Australian Biological Resources Study

Australian Faunal Directory




Regional Maps

Phylum PORIFERA Grant, 1836


Compiler and date details

February 2012 - John N.A. Hooper, Queensland Museum

2005 - Updated by John N.A. Hooper

30 December 2002 - John N.A. Hooper, Queensland Museum, Brisbane, Queensland, Australia

1994 - Felix Wiedenmayer, Naturhistorisches Museum Basel, Basel, Switzerland


1994 Introduction to the Zoological Catalogue of Australia, Porifera

The sponges, phylum Porifera, are predominantly marine with a relatively small number of incursives to fresh water ecosystems. They represent a level of cellular organisation unique in the animal kingdom, with a layer of collar cells (choanocytes) forming chambers within an internal water-filled system of canals. Early accounts of their structure likened sponges to 'clones' of single-celled animals and their recognition as a distinctive phylum was adopted widely only from the early twentieth century. Indeed, they were included as the Spongida, Spongiida or Spongiae in the Protozoa in earlier volumes of the Zoological Record. In the 1872 volume, Lütken noted that the Spongozoa 'should probably form a class by themselves', and only in 1876 were they elevated to their own section of the Zoological Record.

Adult sponges are sessile—although slow movement has been recorded for some species (Fishelson 1981). Dispersal occurs through a larval stage which is predominantly demersal, possibly also pelagic in a few groups. Dispersal may also occur through fragmentation and attachment of adults and this method may be the predominant strategy for recruitment, at least in shallow water populations. Despite their unique structural and biological characteristics, sponges have attracted the attention of few systematists. In consequence, knowledge of their diversity and distribution is patchy.

The taxonomy of Porifera is still not resolved, with the validity of many species and higher taxonomy not satisfactorily established. Differences in interpretation of the importance of various characters have produced many widely divergent taxonomies. These, together with the frequent character convergences and secondary losses, make sponge identifications difficult even for the specialist biologist. Similarly, external characters such as growth form and colour are sometimes affected by local conditions, producing an apparent morphological plasticity in some groups.

Early studies of live populations (e.g. Vosmaer 1932–1955) perhaps over-emphasised the intra-specific variability, to the point where identification of particular taxa became nearly impossible (i.e. with sibling species and allied genera merging into each other). More recent empirical evidence suggests that even subtle differences between populations may be indicative of well defined biochemical and fixed genetic differences, with the likelihood that many so-called geographic variants or varieties of species—and certainly many of the so-called cosmopolitan species—consist of divergent sibling species (e.g. Hooper et al. 1990; Solé-Cava et al. 1991, 1992; Van Soest et al. 1991).

A few groups of sponges have characteristic growth forms (Hooper & Wiedenmayer 1994: figs 3, 4, 146, 183), although most lower level taxa display considerable variety in growth form among allied species and only rarely are higher taxa characterised by ‘typical’ morphologies. The existing taxonomy is based predominantly on skeletal features, with skeletal architecture, spicule geometry and spicule localisation traditionally considered of most importance. Within the three groups of sponges there are many hundreds of spicule geometries, with a complex associated nomenclature. The major spicule forms are illustrated in Figs 5–145, 147–182, 184–230. Although other non-skeletal characters are becoming increasingly useful in definition of taxa at all levels, it is still true, as stated many years ago by Dendy (1924), that for most groups spicules are excellent indicators of phylogeny. One major problem within existing sponge taxonomy, however, is a prevalence of poorly known, inadequately or inaccurately described species making the literature—especially the older Australian literature—unreliable.

Interest in Porifera has escalated phenomenally during the last decade, largely as a result of the rapidly growing marine natural products industry, the abnormally high proportion of ‘biologically active’ members (as compared with other phyla—Garson, 1993), and the remarkable resurgence of interest in marine biodiversity during the 1990s.

Worldwide, the described sponge fauna numbers around 6000 species, although three times this number of species may be extant. Division of the phylum into three classes—the Calcarea (the calcareous sponges), the Hexactinellida (the glass sponges) and the Demospongiae—is generally accepted, but taxonomic ordering at lower levels is not yet clearly resolved. All three classes are represented in Australia. This work records 1416 'valid' species in the Australian fauna, including representatives in Australian extra-territorial waters, with a nominal 2378 species available names.

At the outset of work towards this volume in 1981, the Australian Porifera had not been worked on substantially since the late 1800s, and there were already more than 2200 species names relating to the fauna in the literature. Our common experience, when trying to identify Australian sponges in a contemporary framework, was that reliable identifications could be achieved only with reasonable confidence if preceded by a comprehensive review of all the literature on marine sponges of Australia, its territories, and adjacent regions, and a working knowledge of the vast and scattered type collections.

Until the present volume, no useful modern review has been available covering or including the vast Australasian sponge fauna except for several higher taxa (Calcarea (Burton 1963) and the Keratose orders (Bergquist 1980a)), and isolated genera and families (Hooper 1984a, 1984b, 1986a, 1990b, 1991, 1992; Van Soest et al. 1991). The world review of sponges in de Laubenfels (1936a), and his work on the keratose sponges (de Laubenfels 1948), were too idiosyncratic and fraught with errors to be useful, although they did provide the first comprehensive access to the sponge nomenclature.

Two unpublished checklists of sponges of mainland Australia formed the basis of the present study, one by Ian G. Skinner (formerly of Roche Research Institute of Marine Pharmacology, Sydney), and the other by Avril L. Ayling (Sea Research, Daintree). Together the lists included only about 150 taxa. Subsequent studies (1994) found 2410 species and subspecies ('variety') names, and 708 generic names recorded for the Australian fauna. and since then (1999) these numbers have risen to over 2500 and 730, respectively. This figure is probably the closest we will achieve to having documented all the Australian sponge literature of the past (although there are still probably one or two we have missed), which now provides us with a sound platform for the future.

The Porifera section of the Zoological Catalogue of Australia database, and the book derived from it (Hooper & Wiedenmayer 1994), developed from the list of published nominal taxa to include a comprehensive, independent and critical survey of type collections (where possible). Thus, 1416 species group names, and 337 genus and subgenus group names are accepted here as valid; the remainder are junior synonyms, homonyms and nomina nuda. Yet the Australian sponge fauna is still relatively poorly known, with many unpublished taxa now housed in the Australian museum collections (especially QM). Hooper & Lévi (1994) suggest that probably less than one third of the fauna has been documented. Moreover, since most of the original systematic-descriptive work for Australian sponges was published before 1916, revisionary work on the fauna is as important a task as discovering and documenting numerous new taxa.

This work does not purport to present a phylogenetic treatment of the Australian Porifera. Rather, it is a documentation of available information on species group names in or associated with the Australian sponge fauna. Taxa are arranged alphabetically within classes and families.

1994 - Nomenclature and Classification published in the ZCA Porifera
This work is not a guide to the classification of Porifera, although such a 'Systema Porifera' is exactly what the phylum now needs, including diagnoses and illustrations of all supra-specific taxa. Rather, this is a compilation and analysis in a contemporary framework, of all primary and subsequent records of Porifera, for Australia and its territories, with extra-limital notes, records, references, and synonymies relevant to this fauna. Its intent is to open the phylum to future workers, without them having to 'reinvent the wheel'. Inevitably, because many Australasian sponge species have not been revised since they were first described, it was impossible to avoid creating new names and new combinations in this volume, although we explicitly tried to refrain from doing so. Some of the nomenclatural changes proposed in this volume, necessitated by compliance to the ICZN (1985), are likely to be challenged by some contemporary workers, but changes have only been made where they cannot be avoided (i.e. where there is direct conflict with the Articles of the ICZN). Consequently, an attempt has been made to include the most recent revisions in the classification presented here.

The higher systematics of most sponge groups are still not fully resolved, with contrary opinions being driven by such things as the discovery of many new, useful, non-morphological characters, the controversy between acceptance of a phylogenetically based classification versus an evolutionary biology classification, as well as the previously poor adherence to the ICZN. There is no consensus amongst modern workers concerning many details in a classification (e.g. valid genera, families, orders), and it is unlikely that this will be resolved in the short term. The classification presented here is certainly not definitive, but is intended to offer a sound base upon which to build. To this end we have followed the most recently presented comprehensive scheme for the group (Hartman 1982), with several more recent emendments.

Sponge taxonomy, especially of the Australian fauna, has been characterised by an ignorance of, or a disregard for, the rules of zoological nomenclature. This is especially evident for: the rule of priority (or seniority) of taxon names at the family level and below; the use of junior synonyms, or introduction of new names, simply because the scope of the original description was inadequate; and consideration that a genus is objectively defined by its type species (and only subjectively defined by its original and revised diagnoses). These problems are mostly confined to the systematic papers of Lendenfeld and Hallmann, as outlined above, but these authors were amongst the most prolific. Some other cases occur elsewhere, such as Carter (1886h: 455), replacing Esperia parasitica by Pseudoesperia enigmatica, for no valid reason, but these instances are rare. In the present work we follow the rules and recommendations of the ICZN. Some workers may see our actions as 'destabilising' the nomenclature (again), but we accept that this consequence is of short term.

In the last two decades, several publications have contributed significantly to new perspectives on the higher taxonomy of Porifera. Incorporation of new characters discovered through use of modern techniques (such as transmission and scanning electron microscopy), as well as studies on hitherto neglected characters (for example reproductive and embryological, biochemical and molecular characters) has contributed much to the development of a more heuristic classification of sponges. The authors at the forefront of this research are Prof. Claude Lévi (MNHP, Paris) and Prof. Dame Patricia R. Bergquist (University of Auckland). In a different manner, the taxonomic framework of sponges has been irrevocably changed by the cladistic school of systematics. Although not universally expounded (or understood) by many contemporary workers, it is incontrovertible that the interpretation of all characters used in sponge taxonomy be based on objective, repeatable, scientific principles as offered by phylogenetic systematics. The author responsible for most progress in this regard is Dr. Rob Van Soest (University of Amsterdam).

The present work builds upon the classification summarised by Hartman (1982), which in turn was based on the earlier works of Lévi in Brien et al. (1973) and Bergquist (1978), although we are limited to presenting a classification only as it pertains to the Australian fauna. Thus, these major works are included here, whereas Burton's (1963) controversial classification of is rejected outright. The arrangement within the Calcarea in the Catalogue essentially follows Dendy & Row (1913), as modified by Hartman (1958) and Borojevic (1966–1990). It should be emphasised again that, given the preliminary state of taxonomic knowledge of the marine sponge fauna of Australia, the present arrangement is merely a beginning. It leaves much room for subsequent revisions, for which the computer-based format of this work is particularly suited.

