stellata, Callyspongia 105 spirastrelloides, Spirastrella 34 Stelletta 19 Spirophora 53 S. arenaria 20 S. bacterium 53 S. arenitecta 20 S. digitata 53 S. bacillifera var. robusta 17 Spirophorella 53 S. discolor 23 S. digitata 53 S. vestigium 20 Spongelia 144 Stellettinopsis 14 S. avara 144 S. carteri 26, 28 S. S. cactos 1 52, 1 53 coriacea 28 S. elegans: Lendenfeld 87 S. lutea 26, 28 S. far/ovi 143 S. purpurea 28 S. horrens 153 S. stellifera 26 S. incerta 128 S. tuberculata 26, 28 S. kirki 73 stelliderma, Burtonispongia 68 S. nodosa 79 stelliderma, Desmacidon 68 S. pallescens 144, 146 stelliderma, Halichondria 68 S. pallescens: Schulze 144 stelliderma, Strongylacidon 68 S. spinosa 128 stellifera, Amorphina 26 Spongia 121 stellifera, Coppatias 26 5. hispida 76, 121 stellifera, Jaspis 26, 28 5. irregularis 122 stellifera, Stellettinopsis 26 S. zimocca 123 Stelodoryxl 98 S. zimocca irregularis 122 S. flabellum 96

Spongia anatipes . 87 Stelospongia 1 34, 141 S. asparagus 106 S. canalis 141,142,143 S. bilamellata 106, 108 S. flabelliformis 142

S. cellulosa . 59 S. flabellum 141, 142, 143, 144 S. compacta 131 S. reticulata 141, 142 S. compactella 131 S. rimosa 141, 142

S. crassiloba Ill, 1 12 Stelospongos 141 S. foliacea 104, 131 S. friabilis 141 S. foliascens 123 Stelospongus 141 S. infundibuliformis 131 S. cribrocrusta 139 11 1

SPONGES FROM BASS STRAIT 241

S. flabelliformis 137, 138, 139, 142, 143, 144 T. ignis 89, 92 S. flabelliformis var. lata 137 T. ignis 90 S. latus 137, 138 T. ignis pacifica 91 S. tuberculatiis 140 T. leptoderma (correctly Lissodendoryx isodic- stipitata, Acanthella 51 tyalis) 100 stipitata, Lissodendoryx 96, 98 T. nigrescens 89 stipitatum, Psammoclema 78, 8182 T. oxeata 96 slrobilina, Ircinia 134 T. panis 90 Strongylacidon 67, 69, 7 T. rubicunda 90, 91 S. friabilis 69 T. rubra 90 S. frutex 68 T. tantula 87, 97 S. inaequalis 69 Tedaniopsis 87 S. intermedia 68, 69 Teichaxinella 48 S. sansibarensis 68 tenacior, Anchinoe 102 S. stelliderma 68 tenacior, Anchinoe coriaceus var 102, 103 strongylata, Lissodendoryx 94 tenacior, Phorbas cf 102 strongylatus, Trachycladus digitatus var. ... 54 tenella, Ceraochalina 113 stylifer, Trachycladus 54 tenella, Chalinella 113 Stylinos 66 tentorioides, Spirastrella 34 S. scariola 66 tenuifibris, Chalinissa 115 styloderma, Lissodendoryx 96 tenuifibris, Cladochalina 115 Stylotella 67, 68, 69 tenuirhabdus, Gelliodes 119 S. agminuta 71 tenuirhaphis, Cladochalina 112, 113 S. inaequalis 67, 69, 7 tenuis, Clathria 62 Stylotella auctorum 69 tenuis, Echinoclathria 59, 62 subclavata, Spongia rimosa var 141 tenuis, Euspongia irregularis var 121, 122 subcylindrica, Callyspongia 105 tenuis, Ophlitaspongia 62 Suberites 33 tenuis, Rhabdastrella 23 Suberites biceps 93 tenuispina, Antherochalina 62 S. panis 90 tenuispina, Chalinopora typica var 110 S. wilsoni 38 ternatensis, Hamigera 97, 99 S. wilsoni var. albidus 34 ternatensis, Lissodendoryx 97, 99, 100 Suberotelites 102 Tethya 29 subfibrosa, Psammopemma densum var. ... 76 T. cliftoni 30 suiiia. Ancorina 18, 19 T. diplodcrma 31, 32 sulfurea, Aplysilla 146, 147 T. inflata 30 sulphurea, Aplysilla 146 T. ingalli 29, 31, 32 sulphurea, Dysideopsis 146 T. japonica 30 suriganensis, Euspongia irregularis var. ... 122 T. laevis , 30 syringiana, Phoriospongia 75 T. mullistella 30 tantula, Tedania 87, 97 T. phillipensis 30 Taonura flabelliformis 143 T. robusta 30, 3 tarentina, Latrunculia (Negombata) .... 38, 40 T. seychellensis 30, 31, 32 Tedania 87, 96, 97, 98 Tetillu 28 T. anhelans 87, 89, 92, 96, 100 T. microxea 29 T. anhelans forma assabensis 90 T. inonodi 29 7". assabensis 90 T. oxeata 29