At both genus and species levels, where the extremes of splitting (as in early work of Carter, Lendenfeld and Dendy) and lumping (for example by Vosmaer and Burton) are particularly prevalent in works on the Australian fauna, we decided to lean towards the splitting side. This decision was made chiefly because we consider that, in the absence of specific revisions for all components of this fauna, it is preferable to distinguish differences than to submerge them (Hartman 1964: 712). There is also empirical evidence now to show that even small morphological differences between allopatric populations translate to relatively major biochemical or genetic differences indicative of sibling species relationships (e.g. Hooper et al. 1991; Solé-Cava et al. 1991). However, there is still a great deal to be done in the interpretation of biochemical/genetic data in a biological context. Similarly, computer-based arrangement makes data-retrieval, particularly on distributions, more meaningful with higher resolution of valid taxa at this level.

1994 - Historical Overview
In order to assess the status of contemporary sponge systematics, it is important to know the contributors, the scope of their work, and the fates of their collections. The contributions of the major early workers and their collections, therefore, are discussed here in some detail. By grouping publications separately on regional and systematic bases, the strengths and deficiencies in knowledge and the small number of contributors to earlier and contemporary taxonomy of Australian sponges are apparent.

Grouped geographically, the major original contributions to local Australian faunas are:

Victoria (mainly Port Phillip and Westernport Bays: Carter 1885–1887; Lendenfeld 1883–1889; Dendy 1891–1897.

Bass Strait and Tasmania: (Lamarck 1813–1814 (redescribed by Topsent 1930, 1933); Haeckel 1872; Selenka 1867; Marshall 1880; the Challenger authors 1884–1888; Shaw 1927; Guiler 1950; Wiedenmayer 1989, 1994.

New South Wales (particularly Port Jackson): Lendenfeld 1884–1889; Whitelegge 1901–1907; Hallmann 1914–1920; Bergquist 1980b.

East coast of Queensland (inter-reef and shelf): Ridley 1884a; Lendenfeld 1889b; Hooper 1987, 1991, 1996; Hooper & Bergquist 1992; Hooper & Lévi 1989; Van Soest et al. 1991.

Great Barrier Reef (coral reef associated sponges): Burton 1934; Bergquist 1969; Wilkinson 1978; Ayling 1982; Pulitzer-Finali 1982; Thompson et al. 1987; Bergquist et al. 1988; Stoddart 1989; Wilkinson & Cheshire 1989; Fromont 1989, 1991, 1993; Hooper 1990b; Bergquist & Kelly-Borges 1991; Hooper & Bergquist 1992; Hooper & Lévi 1993a, 1993b.

Western Gulf of Carpentaria, Cape York and Torres Strait: Ridley 1884a; the Challenger authors 1884–1888; Kieschnick 1900 (some revised in Hooper 1991, 1996).

Northwest coast (Cape Londonderry to eastern Gulf of Carpentaria): Ridley 1884a; Bergquist & Tizard 1967; Hooper 1984a, 1986a, 1986b, 1987, 1988, 1991, 1996, Hooper, Capon & Hodder 1991; Hooper & Bergquist 1992; Hooper et al. 1997.

Northwest Shelf: (Northwest Cape to Bonaparte Archipelago): Hooper 1984a, 1984b, 1986a, 1986b, 1991, 1996.

Southern Western Australia—Albany to Shark Bay: Lamarck 1813–1814 (revised by Topsent 1930–1933); Bowerbank 1841–1876; German expeditions (Hentschel 1909–1911; Lendenfeld 1907; Row & Hôzawa 1931—Houtman Abrolhos Dendy & Frederick 1924; Hooper, 1996; Hooper & Krasochin 1989; Hooper & Bergquist 1992.

South Australia: Lamarck 1813–1814 (revised by Topsent 1930–1933); Haeckel 1872; Bergquist & Skinner in Shepherd & Thomas 1982; Hooper 1991, 1996; Reiswig 1992; Wiedenmayer, 1994.

Tropical islands and island territories of Australia: Kelly-Borges & Valentine 1996.

Hooper & Lévi (1994) hypothesised that sponges have much more restricted distributions than recognised (e.g. Burton 1934), and that there are many more 'pockets of endemism' around the Australian coast. This hypothesis has empirical support from comparative studies of many regional faunas in tropical Australia, but further corroborative evidence is needed. Several substantial and well curated collections of sponges from most Australian marine provinces are now available in Australian museums, and it is hoped that future effort will be directed to utilising and documenting this existing resource.

Faunas for which good contemporary collections are held include:

Northwest continental (from Cape York all the way into the Northwest Shelf, to Northwest Cape, WA). Collections of over 4000 specimens from this region, including some 800 species, are in the Northern Territory Museum of Arts and Sciences, Darwin (NTM), but only a small proportion of these have been published (Hooper 1984b et seq.; Hooper et al. 1997).

Northwest oceanic coral reef. Good collections of sponges, from oceanic coral reefs on the northwest margin of the continent, as well as Christmas and Cocos-Keeling Islands, are now in the Western Australian Museum (WAM) and NTM. Reports on some of these species are published (Ashmore, Cartier, Hibernia Reefs: Hooper 1994), but most taxa have been sorted only to genus.

Gulf of Carpentaria. Several recent major trawling surveys in this region by PIBOC RV Akademik Oparin (1989) and CSIRO RV Southern Surveyor (1990–1998) revealed patchy distributions of sponges, associated mostly with the hard benthos. Most sediments in deeper waters of the Gulf (›40 m) consist of soft mud and gravel, and few species appear to live in these softer sediments. The collections, now comprising about 300 species, are still largely unpublished and are housed in the NTM and Queensland Museum, Brisbane (QM). Work on some families in this fauna is underway (also Hooper 1991, 1996).

Northeastern continental shelf (inter-reef), and deeper water continental slope. Cannon et al. (1987) made substantial collections of inter-reef sponges from the Cairns region. These were sorted roughly to genus, but remain largely unidentified (Hooper 1987, 1991, 1996). Subsequent large, fully documented collections from this same region, indicate that the fauna of the inter-reef region comprises over 200 species and is largely different in species composition from both the adjacent coral reef fauna on the Great Barrier Reef and the Northwest Shelf fauna at equivalent latitudes and depths. Work on these collections in the QM is in progress. Several collections of deep water sponges from the continental shelf and slope are housed in the Museum of Tropical Queensland, Townsville (CIDARIS collections).

Northeastern coral reef. The macrobenthic sponge fauna associated with coral reefs is relatively well published in comparison to other habitats (see list above). Only recently has this fauna been actively documented in a contemporary manner, with over 800 coral reef sponge species now contained in voucher collections in the Queensland Museum. Most type specimens of coral reef-associated sponges are housed in the Natural History Museum, London (BMNH), but many more types from contemporary collections are being accumulated in the QM Brisbane and MTQ Townsville. This fauna is currently the most actively researched, possibly containing the greatest species diversity of all Australasian provinces (Fromont 1991, 1993; Hooper 1991, 1996; Hooper & Bergquist 1992; Hooper & Lévi 1993a, 1993b, 1994).

Southern Queensland and northern New South Wales (between Fraser Island and the Solitary Islands). The QM now holds extensive 'sorted collections' from this region (to the level of genus level and a 'unique species number'). Investigations on this fauna are also actively in progress. This region is of particular biogeographic significance, being an overlap zone between the Solanderian and Peronian provinces, because there is virtually no overlap in species' distributions, and very sharp species' boundaries between the faunas have been discovered (Hooper & Lévi 1994; Hooper, unpublished data).

Southwestern Australia. Large collections from this fauna are in the WAM, but until recently much of it unidentified, or has been sorted only to genus. More recently Jane Fromont has been researching and documenting certain sponge taxa from this region (e.g. Fromont 1998). This region is also particularly interesting given the many pockets of apparent endemism in sponges along the western continental margin (Hooper & Lévi 1994).

Great Australian Bight. Substantial collections from this region are now held in the South Australian Museum (SAMA); most taxa have been sorted only to genus.

Bass Strait, eastern Victoria, southern New South Wales, Norfolk and Lord Howe Islands. Good collections of macrobenthic sponges now exist for southern Australia in general (Museum of Victoria (NMV), Roche Research Institute of Marine Pharmacology Collection (RRIMP, now in the Australian Museum (AM), Sydney), and Australian Institute of Marine Sciences, Townsville (AIMS NCI) collections), but the precise affinities of these faunas are still uncertain (see Hooper 1991). Only a very small proportion of these contemporary collections has been published (e.g. Bergquist 1980a et seq.; Wiedenmayer 1989). Extensive collections of sponges from recent trawl surveys in Bass Strait and Victorian waters remain largely unsorted and unidentified, and similarly, much of the early material from this region collected by Carter and Dendy and housed in the NMV also remains unsorted and unregistered. These southern Australian collections (South Australia, Victoria, Tasmania) require urgent attention.

Antarctic territories and Subantarctic islands. In comparison with the tropical fauna, the sponge fauna of the Antarctic and Subantarctic territories is relatively well known. This fauna comprises about 500 described species, of which over 200 are known to live in the Australian territories. Most published collections are deposited in overseas institutions, although a small collection of Koltun (1976) and duplicate samples of Burton (1929) are housed in the SAM and AM, respectively. Principal references for this fauna are: Brøndsted (1923, 1926); Burton (1929, 1932, 1934, 1938); Desqueyroux (1972, 1975); Hentschel (1914); Koltun (1964, 1976); Sarà (1978); and Topsent (1901, 1908, 1913, 1916, 1917).

There is no useful secondary literature available for identification of sponges, save for the comprehensive treatment of families by Hartman (1982), and a (still incomplete) 'Sponguide' to genera and families on the internet (; Hooper 1991-8). Access to the generic and species level classifications is still more or less restricted to specialist workers. Reviews and revisions of supra-specific taxa either containing Australian representatives, or useful for the taxonomy of this fauna, include the following:

Hexactinellida: Ijima 1927; Lévi 1964; Reiswig 1992.

Calcarea: Dendy & Row 1913; Burton 1963; Borojevic 1968b; Borojevic, Boury-Esnault & Vacelet 1990.

Astrophorida and Spirophorida: Lendenfeld 1903.

Hadromerida: Tethyidae, Bergquist & Kelly-Borges 1991; Hemiasterellidae, Hooper 1986a; Voultsiadou-Koukoura & Van Soest 1991; Polymastiidae, Kelly-Borges & Bergquist 1997; Latrunculiidae, Kelly-Borges & Vacelet 1995; 'Lithistida', Kelly-Borges & Pomponi 1994; Suberitidae, Kelly-Borges & Bergquist 1994, Fromont 1998.

Haplosclerida and Petrosida: Lendenfeld 1887a; Burton 1927, 1934; Bergquist & Warne 1980; Van Soest 1980; Hooper 1984b; Fromont 1991, 1993.

Dictyoceratida, Dendroceratida and Verongida ('keratose' sponges): Bergquist 1980b, 1995; Bergquist et al. 1988, Bergquist & Kelly-Borges 1995.

Halichondrida (including Axinellidae, Desmoxyidae): Hallmann 1916-1917; Bergquist & Hartman 1969; Van Soest et al. 1990; Hooper et al. 1992; Hooper & Bergquist 1992; Hooper & Lévi 1993b; Hooper et al. 1997.

Poecilosclerida: general, Van Soest 1984; Bergquist & Fromont 1988; Microcionidae, Hallmann 1920; Hooper 1988b, 1990a, 1996, Hooper & Lévi 1993a; Raspailiidae, Bergquist 1970; Hooper 1991; Rhabderemiidae, Hooper 1990b; Van Soest & Hooper 1993; Desmacellidae, Bergquist 1970; Hooper 1984b; Hooper, Capon & Hodder 1991; Myxillidae , Hooper 1987; Van Soest et al. 1991; Desmacididae, Wiedenmayer 1989; Anchinoidae, Voultsiadou-Koukoura & Van Soest 1991.

Early taxonomists and their collections
Six prominent names are associated with early work on Australian sponges: Lamarck, Carter, Dendy, Lendenfeld, Whitelegge and Hallmann.

J.B.P. de Monet Lamarck. A 1988 survey of Lamarck's (1813-1814, 1814-1815, 1816) important Australian sponge collections in the MNHN Paris, found that most material was intact, in relatively good condition (behind glass), and probably largely untouched since revised by Topsent (1930, 1932, 1933). This material is presently in the new underground zoothéque of the MNHN, with replacement computer-printed labels now included for each specimen. All Lamarck's sponge material has been re-photographed and microscope slide preparations have been made (QM Brisbane). There is also a nearly complete set of microscope slide preparations of this material in the BMNH, although most of the section preparations were found not to be particularly useful. Some of the Lamarck material has already been redescribed in modern terms (Raspailiidae: Hooper 1991; Axinellidae: Hooper & Lévi 1993b), or is in preparation for publication (Microcionidae: Hooper, in preparation). However, this task is not yet completed. Despite earlier examinations of Lamarck's species by Ridley (unpublished notes in the BMNH; some revised taxa in 1884a) and Topsent (1930-1933), there are still a number of senior synonyms (Lamarck names) for Australian taxa which have not yet been settled (particularly names for several of Lendenfeld and Hallmann's southern Australian species).

H.J. Carter. Carter worked in retirement (see Wiedenmayer 1977: 253), publishing in his characteristic piecemeal fashion, almost exclusively in the Annals and Magazine of Natural History (see Dendy 1889), in which plates were a luxury. Carter never designated holotypes, but many of his descriptions were based on particular specimens (often including external measurements) and he was careful in labelling specimens. For the J.B. Wilson Collection (from Port Phillip Heads and Westernport Bay, described by Carter 1885–7), he wrote a Catalogue, listing all specimens, wet and dry, with his own identifications and running numbers (his own and J.B. Wilson's). The Manuscript Catalogue also contains additions in Ridley's hand (then Assistant Keeper in the BMNH), and a copy of it is now deposited in the Sponge Archive (NMV Sponge Archive 9/3).

Good agreement has generally been found between this catalogue, the register entries, and surveys of the specimens, the dry ones all carefully labelled by Carter himself. In a few cases, more specimens are registered than are listed by Carter, which probably means that Carter had already sectioned these specimens, and the fragments were inadvertently registered separately. However, there are occasionally more serious discrepancies between the Manuscript Catalogue and Carter's published descriptions, in the number of specimens and mode of preservation (wet/dry), invariably with Carter mentioning fewer specimens in publication. Yet Carter mentions in his introduction to the Manuscript Catalogue that all the specimens listed are labelled in accordance with his published accounts. The salient examples of such discrepancies are Stelospongus flabelliformis, sensu lato, and Pseudoceratina durissima. The inescapable conclusion is that if Carter mentioned a number of specimens in a description, he referred only to the specimens dissected and examined microscopically, and he probably identified the remaining ones only superficially.

Specimens of Carter's Australian species fall into three groups. The first is the Cawne Warren Collection from Bass Strait (Carter 1881), formerly in the Liverpool Free Museum (LFM; now Merseyside County Museums). Most of the specimens were destroyed by bombing raids during World War II, but some of the type material survives in the form of slides in Carter's cabinet at the BMNH, and a small number of specimens are also held at the BMNH.

The second group comprises species described before 1885 from old collections in the BMNH, mostly Bowerbank's material. The specimens are mostly dry, and are also represented by slides in Carter's cabinet. This cabinet is kept separately from the main systematic slide collection in the BMNH, and Carter's slides (including extra-limital species, some with mounted fragments) are registered as BMNH 1954.3.9.1–627 (balsam mounts) and 1954.3.9.628–823 (fragments on wooden slides) (NMV Sponge Archive 9/2).

The J.B. Wilson collection from Victoria is the third group and by far the largest. Parts of this collection were described from 1885–1887 (see Wilson 1895; much more unregistered material of J.B. Wilson still exists in the NMV). The named demosponge specimens of this group are registered as BMNH 1886.12.15.1–151 (wet), 1886.12.15.152–508 (dry); and 1887.7.11.1–24 (wet), 1887.7.11.25–30 (dry). A few are registered with the prefix 1884.10.10 (NMV Sponge Archive 9/2 10/2, 3). The Calcarea of this group are registered BMNH 1887.7.12.1–81 (see Burton 1963: 577).

The BMNH also holds a large collection of microslides of Victorian sponges, mostly from Port Phillip, purchased from the estate of J. Bracebridge Wilson, Geelong (registered 1897.5.4.1–801). It is not known whether these slides were used by Carter (1885–1887) and/or Dendy (1891–1897) for their descriptions. Similarly, a collection of fragments of BMNH type specimens from Victoria, collected by J.B. Wilson and described by Carter, was given to A. Dendy in Melbourne by Ralph Kirkpatrick (at the time, Assistant Keeper). Dendy obviously made his own slides from these, but later gave the remaining fragments to the Australian Museum, Sydney (AM), where T. Whitelegge produced a complete set of slides. This latter set is still in the AM, and registered AM G2731–2954, almost all with references to the BMNH register numbers (NMV Sponge Archive 9/1, 10/1).

A. Dendy. Dendy's descriptions of southern Australian species, like those of Carter, suffer from brevity and lack of illustrations, many being inadequate for present needs. The only exception is Dendy's earliest work on Victorian Calcarea (1891), which is lavishly illustrated. As Dendy himself made clear (1893a: 69; 1895: 232), the decline in standard of his presentations was directly related to the decline in funds available for publications during this period, and to the overwhelming diversity of material remaining to be described. Dendy was unusually meticulous for his time in publishing his own 'RN' (or Register Number) running numbers at the end of each description, so that most of his type specimens and even slides can be collated with museum register numbers (see Ayling et al. 1982).

Dendy's type material is now split between the BMNH London and the NMV Melbourne; information on this is easily available (Ayling, Stone & Smith 1982). All material previously housed by the University of Melbourne has now been deposited in the NMV collections (NMV Sponge archive 23). It comprises 91 slides of Calcarea (including probable fragments of type material of Carter, Poléjaeff, and Dendy) and eight slides of Demospongiae.

R. von Lendenfeld. Lendenfeld's original work on Australian sponges (1883–1889) is generally considered to be careless by contemporary workers. There are documented instances of where his work is completely wrong (e.g. where published descriptions do not match type specimens; type series of a single taxon consist of several species, none or only some of which match original descriptions; or type specimens do not even belong to the alleged order (Hooper 1991, and work in preparation on Microcionidae)). Conversely, Lendenfeld also produced excellent, perfectly recognisable descriptions of many species, particularly many —keratose' sponges, accompanied by field notes that are accurate even by modern standards.

Many of Lendenfeld's species have been revised by subsequent authors: Demospongiae (Whitelegge 1901–1907; Hallmann 1912–20; Burton 1927, 1934; Bergquist 1980b; Hooper, 1991) and Calcarea (Dendy & Row 1913; Burton 1963). These revisions are far from complete. Lendenfeld's types in Sydney, London and East Berlin need to be re-examined; their labelling is often ambiguous, so that matching 'apparent types' with descriptions (if at all possible) is essential.

Hallmann (1912) noted that in cases where Lendenfeld based his descriptions on two or more specimens, he appears to have taken the microscopical characters from one, and the external traits from the remainder or from all. Our collective surveys of Lendenfeld's types in the AM and BMNH confirmed that this was indeed likely. Thus, for many of Lendenfeld's species, the type series exceeds two specimens and is often composite, even to the unaided eye. Unfortunately much work is still required to determine the status of many of Lendenfeld's 'types' in the BMNH, AM and the Museum für Naturkunde an der Universität Humbolt zu Berlin (ZMB).

In the introduction to his largest monograph, Lendenfeld (1889b) gives a short biogeographical account of his years in Australia and New Zealand (1882–6) and the intervening three years before the publication of the monograph. He states that he examined every specimen in the BMNH belonging to the horny sponges (i.e. his sense of the 'keratose' sponges, including siliceous poecilosclerid and haplosclerid genera Echinoclathria, Phoriospongia, Psammoclema and Dactylia). Among these were type specimens of Gray, Bowerbank, Poléjaeff, Ridley, and Carter, and also countless undescribed specimens deposited there in Gray's and Bowerbank's time, and before 1873 (the year of the Challenger Expedition). Many of the latter are now primary types of Lendenfeld's (1889b) species, as he acknowledges at the end of relevant descriptions, under the heading 'Distribution'.