T. commixta . 92 T. praecipua 28, 29 T. digitata 89, 92 Thalassomora 14 T. digitata var. fibrosa 90 Thalyseurypon 56 T. digitata forma inermis 90 Thalysias 71 T. digitata forma polytyla 90 T. juniperina 58 T. digitata var. rubicunda 90 T. transiens 57 T. digitata var. rubra 90 Thoosa 38, 39 T. digitata var. verrucosa 90 T. laeviaster 38 1

242 <. WII 1)1 NMAVCK

140 I . mollis 38, 39 tubercutatus, Stelospongus Thorecta 134, 137, 139 tuberculatus, Thorecta 140, 141

r. boleta 137 Tubulodigitus 104 /'. boletus 137, 139 T. communis 115 '/'. choanoides 130, 137 luhulosa, Echinoclathria 66 /'. exemplum var, secunda 139 tubulosa, < )phlitaspongia 66 /'. exemplum var. tertia 139 tumlda, Phakellia 4X /'. farlovi 1.39, 142, 143 tumida, Pseudaxinellal 4X

I tylostrongylal, Mycale X4, 87 . freija (correctly Aplysina lendenfeldi) 155 105, /. glomerosus 13'>, 1 40 typica, Ceraochalina 104, 115 T. horridus (correctly Mycale (Arenochalina) typica, Chalinopora 104, 110, 113 laxissima)

/'. ? murravi'! 140 typica, Euchalina 104, 1 13

var. haphis, ( I. rimosa (correctly Fasciospongia) . ... 141 typica mcgai lialinopora .... 110 /'. tuberculatus 140, 141 typica, Rhaphoxya 53 /. typicus 139 typica var. tenuispina, Chalinopora IK) Thorectandra 134, 137, 139, 140 typicus, Thorecta 139

I choanoides 1 37 vagabunda, Spiraslrella 34

1 . corticatus 139 validissima, Dasychalina 105 topsenti, ( 'orticium 13 vansoesti, Psammoclema 79, 80, 82 torresia, / endenfeldial 127 vusijorims, Geelongia 137 tortugasense, Merriamium 94, 95 vasiformis, Phyllospongia 13 roxadocia 105 venela, Myxilla 97 toxifera, Callyspongia 114, 117 vermicularis, Carteriospongia 127 Toxochalina 104 vermicularis, ( 'arteriospongia cf 127

I vcrmicularis, 127 . difficilis 113 Phyllospongia Trachycladus 39, 53, 54 vermicularis, Phyllospongia (Carteriospongia) 127 /. bacterium 54 Verongia 154, 155

/ V. 146 . digitatus 53 rosea (recte Aplysilla)

/'. i l> i t 1 1 ; 1 1 us var. clavatus 54 verrucosa, I'edania digilala var 90 /'. digitatus var. gracilis 54 vestigium, Stelletta 20 r. digitatus var. strongylatus 54 villosu, Aulena 144 /'. fastigatus 54 villosa, Dysidea 61 /'. taevispirulifer S3, 54, 61, 97 villosa, Euspongia irregularis var 121, 122 /'. pustulosus 54 villosa, Halme 61 /'. reteporosus 54 virgula, Rhabdastrella 23 /'. scabrosus 54 viscosa, Darwinclla 148, 149 /'. stylifer 54 Waldoschmittia 94, 95, 96 Trachytedania 87 W. schmidti 95 transiens, ( 'lathria 57 wellsi, Ax'mosiu 62 transiens, Thalysias 57 wellsi, Phakettia 62 transitoria, Spiraslrella 34 Weinioreus? 56 trichotoma, Chalina polychotoma var. 112, 113 Wilsonella 56 trilabis, Forcepia (Ectoforcepia) 93 w ilsoni, Suberites 34, 38 truncata, ( allyspongia 105 wilsoni var. albidus, Suberites 34 truncatella var, laxa, Cacochalina 84 wflsooi, Clatbria 57, 58 truncatella var, mollissima, Cacochalina 69, 71 wilsoni, Phoriospongia 65, 76 luha 103, 104 Zetekispongia 94, 95, 96 r. armigera 105 /. zonca 94, 95, 96 tubaria, Sigmatella australis var 74 zimocca irregularis, Spongia 122 tuberculata, ( 'allyspongia 105 zimocca, Spongia 123 tuberculata, Jaspis 26 zonca, Zetekispongia 94, 95, 96 tuberculata, Stellettinopsis 26, 2S Zygomycale 84 tuberculatus, Coppatias 26