During the three years before the monograph appeared in print, Lendenfeld also completed the manuscripts of his two other major monographs (1887a, 1888), which are based largely on his own collection (1887a, now scattered among AM, BMNH, and ZMB), and that of the AM (1888). Both publications were delayed, we are informed in the respective introductions, because Lendenfeld wished to update his identifications and descriptions following his survey of Australian specimens in the BMNH described by others. The result was idiosyncratic: whenever he discovered that one of his species had already been described, he either adopted the old name, but with himself as author (in cases where the scope of a single senior synonym was changed), or applied a new name (whenever he combined different senior synonyms in one species). Dendy, in his 1887a work, criticised this procedure, but Lendenfeld (1887b) defended himself by claiming that his nomeclature was more logical than that of Ridley & Dendy (1887).

Lendenfeld was also renowned for his habit of changing names of species and genera between the time of labelling specimens and publication of descriptions. This sometimes occurred twice, with a first version upon collecting, changed at cataloguing, and again in publication. Whitelegge (1901: 64) complained about this practice, and Lendenfeld, approached by the Trustees of the AM, then supplied a 'Key List' collating manuscript and published names. This 'Key List' is mentioned repeatedly by Whitelegge (1901–1907) and Hallmann (1912–1920), but it now appears to be lost (Elizabeth Pope and Frank Rowe, personal communication). Fortunately, many of the name changes can be reconstructed from subsequent collations by Whitelegge (who worked with Lendenfeld in Sydney), and from Lendenfeld's specimen labels (often multiple) with specimens in the BMNH and in the AM. Nevertheless, many of Lendenfeld's meaningless manuscript names remain on specimen labels and in registers in the AM, BMNH and ZMB. Unless the 'Key List' is discovered these problems are unlikely ever to be resolved completely. More importantly, a considerable number of Lendenfeld's species are virtually unrecognisable, and the status of many of his type specimens is still uncertain. In this Catalogue, Lendenfeld's 'uncertain' material is stated as such when we have been unable to resolve a taxon's identity.

The nomenclatural confusion created by Lendenfeld is reflected in his 1887 classification (Lendenfeld 1887d), where many subfamilies (such as Suberamatinae, Spongissinae, Aplysissinae, Hircinissinae, Psammopessinae) are based on manuscript names of genera still found on labels. Similarly, Lendenfeld usually disregarded type specimen collection data in his major publications (especially 1889b), with the result that about 50% of published localities differ from those on specimen labels (and in some instances a third, different locality is written in the AM register). Many of the specimens figured in 1889b are primary types of Carter (1885–1887) and of Ridley (1884a, 1884b); others are those from the older (dry) BMNH Collection mentioned above, while again others (mostly wet) are from Lendenfeld's own collection, which he sold to the BMNH in 1886–1887. Some of these specimens have been refigured by Bergquist (1980b) in an important monograph on the `keratose' sponges, and she has rediscovered and redefined many of Lendenfeld's `species'. However, as stated by Bergquist (1980b), many of Lendenfeld's species names are still mystifying. Collations of problematic specimens with figures are cited in this Catalogue under type-specimen identifiers whenever possible.

The great haste with which Lendenfeld worked is reflected in the confusion of figures in his monograph of 1888. Also, the figures of anatomical details in Lendenfeld (1889b) should all be viewed with caution. More sophisticated, modern histological techniques have since demonstrated that much of the `anatomical detail' was largely imaginative, and unlikely to have been visible given the available (often dry) material and the microscope technology of that era. Conversely, Lendenfeld's accuracy has been proven on several occasions. One such case is Druinella rotunda, where it was found from a re-examination of a type slide (ZMB 10403) that Lendenfeld (1889b, pl. 34, fig. 3) was very accurate in reconstructing and depicting the general aquiferous system of the sponge (contrary to doubts by Bergquist (1965: 138), who was probably unaware of the existence of this type material), although the finer cytological details including choanocyte flagella (Lendenfeld 1889b, pl. 34, fig. 10) are obviously fabricated.

It is interesting to note that F.E. Schulze, Lendenfeld's teacher, was far more conscientious in his published systematics, whereas E. Haeckel, a contemporary of Lendenfeld, was even more idiosyncratic than Lendenfeld in his systematic outlook.

From what is known of Lendenfeld's biography and published works, the Australian specimens described as new species and varieties were in three original collections. These were his private collection (the largest), the collection of the AM at the time of his departure from Australia in 1886 and the previously undescribed dry specimens in the old (pre-1880) collection in the BMNH. He sold his private collection piecemeal to the BMNH, presumably in 1886–1887 (the two years of registry), and to the ZMB. These specimens are documented by the respective register-entries in the BMNH (1886.8.27.1–679, 1887.4.27.1–125) and ZMB (specimens and slides, register nos. 1101–1223, 2254–2331, 3009–3017, 6188, 6200, 6396–8, 6431–7, 644–63, 6467, 6507–10, 6514–9, 6537–45, 6582, 6587, 6591, 6627, 6655, 6684–9, 6693, 6695, 6719–49, 6813–9, 6894–9, 6906–13, 6922–5, 7062, 7071–7, 7086, 7103, 7108–7120, 7130, 7133–6a, 7146–51, 7156, 7166–91, 7221, 10188–93, 10363–10378, 10381–10448, 10453–87). Photocopies of the relevant pages from the BMNH registers are lodged in the NMV Sponge Archive 9/2, and the data from the ZMB were copied out by hand and appear in NMV Sponge Archive 10/3.

During our surveys of type specimens in the BMNH, all data on the labels were checked against the register entries and copied into notebooks (NMV Sponge Archive 10/1–3) or a computer database (QM). Most of Lendenfeld's and Carter's labels also appear on photographs of type specimens, taken by Felix Wiedenmayer, appearing in NMV Sponge Archive 7. The same procedure was adopted for Lendenfeld's types in the AM (NMV Sponge Archive 7, 10/1, 14). A photocopy of many pages from the AM register with Whitelegge's list of collations between BMNH numbers and AM numbers, for fragments of Lendenfeld's BMNH types donated by Dendy, is now held in the NMV Sponge Archive 9/1. The original labels on several of the Lendenfeld types in the AM have been lost or have become illegible, but they are nevertheless documented as such by labels and register entries in Whitelegge's and Hallmann's handwriting.

Unfortunately, it was not possible to survey the entire Lendenfeld type collection in the ZMB, but there is no reason to doubt the accuracy of relevant entries in the register (which also state type localities). Some of these entries, as for some in the BMNH register and specimen labels, specify localities not given by Lendenfeld in the descriptions of the respective species, or in other instances they contradict the published type locality. Some of these have been noted already in publications of Whitelegge (1901–1907), Hallmann (1912–16) and Hooper (1991), and similar discrepancies are noted in this work.

Among Lendenfeld's many manuscript names in the three museum collections, some could be collated with published names, and thus are accepted as referring to type specimens. In part this was possible through Whitelegge's and Hallmann's indications (mostly published) referring to the `Key List' at their disposal (but now lost); partly through double entries on labels and in registers in the BMNH; but the majority of such names remain meaningless in this context.

Most of Lendenfeld's own slides in the BMNH had deteriorated so much that they were destroyed as worthless in 1958. Only 26 were retained, several of which relate to manuscript names (NMV Sponge Archive 9/2).

T. Whitelegge. Both Whitelegge and Hallmann had the unenviable task of trying to identify many species from the extensive trawl and dredging surveys being undertaken at that time in southeastern Australia, with Lendenfeld's published taxa. Each expended large amounts of effort trying to revise (or interpret) his taxa, but neither was completely successful.

Whitelegge (see 1901: 64) co-operated with Lendenfeld during the latter's work in Australia, helping him with collecting, sorting and preliminary curation of the material. This `curation' included provisional cataloguing of Lendenfeld's material by running numbers. Although these numbers were characteristically disregarded by Lendenfeld in his 1888 publication, they were subsequently cited in Whitelegge's 1901–1907 and other revisions. They also appear on Whitelegge's labels with the AM specimens and thus provide, with the remaining data, the only useful pointers to the type status of many of Lendenfeld specimens. Definitive registration of sponge material in the AM was undertaken by Whitelegge, from about 1900 onwards, and was probably completed by Hallmann and other anonymous helpers. Lendenfeld's open, flamboyant handwriting is easily recognisable.

Whitelegge (1901–1907) based his revisions of many of Lendenfeld's species on re-examination of the type specimens in the AM, and of fragments (`schizotypes') of Lendenfeld's other type specimens he had previously sold to the BMNH. These fragments were donated to Dendy by Kirkpatrick from the BMNH, and they were subsequently passed on to the AM where they now exist mainly as microscope slides. To be fair, Whitelegge's lack of success in revising Lendenfeld's taxa, and in the placement of many of his own species, is not solely attributable to the problems that Lendenfeld created for him, but probably stemmed equally from his own relative inexperience with sponges.

E.F. Hallmann. Hallmann was associated with the Australian Museum in 1912–14 (probably as an honourary researcher), and a Linnean Macleay Fellow of the New South Wales Linnean Society from 1914–16. As far as we can ascertain, he was never officially employed by the AM, but he was the last Australian-based sponge worker before 1982. During his brief publication period (1914–20) he contributed several significant revisions (including some of Lendenfeld's species), and descriptions of the fauna mostly in the vicinity of Sydney. There is no doubt that Hallmann's taxonomy was far better than most of his predecessors, although he had the advantage of specialising mainly on a few groups (Poecilosclerida and Halichondrida).

Through his novel method of dealing with the type status of many of Lendenfeld's specimens and taxa, Hallmann created a number of new problems for subsequent workers. When a type specimen was found to disagree substantially with the original description, Hallmann negated its status as a primary type. He usually redescribed such specimens as new species, with Lendenfeld's name conserved only for syntypes found to agree with the original description. Where a holotype only was involved, this was either redescribed by Hallmann as a new species, with Lendenfeld's species being declared unrecognisable (or of doubtful generic affinity), or it was described as a junior synonym.

Two notable examples of this arrangement are Axinella inflata Lendenfeld, 1888 and Reniochalina stalagmitis Lendenfeld, 1888. The wet holotype of Axinella inflata was redescribed and figured by Hallmann (1914c: 427, pl. 23, fig. 5) as a synonym of Chalinodendron dendrilla Hallmann. He noted this decision on a separate label left with the other labels of A. inflata, removed the holotype, and probably placed it in another jar with a new label headed 'Chalinodendron dendrilla'. As a result, all that now remains in the type collection of Axinella inflata is a jar with the labels. It is possible that the holotype is extant in the general collection with the other specimens (`hypotypes'), but neither of us has found it. In a similar way, Lendenfeld's type species and genus of Reniochalina was divested of its type status and redescribed as Axiamon folium gen. et sp. nov. by Hallmann (1914c). This procedure rested on a subjective decision, contrary to the rules of nomenclature, with both Hallmann's new genus and new species names becoming objective synonyms of Reniochalina stalagmitis.

Under our interpretation of the International Code of Zoological Nomenclature (1985; henceforth referred to as the ICZN), Hallmann's new names are valid only if they are based on part of the revised type series of Lendenfeld, where more than one type specimen exists (most species of that era were based on a series of syntypes). If only one unrevised syntype remains, then this is implicitly the lectotype of the original species, and if more unrevised syntypes remain, a lectotype for Lendenfeld's restricted species should eventually be chosen from among these. However, Hallmann's procedure was not quite consistent and in some cases he accepted names and Lendenfeld's type specimens even though he stated in publication that the original description was known to be inaccurate or misleading. Hallmann's actions are understandable given the frustration of dealing with Lendenfeld's nomenclature and type material, but unfortunately many of his decisions are now unacceptable under the rules and recommendations of the ICZN.

Hallmann's frustration with the Lendenfeld collection is further reflected in the discovery of several entries in the AM Register, regarding some 'redundant' 'schizotypes' of Lendenfeld from the BMNH: they were destroyed as worthless because the species was already well represented by complete specimens in the AM. These transgressions contrast with Hallmann's abilities as a taxonomist. His descriptions are explicit, mostly well illustrated, and are generally still useful.

Hallmann (1914) attributed Lendenfeld's numerous mistakes to his poor powers of observation, but for the many Lendenfeld species represented by more than one type specimen, and particularly where the syntypes are obviously disparate to the unaided eye (consisting of more than one species), the reasons for Lendenfeld's failures are probably more complex. Our experiences in surveying Lendenfeld's collections, specimen labels and register notes in the BMNH and AM suggest to us that these problems were probably due to Lendenfeld's haste, his poor organisation of manuscript notes, his excessive reliance on his memory, his use of external traits only in identification of most syntypes, and probably the fact that he was actually paid for his collections by museums and collectors throughout Europe, presumably on a basis of quantities of specimens—hence the proliferation of syntypes of ostensibly the same species (but in many cases composite type series) throughout the European collections.

Hallmann corrected many of Lendenfeld's more flagrant mistakes, noting that they seem to be derived from simple errors of mislabelling. Undoubtedly more errors are as yet undetected, particularly with wet `type specimens' where several are now together in one jar, with labels detached. Many type specimens have printed labels as well as manuscript labels, because they were once on exhibit. During World War II, all sponges were removed from the AM to safer storage, and were returned to the Museum after the war; some of the printed labels became detached (Elizabeth Pope, personal communication when the Curator in charge). The specimens generally have separate little labels with register numbers attached (in tin, with wet specimens). No discrepancies were found during the authors' surveys.

Species described by both Whitelegge (1889, 1897, 1901, 1902a, 1902b, 1906, 1907) and Hallmann (1912, 1914a, 1914b, 1914c, 1916a, 1916b, 1917a, 1917b, 1920), including revisions of Lendenfeld's `type material' and many of their own species, are housed predominantly in the AM. Some duplicate (donated/exchanged) material is in the BMNH. A survey of this material in both museums found most specimens still extant; most of the specimens that could not be found were Lendenfeld's own specimens, nominated as types by Whitelegge or Hallmann, and it is possible that these have not yet been separated from the general collections in the AM.

Other Authors
As far as we are aware, from first hand examination of museum specimens and/or registers, or from personal communications with contemporary sponge workers, the type material of other major published collections of the Australasian sponge fauna is accurately accessioned into the Australian state museum collections or in major museums overseas. Since about 1960 it has been usual practice for Australasian sponge workers to cite voucher specimen numbers in their publications, making the task of locating and checking this material relatively easy.

With few exceptions, Australasian material described by M. Burton, R. Kirkpatrick, S.O. Ridley, S.O. Ridley & A. Dendy, R.W.H. Row, and others is in the BMNH. M.E. Shaw's (1927) and E.R. Guiler's (1950) material from Tasmania is also in the BMNH, with secondary fragments probably retained in the Queen Victoria Museum (QVM), Launceston (although this material has not yet been audited). V.M. Koltun's described Antarctic material, from the BANZARE expeditions (Koltun 1976), is housed in the SAMA. Clive R. Wilkinson's species from the Great Barrier Reef are in the AM, with many more specimens in the AIMS (probably destined for the QM or Museum of Tropical Queensland (MTQ), Townsville).

Patricia R. Bergquist was the first contemporary systematist to work on the Australian sponge fauna. She dealt mainly with species from southeastern Australia and the Great Barrier Reef (e.g. Bergquist 1980b), whilst associated with the Roche Research Institute of Marine Pharmacology, Sydney; several other publications included other faunas (Bergquist & Tizard 1967; Bergquist 1969). Earlier publications of Bergquist and co-workers, including records from the Australian fauna or describing species present in both New Zealand and Australian faunas, noted that type specimens and voucher material was deposited in the National Museum of New Zealand (NMNZ and formerly the Dominion Museum, Wellington) (Bergquist 1961a, 1961b, 1961c, 1965, 1972; also her monographs on the New Zealand fauna: Bergquist 1968, Bergquist et al. 1970, 1980, 1988). Types of other described species are in the AM (Bergquist 1969, 1980b; Bergquist & Tizard 1967; Bergquist et al. 1971, 1988; Thompson et al. 1987; Kelly-Borges & Bergquist 1988). Dawson (1993) recently compiled a comprehensive catalogue for the New Zealand regional fauna.

The year 1981 marked a turning point in Australian sponge taxonomy. The pivotal event was the Sponge Taxonomy Workshop held at the NMV Melbourne in March, co-convened by Patricia Bergquist and Felix Wiedenmayer, and funded by the Australian Biological Resources Study (ABRS). The workshop was devoted to the status and future of sponge taxonomy in Australia. This Catalogue is an indirect result of this workshop, and would never have been attempted without the support and encouragement of ABRS.

In 1981, too, John Hooper began work on sponges at the Northern Territory Museum, Darwin. He and co-workers split their Australian type material between the NTM (Hooper 1984a, 1986a, 1986b, 1987, 1990b, 1991; Hooper et al. 1989, 1990, 1991, 1992) and QM (including MTQ) (Hooper 1987, 1996; Hooper et al. 1992, 1997; Hooper & Lévi 1989, 1993a, 1993b), with several other types in the NMV (Hooper 1984b). In that same year, Jane Fromont commenced studies on Australian sponges, initially associated with the WAM, subsequently at James Cook University, Townsville, and now again at the WAM. Her type material (Fromont 1991, 1993, 1998) is lodged with the MTQ and WAM.

From 1981 through to 1985, Felix Wiedenmayer worked on the documentation and description of the Victorian and Bass Strait faunas. The outcome was a monograph on dominant species of the Maugean sponge fauna (Wiedenmayer 1989), type and voucher material from which was deposited in the NMV, and more recently a treatise on the post-Palaeozoic sponges, with a synthesis of the worldwide fauna and applications to the biogeography of southern Australian species in particular (Wiedenmayer, 1994), and aimed mainly at the higher taxonomic ecological, biogeographical and stratigraphic distributions of sponges. In addition, he prepared the initial data for the Catalogue, lodging much of the raw data accumulated in his survey of sponge types in a Sponge Archive in the NMV. Among these are a large collection of unpublished notes on the BMNH and European museum type collections; photographic negatives of most of these type specimens (with film and frame number referred to in the Catalogue, corresponding to NMV Sponge Archive 7); and an annotated bibliography of about 1500 references, up to 1984, originally intended for publication with the Catalogue (NMV Sponge Archives 1–23). {Hooperj1996a}{Hooperj1997a}


Preliminary gathering of the data, assessment of much of the type material, many taxonomic decisions for the Australian fauna and definitions for higher taxa were completed by Felix Wiedenmayer in 1985. His tenure at the NMV, his travel to London (BMNH), East Berlin (ZMB) and Jena (PMJ) in 1983 to examine type material, and employment of Bronwen J. Scott to transfer most of the original data into computer format, eventually leading to the initial preparation of the Porifera section of the Zoological Catalogue of Australia, were funded by Marine Sciences and Technologies Grant 80/0267 and Australian Biological Resources Study Grants 82/1643 and 82/1642. Volunteer assistance of Natalie Schroeder is also acknowledged for helping with the data transfer process. Dr A.A. Racek compiled the original data sheets for the freshwater sponges.

Dr Barry R. Wilson and Dr Brian J. Smith, both formerly of the NMV, are thanked for devising and formulating the original project. Dr Smith was Principal Investigator in the MST Project supporting Felix Wiedenmayer's tenure. Dr Avril Ayling (Sea Research, Daintree, Queensland) made available her manuscript checklist for mainland Australia and alphabetical file of species, which proved to be very helpful in all phases of compilation. During the period 1981–1984 both Avril and John Hooper assisted with many of the initial taxonomic decisions.

Dr Barry Richardson and Dr Dan W. Walton, formerly of the Australian Biological Resources Study, are thanked for their encouragement and guidance with adaptation of the original Checklist of Australian Sponges for publication as a Catalogue in the present series. Dr Carden Wallace and Dr W.W. Keith Houston undertook initial editing of the Catalogue; they were succeeded by Dr Alice Wells. Frank Coffa of the Museum of Victoria is thanked for processing the many photographs of type specimens, now in NMV Sponge Archives. The staff of the NMV Library, particularly Lily Stefani, were instrumental with the many interlibrary loans. Judith Odgers, Biomedical Library, Monash University, provided useful references and abstracts of recent papers (1970–1982) obtained through a computer search.

This volume has been some nine years in preparation during which time considerable attention has been given to both lower and higher systematics of sponges. John Hooper added to the volume information from all pertinent publications on the Australasian sponges between 1985 and December 1993, revising all the taxa in light of recent revisions, and rewriting most of the definitions and other information given here in accordance with more recent publications and contemporary interpretations of sponges and their characters. For their assistance in ensuring that contemporary taxonomic arrangements were included in the final version of the Catalogue, and for other useful comments on the manuscript, we thank Dr Nicole Boury-Esnault (Marseille), Dr Henry Reiswig (Montreal), Dr Jean Vacelet (Marseille), Prof. Patricia Bergquist (Aukland) and Dr Rob Van Soest (Amsterdam).

John Hooper acknowledges a Post-Doctoral Fellowship (1988) from Professor Claude Lévi (Laboratoire de Biologie des Invertébrés Marins et Malacologie, Muséum National d'Histoire Naturelle), a Churchill Fellowship (1988) from the Sir Winston Churchill Memorial Trust (Darwin and Canberra), an Australian Biological Resources Study grant (86/0724) which enabled detailed studies to be carried out on collections at the Paris (MNHN) and London (BMNH) Museums and additional support from ABRS for updating of the data files. John acknowledges both his previous (NTM, Darwin) and present employers (QM, Brisbane) for supporting his participation in the Catalogue. Technical assistance by Ms A.M. Mussig, Ms J. Baxter, Ms C. Johnston, Mr R. Williams, Mr D. Low Choy, Mr A. Howard and Mr S. Cook is gratefully acknowledged. Ms Leonie Hooper prepared the illustrations.

Surveys by both authors, of types and other collections from museums throughout the world, would not have been possible without the valuable assistance of numerous people: Dr Lester Cannon (QM, Brisbane), Dr J.W. Pickett (NSW Geological Survey), Dr Frank W.E. Rowe (formerly AM, Sydney), Ms Shirley Stone and Dr David George (BMNH, London), Dr and Mrs Dieter Kühlmann (ZMB, East Berlin), Dr Ruth Desqueyroux-Faundez (MNHG, Geneva), Dr D. von Knorre (PMJ, Jena), Prof. Claude Lévi (MNHN, Paris), Dr J.A. Devidts and G. Hildwein (MZUS, Strasburg), Dr E. Kreissl (MJG, Graz), Dr M. Dzwillo (ZMH, Hamburg), Dr P. Kuenzer (Göttingen), Dr I. Hasenfurs (Erlangen), Dr Manfred Grasshoff (SMF, Frankfurt), Ms B.L. Burch (BPBM, Honolulu), Ms Susan Chambers (RSM, Edinburgh), Dr Frank Climo (NMNZ, Wellington), Dr J.C. Den Hartog (RMNH, Leiden), Dr T. Hoshino (deceased; MMBS, Hiroshima), Dr C.C. Lu and Ms Susan Boyd (NMV, Melbourne), Dr A.K. Mandal (IM, Calcutta), Mrs Loisette Marsh (WAM, Perth), Dr C. O'Riordan (INM, Dublin), Dr Urs Rahm (NM, Basel), Dr Klaus Rützler and Ms Kate Smith (USNM, Washington), Dr Rob Van Soest (ITZ, Univ. Amsterdam), Dr B.R. Stuckenburg (UM, Natal), Dr Ole Tendal (UZM, Copenhagen), Dr Jean Vacelet and Dr Nicole Boury-Esnault (SME, Marseille), Mr Shane Parker (deceased) and Mr Wolfgang Zeidler (SAMA, Adelaide).

To all persons and institutions mentioned, and to any we may have overlooked, we express our deep gratitude.

Database Notes

The Map and illustrations cited in the text refer to figures provided in the published Catalogue (Hooper & Wiedenmayer 1994). They will be included in this web site in due course.

Type Specimens
A legacy of the early work and practices in sponge taxonomy is the difficulty of locating types. Our surveys of the Australian Museum (AM) collections revealed that several type specimens (mainly of Lendenfeld, Whitelegge and Hallmann) are still missing from the type collection, which was only separated from the general collection within the past 20 years. These may still be amongst the general collection, but casual searches have been unsuccessful. Likewise, a number of type specimens purportedly in the BMNH (mostly those of Lendenfeld 1889), could not be found. These are part of the old (pre-1880s) dry collection and will probably be found eventually in the presently unsorted 'exotic' collection.

Where type material is not available, and where corresponding published descriptions are not informative, some of our placements of these taxa are speculative and are noted.

Prior to Burton's (1929, 1934) work on the Australian fauna, it was not usual practice for taxonomists to designate holotypes of new species in their publications, or to detail designations with reference to a figure, to dimensions or the salient traits, to a register number, or even to a single specimen of the species. A few papers by Carter, Ridley, the Challenger monographs, Kirkpatrick (1908), and Dendy (1922, 1924) are exceptions, but even in these the authors are inconsistent. Likewise, it was not common practice to designate lectotypes. Topsent (1930, 1932, 1933) implied such designations consistently in his revisions of Lamarck's sponges by the statement 'le type'. In his list of specimens in the BMNH, Burton (1963: 577) was still ambiguous in his type material designation, usually only noting indiscriminantly that specimens were `cotypes', meaning paratypes, paralectotypes, and syntypes. Similarly, the indication 'holotype' in Bergquist (1980b), in both her figure captions and in text, mostly implies a lectotype designation (given that most of Lendenfeld's type specimens consisted of syntypes). Our recent surveys of the AM and BMNH type collections, particularly of Lendenfeld's material, resulted in many more lecto- and paralectotype specimen designations (Wiedenmayer 1989; Hooper 1991). Most of the remaining 'undesignated material', where known, is nominated in the present volume.

It should be noted that Whitelegge and Hallmann, in their revisions of Lendenfeld species, rarely, if ever, referred to a ‘type’ in a type series (i.e. implying a lectotype), although distinctions between ‘type’ and ‘cotype’ are often found on their labels. Similarly, Lendenfeld only occasionally distinguished ‘type’ and ‘duplicate’ on labels of his syntypes, and similar statements are found on specimen labels and in register entries in the BMNH, often made by subsequent curators. Although such statements have no status as subsequent designations under the ICZN, we have often followed their actions in our designation of lectotypes in the present work (in the absence of any overriding considerations, such as published figures or condition of material or misidentifications of specimens and types by these authors).

In many cases, subsequent curators or other taxonomists (especially Ridley, Kirkpatrick, Dendy and Burton at the BMNH, and Whitelegge and Hallmann at the AM) wrote notes about a specimen’s identity or type status directly on the museum specimen labels, and/or registers. These annotations are recorded in NMV Sponge Archive 10/1–3. In a few cases, where we have been uncertain of the higher taxonomic placement of particular specimens, we have used these annotations to assist in our taxonomic decisions, and these instances are acknowledged here. In situations where more than one primary type specimen exists (syntypes), we have considered that the comment ‘type’ written on a museum label or register pertaining to a Carter or Lendenfeld specimen, is good indication that the responsible curator regarded that specimen as being most representative, agreeing best with the original description, or that he had access to some overlooked or now lost manuscript indication to that effect by the original author. However, having remained unpublished, such a remark cannot be construed as referring to the holotype or as a subsequent designation of a lectotype. Other specimens of the same type series may or may not be designated as ‘cotypes’. A remark ‘type?’ on the register, therefore, does not necessarily mean that the particular specimen has been excluded from the type series by the responsible curator, but merely is an annotation of his uncertainty as to which is the ‘type’ in his or the author’s unstated sense.

Collation of particular slides with specimens can be difficult or doubtful, especially with type series, or where a species has been redescribed by the author from new records. Many slides in the BMNH have been made subsequently from specimens housed there, or from fragments of types received in exchange from abroad, or were received as such through donations, purchase or exchange. A number of such slides have probably been missed in this work. Unfortunately, earlier curators at both the BMNH and AM often registered microscope slide preparations of type material with new numbers, different from the type itself. Interpretation of the status of a slide by this means was found to be extremely unreliable, particularly where corresponding register entries are not explicit.

In retrospect, most problems in preparing the Catalogue, and shortcomings of the final product, are ultimately related to finding and interpreting the status of scattered type material, and the authors’ lack of expertise in particular groups. (Since the greater part of the known Australian sponge fauna was described prior to 1900, relatively few taxa are now considered to be well known by contemporary authors.) Where discrepancies and doubts remain, they are indicated.

Extralimital Distributions
For extralimital species distributions, the selection of reference sources used was made on biogeographic criteria. Thus, the northern, eastern and southern Atlantic and the northern Pacific regions are neglected completely. One general guideline for the selection of extralimital titles is that the zoogeographical affinity of the sponge fauna of mainland Australia is assumed to reflect a disjunct circumtropical and warm-temperate distribution with Tethyan palaeogeographic roots (Wiedenmayer 1977). This assumption is probably incorrect, as suggested by Hooper & Lévi (1994) in their analysis of three families of demosponges in the Australian and New Caledonian regions, but sponge zoogeography is still largely in the realms of speculation rather than science. Our present interpretation, and our restriction of the extra-limital fauna, is a compromise with regard to amount and detail of data to be included, particularly for the more important systematic-descriptive accounts. The world marine zones referred to in the distribution records in this work are illustrated in Fig. 2.

 Fig. 2. Major world biogeographic provinces related to sponge distributions

Fig. 2. Major world biogeographic provinces related to sponge distributions (Australian provinces after Wilson & Allen (1987) and Hooper (1991); other provinces after Wiedenmayer (1989) and Hooper and Lévi (1994)); 1 Solanderian; 2, Peronian; 3, Maugean; 4, Subantarctic; 5, Flindersian; 6, Damperian; 7, New Zealand; 8, New Caledonian; 9, SW central Pacific; 10, NW central Pacific; 11, NW Pacific rim; 12, E Indonesia; 13, Philippines; 14, S China Sea; 15, NE Indian Ocean; 16, Arabian Gulf — Red Sea; 17, W Indian Ocean; 18, S and SE Africa; 19, SE Atlantic; 20, tropical NE Africa; 21, Mediterranean; 22, boreal NE Atlantic; 23, NW Atlantic; 24, Caribbean; 25, tropical SW Atlantic; 26, temperate SW Atlantic; 27, Austral; 28, SE Pacific rim; 29, NE Pacific rim; 30, Boreal. (Mercator cylindrical projection.) [Leonie G. Hooper]

The inclusion of the Australian Antarctic and Subantarctic fauna made it desirable to include references on the Arctic and boreal sponge fauna, because of the supposed (but probably completely incorrect) bipolar distribution of many elements (Koltun 1970). Fortunately, the extensive bibliographic coverage in Lundbeck (1902–1910), Hentschel (1929) and especially Koltun (1959–1966) restricted the choice of titles for these areas.

Another consideration in this choice was that the more comprehensive monographs, with ample synonymies, discussions and good data on distribution, largely compensate for the omission of small and strictly local papers appearing before 1984. However, a relatively larger number of descriptive papers on restricted sponge faunas within Australasia have been published since 1984.

Reference Citation
Beyond systematic descriptive accounts, significant works on biology (reproduction, ontogeny, physiology, cytology, histology, dissociation-reaggregation, microbiology-symbiosis, biochemistry, molecular biology, genetics, chemotaxonomy), ecology and zoogeography are cited. As yet, little research of this type has been carried out in Australia and so much obviously remains to be done. Again, a compromise had to be reached in this volume and extralimital works in these fields are restricted to the period 1970–1991. Such references were drawn from the Zoological Record and from our personal reprint exchange collections and, although our selection is far from complete, it is nevertheless a lead into the various fields. Biochemical and marine natural products literature dealing with sponges was only considered for those works which utilized taxonomic expertise to identify material, following the recommendations of Bergquist & Wells (1983).

Many dates of publications, particularly of the works of the earlier authors and the more obscure publications, vary in their citation between different contemporary authors, and some are still disputed. We follow Vosmaer (1928) in this work, being the most thorough, complete, and comprehensive bibliographical work on sponges for all literature prior to 1913. Publications subsequent to 1913 follow the Zoological Record.


Between publication of the Porifera section of the Zoological Catalogue of Australia database on 21 November 1994 ((Hooper, J.N.A. & Wiedenmayer, F., 1994)) and 1999 a substantial amount of taxonomic revision and nearly 200 additional species were described for the Australasian fauna. J.N.A. Hooper incorporated these revisions and new species into the first web-based version of the Catalogue database, including publications up to and including 30 March 1999, based on the original published work.


In 2002 a major systematic revision of Porifera was published Systema Porifera. A guide to the classification of sponges, as well as many new species and higher taxa added to the Australian marine sponge inventory during this period. Consequently, J.N.A. Hooper restructured the entire existing Catalogue database according to the revised Systema classification, including diagnoses of higher taxa, some nomenclatural corrections and taxonomic mistakes overlooked during the previous editions of the Catalogue.

In addition to the new taxonomic and nomenclatural decisions pubished in the Systema Porifera, there are also overviews of the recent advances in several biological aspects of Porifera, including a summary of the debate concerning the monophyly of Porifera itself. The reader is referred to that publication, which will eventually appear as an interactive web publication.

Table 2: The revised higher level classification of families (in alphabetical order) recorded in the Australian fauna. (After recent emendments published in Hooper & Van Soest (2002))


Plakinidae Schulze, 1880

Agelasidae Hartman, 1980
Astroscleridae Lister, 1900

Darwinellidae Merejkowsky, 1879
Dictyodendrillidae Bergquist, 1980

Dysideidae Gray, 1867
Irciniidae Gray, 1867
Spongiidae Gray, 1867
Thorectidae Bergquist, 1978

Axinellidae Carter, 1875
Bubaridae Topsent, 1894
Desmoxyidae Hallmann, 1917
Dictyonellidae Van Soest, Diaz & Pomponi, 1990
Halichondriidae Gray, 1867

Halisarcidae Schmidt, 1862

Callyspongiidae de Laubenfels, 1936
Chalinidae Gray, 1867
Niphatidae Van Soest, 1980
Petrosiidae Van Soest, 1980
Phloeodictyidae Carter, 1882
Metaniidae Volkmer-Ribeiro, 1986
Spongillidae Gray, 1867

Latrunculiidae Topsent, 1922
Acarnidae Dendy, 1922
Microcionidae Carter, 1875
Raspailiidae Hentschel, 1923
Rhabderemiidae Topsent, 1928
Cladorhizidae de Laubenfels, 1936
Desmacellidae Ridley & Dendy, 1886
Esperiopsidae Hentschel, 1923
Guitarridae Dendy, 1924
Isodictyidae Dendy, 1924
Mycalidae Lundbeck, 1905
Podospongiidae de Laubenfels, 1936
Chondropsidae Carter, 1886
Coelosphaeridae Dendy, 1922
Crambeidae Lévi, 1963
Crellidae Dendy, 1922
Dendoricellidae Hentschel, 1923
Desmacididae Schmidt, 1870
Hymedesmiidae Topsent, 1928
Iotrochotidae Dendy, 1922
Myxillidae Dendy, 1922
Phellodermidae Van Soest & Hajdu, 2002
Tedaniidae Ridley & Dendy, 1886

Aplysinellidae Bergquist, 1980
Aplysinidae Carter, 1875
Ianthellidae Hyatt, 1875
Pseudoceratinidae Carter, 1885

Verticillitidae Steinmann, 1882

Ancorinidae Schmidt, 1870
Geodiidae Gray, 1867
Pachastrellidae Carter, 1875

Chondrillidae Gray, 1872

Acanthochaetetidae Fischer, 1970
Alectonidae Rosell, 1996
Clionaidae D'Orbigny, 1851
Hemiasterellidae Lendenfeld, 1889
Placospongiidae Gray, 1867
Polymastiidae Gray, 1867
Sollasellidae Lendenfeld, 1887
Spirastrellidae Ridley & Dendy, 1886
Stylocordylidae Topsent, 1928
Suberitidae Schmidt, 1870
Tethyidae Gray, 1867
Timeidae Topsent, 1928
Trachycladidae Hallmann, 1917

Desmanthidae Topsent, 1894
Theonellidae Lendenfeld, 1903

Samidae Sollas, 1888
Tetillidae Sollas, 1886


Baeriidae Borojevic, Boury-Esnault & Vacelet, 2000

Clathrinidae Minchin, 1900
Leucaltidae Dendy & Row, 1913
Leucascidae Dendy, 1892
Leucettidae de Laubenfels, 1936
Levinellidae Borojevic & Boury-Esnault, 1986
Soleniscidae Borojevic, Boury-Esnault & Vacelet, 1990

Achramorphidae Borojevic, Boury-Esnault, Manuel & Vacelet, 2002
Amphoriscidae Dendy, 1892
Grantiidae Dendy, 1892
Heteropiidae Dendy, 1892
Jenkinidae Borojevic, Boury-Esnault & Vacelet, 2000
Lelapiidae Dendy & Row, 1913
Leucosoleniidae Minchin, 1900
Sycettidae Dendy, 1892

Minchinellidae Dendy & Row, 1913


Hyalonematidae Gray, 1857
Monorhaphididae Ijima, 1927
Pheronematidae Gray, 1870

Aulocalycidae Ijima, 1927
Euretidae Zittel, 1877
Farreidae Gray, 1872
Aulocystidae Sollas, 1887
Euplectellidae Gray, 1867
Rossellidae Gray, 1872

Limital Area

Distribution data in the Directory is by political and geographic region descriptors and serves as a guide to the distribution of a taxon. For details of a taxon's distribution, the reader should consult the cited references (if any) at genus and species levels.

Australia is defined as including Lord Howe Is., Norfolk Is., Cocos (Keeling) Ils, Christmas Is., Ashmore and Cartier Ils, Macquarie Is., Australian Antarctic Territory, Heard and McDonald Ils, and the waters associated with these land areas of Australian political responsibility. Political areas include the adjacent waters.

Terrestrial geographical terms are based on the drainage systems of continental Australia, while marine terms are self explanatory except as follows: the boundary between the coastal and oceanic zones is the 200 m contour; the Arafura Sea extends from Cape York to 124 DEG E; and the boundary between the Tasman and Coral Seas is considered to be the latitude of Fraser Island, also regarded as the southern terminus of the Great Barrier Reef.

Distribution records, if any, outside of these areas are listed as extralimital. The distribution descriptors for each species are collated to genus level. Users are advised that extralimital distribution for some taxa may not be complete.



Sessile metazoans possessing inhalant and exhalant pores connected by chambers lined by choanocytes. The outside and the canals are lined by pinacocytes (exo- and endopinacocytes respectively). Water is inhaled through small pores (10-100µm in diameter), traversing the afferent canals towards the choanocyte chamber, and is expelled through efferent canals and the larger exhalant osculum. Water currents are unidirectional, maintained by an active beating of a single layer of flagellated cells (choanocytes) usually contained within chambers. Food particles and oxygen are removed from the water by various cells, including the choanocytes. Other cells, including archaeocytes, are instrumental in transporting these respiratory and dietary products throughout the sponge body, in addition to other functions. Cells are highly mobile, such as those that move freely within an extracellular matrix made of fibrils of collagen (the mesohyl): archaeocytes, collenocytes, spiculocytes, spongocytes, glycocytes, cells with inclusions, etc. (see definitions of terms in Boury-Esnault & Rützler, 1997). Most of the cells, especially the archaeocytes, have an ability to continually evolve into several other cell types as required by the individual organism (totipotency), which provide the sponge with a plasticity for its organisation. Firmness of the sponge body is provided by (1) collagen fibrils of the mesohyl, (2) spongin fibres, and (3) an inorganic skeleton consisting of various supporting mineral elements composed of either calcareous (CaCO3) or silica (SiO2) (including discrete spicules, articulated or fused spicules and/or hypercalcified mineralised basal skeleton). Articulated and hypercalcified skeletons are absent in most Recent taxa, but were much more prevalent in fossil faunas ('lithistid' and 'sclerosponge' bauplans). These organic and inorganic materials are manufactured or otherwise engineered by various types of cells. Sponges have free-swimming or creeping larvae, although most groups have considerable means of asexual propagation, and all have extensive regenerative powers that appear to be vital for sustaining local populations. There are three distinct classes (Hexactinellida, Demospongiae and Calcarea), with the extinct class Archaeocyatha having suspected affinities with Demospongiae. The fossil 'classes' Sphinctozoa and Stromatoporoidea are obvious grades of construction and not phylogenetic clades (e.g., Wood 1991a), although distribution of the many lower taxa amongst the four well-established classes remains largely unresolved.


ID Keys

(1) Mineral skeleton absent----------------------------------------------------------------------Demospongiae
Mineral skeleton includes a ‘hypercalcified’ basal skeleton of solid limestone--------------------------2
Mineral skeleton consisting of discrete spicules--------------------------------------------------------------------3

(2) Soft parts contain siliceous spicules-----------------------------------------------------Demospongiae
Soft parts contain calcareous spicules (test with acid)------------------------------------------Calcarea

(3) Spicules siliceous, the larger ones are triaxone/hexactine (six-rayed), occurring both individually and
fused together-------------------------------------------------------------------------------------Hexactinellida
Spicules siliceous, the larger ones are tetraxone or monaxone----------------------Demospongiae
Spicules calcareous (test with acid), usually triactine or tetractine--------------------------Calcarea


General References

Ayling, A.L. 1982. A redescription of Astrosclera willeyana Lister, 1900 (Ceratoporellida, Demospongiae), a new record from the Great Barrier Reef. Memoirs of the National Museum of Victoria, Melbourne 43: 99-103

Ayling, A.L., Stone, S. & Smith, B.J. 1982. Catalogue of types of sponge species from Southern Australia described by Arthur Dendy. Reports of the National Museum of Victoria 1: 97-109

Bergquist, P.R. 1961a. A collection of Porifera from northern New Zealand, with descriptions of seventeen new species. Pacific Science 15: 33-48 18 figs

Bergquist, P.R. 1961b. Demospongiae (Porifera) of the Chatham Islands and Chatham Rise, collected by the Chatham Islands 1954 Expedition. Bulletin of the New Zealand Department of Scientific and Industrial Research 139: 169-206 20 figs

Bergquist, P.R. 1961c. The Keratosa (Porifera) collected by the Chatham Islands 1954 Expedition. Bulletin of the New Zealand Department of Scientific and Industrial Research 139: 207-219 6 figs

Bergquist, P.R. 1965. The sponges of Micronesia, Part I. The Palau Archipelago. Pacific Science 19: 123-204 34 figs

Bergquist, P.R. 1968. The marine fauna of New Zealand: Porifera, Demospongiae, Part 1 (Tetractinomorpha and Lithistida). Bulletin of the New Zealand Department of Scientific and Industrial Research 188: 1-105 15 pls 30 figs

Bergquist, P.R. 1969. Shallow water Demospongiae from Heron Island. Papers from the Heron Island Research Station, University of Queensland 1(4): 63-72 2 pls

Bergquist, P.R. 1970. The marine fauna of New Zealand: Porifera, Demospongiae, Part 2 (Axinellida and Halichondrida). (Mem. N.Z. Oceanogr. Inst. 51). Bulletin of the New Zealand Department of Scientific and Industrial Research 197: 1-85 20 pls 3 figs

Bergquist, P.R. 1972. Deep water Demospongiae from New Zealand. Micronesica 8: 125-136

Bergquist, P.R. 1978. Sponges. London : Hutchinson 268 pp. 12 pls 81 figs 15 tables.

Bergquist, P.R. 1980a. The ordinal and subclass classification of the Demospongiae (Porifera), appraisal of the present arrangement, and proposal of a new order. New Zealand Journal of Zoology 7: 1-6

Bergquist, P.R. 1980b. A revision of the supraspecific classification of the orders Dictyoceratida, Dendroceratida and Verongida (Class Demospongiae). New Zealand Journal of Zoology 7: 443-503 figs 1-25 pls

Bergquist, P.R. 1995. Dictyoceratida, Dendroceratida and Verongida from the New Caledonia Lagoon (Porifera: Demospongiae). Memoirs of the Queensland Museum 38(1): 1-51

Bergquist, P.R. 1996. The marine fauna of New Zealand: Porifera: Demospongiae: Part 5. Dendroceratida and Halisarcida. New Zealand Oceanographic Institute Memoir 107: 1-53

Bergquist, P.R., Ayling, A.M. & Wilkinson, C.R. 1988. Foliose Dictyoceratida of the Australian Great Barrier Reef. 1. Taxonomy and Phylogenetic relationships. Pubblicazioni della Stazione Zoologica di Napoli I: Marine Ecology 9(4): 291-320

Bergquist, P.R., Hofheinz, W. & Oesterhelt, G. 1980. Sterol composition and the classification of the Demospongiae. Biochemical Systematics and Ecology 8: 423-435 7 tables

Bergquist, P.R., Morton, J.E. & Tizard, C.A. 1971. Some Demospongiae from the Solomon Islands with descriptive notes on the major sponge habitats. Micronesica 7: 99-121 pls 1-4 3 figs

Bergquist, P.R., Sinclair, M.E. & Hogg, J.J. 1970. Adaptation to intertidal existence: reproductive cycles and larval behaviour in Demospongiae. pp. 247-271 3 figs 2 tables in Fry, W.G. (ed.). The Biology of the Porifera. London : Academic Press Symp. Zool. Soc. Lond. Vol. 25 540 pp.

Bergquist, P.R. & Fromont, J. 1988. The marine fauna of New Zealand: Porifera, Demospongiae, Part 4 (Poecilosclerida). New Zealand Oceanographic Institute Memoir 96: 1-197 pls 1-57

Bergquist, P.R. & Hartman, W.D. 1969. Free amino acid patterns and the classification of the Demospongiae. Marine Biology, Berlin 3: 247-268

Bergquist, P.R. & Kelly-Borges, M. 1991. An evaluation of the genus Tethya (Porifera: Demospongiae: Hadromerida) with descriptions of new species from the southwest Pacific. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 8(1): 37-72

Bergquist, P.R. & Skinner, I.G. 1982. Chapter 3. Sponges (Phylum Porifera). 38-72 pls 1-7 in Shepherd, S.A. & Thomas, I.M. (eds). Marine Invertebrates of Southern Australia. Adelaide : D.J. Woolman.

Bergquist, P.R. & Tizard, C.A. 1967. Australian intertidal sponges from the Darwin area. Micronesica 3: 175-202 6 pls

Bergquist, P.R. & Warne, K.P. 1980. The marine fauna of New Zealand: Porifera, Demospongiae, Part 3 (Haplosclerida and Nepheliospongida). Memoirs of the New Zealand Oceanographic Institute 87: 1-77 17 pls 4 figs

Bergquist, P.R. & Wells, R.J. 1983. Chemotaxonomy of the Porifera: the development and current status of the field. pp. 1-50 in Scheuer, P. (ed.). Marine Natural Products. New York : Academic Press.

Borojevic, R. 1966. Eponges calcaires des côtes de France. I. Amphiute paulini Hanitsch; les genres Amphiute et Paraheteropia. Archives de Zoologie Expérimentale et Générale 106: 665-670 2 figs

Borojevic, R. 1967a. Spongiaires d'Afrique du Sud, (2) Calcarea. Transactions of the Royal Society of South Africa 37: 183-226 pl. 16 25 figs

Borojevic, R. 1967b. Eponges calcaires des côtes de France. II. Le genre Ascandra Haeckel emend. Archives de Zoologie Expérimentale et Générale 107: 357-368 5 figs

Borojevic, R. 1967c. Eponges calcaires des côtes de France. III. Discussion sur la taxonomie des Eponges calcaires: Aphroceras ensata (Bowerbank) et Ute gladiata sp. n. Archives de Zoologie Expérimentale et Générale 107: 703-724 6 figs

Borojevic, R. 1967d. Eponges calcaires recueillies en Nouvelle-Calédonie par la Mission Singer-Polignac In, Expéd. Franç. Récifs Coral. Nouv. Caléd Paris : Fond. Singer-Polignac. Vol. 2. 11 pp. 4 figs

Borojevic, R. 1967e. Importance de l'étude de la répartition écologique pour la taxonomie des Eponges calcaires. Helgoländer Wissenschaftliche Meeresuntersuchungen 15: 116-119

Borojevic, R. 1968a. Eponges calcaires des côtes de France. IV. Le genre Ascaltis Haeckel emend. Archives de Zoologie Expérimentale et Générale 109: 193-210

Borojevic, R. 1968b. Systématique et évolution des éponges Calcaires. Dieuxième Thèse Université de Paris. 33 pp.

Borojevic, R. 1971a. Eponges calcaires. pp. 113-127 in Grua, P. (ed.). Comité National Français des Recherches Antarctiques (CNFRA), No. 3. Territoires des terres australes et antarctiques françaises. Invertébrés de l'infralittoral rocheux dans l'archipel de Kerguelen. Paris : CNFRA.

Borojevic, R. 1971b. Eponges calcaires des côtes sud-est du Brésil, épibiontes sur Laminaria brasiliensis et Sargassum cymosum. Revista Brasileira de Biologia 31: 525-530

Borojevic, R., Boury-Esnault, N. & Vacelet, J. 1990. A revision of the supraspecific classification of the subclass Calcinea (Porifera, Class Calcarea). Bulletin du Muséum d'Histoire Naturelle. Paris 4 12(A,2): 243-276

Borojevic, R., Cabioch, L. & Lévi, C. 1968. Inventaire de la faune marine de Roscoff. Spongiaires. Editions de la Station Biologique de Roscoff, nouvelle série. Paris : Cahiers de Biologie marine 44 pp.

Borojevic, R. & Boury-Esnault, N. 1987. Revision of the genus Leucilla Haeckel, 1872, with a re-description of the type species Leucilla amphora Haeckel, 1872. pp. 29-40 in Jones, W.C. European contributions to the taxonomy of sponges. Publications of the Sherkin Island Marine Station 1

Borojevic, R. & Grua, P. 1964. Eponges calcaires de Kerguelen. Systématique et écologie. Archives de Zoologie Expérimentale et Générale 105: 1-29 12 figs

Borojevic, R. & Peixinho, S. 1976. Eponges calcaires du nord-nord-est de Brésil. Bulletin du Muséum National d'Histoire Naturelle. Paris Zool. 279: 987-1036

Bowerbank, J.S. 1841a. [On the keratose or horny sponges of commerce]. Annals and Magazine of Natural History 7: 72-74

Bowerbank, J.S. 1841b. Observations on a keratose sponge from Australia. Annals and Magazine of Natural History 7: 129-132 pl. 3

Bowerbank, J.S. 1841c. Description of three species of sponge, containing some new forms of organization. The Microscopical Journal, London 1: 161-162 [also published in Ann. Mag. Nat. Hist. 8(1842): 393–394]

Bowerbank, J.S. 1844a. On the keratose or horny sponges of commerce. Transactions of the Microscopical Society of London 1: 32-39 pl. 3

Bowerbank, J.S. 1844b. On three species of sponge containing some new forms of organisation. Transactions of the Microscopical Society of London 1: 63-76 pl. 6 [pl. 7 omitted]

Bowerbank, J.S. 1845. Observations on the Spongiadae, with descriptions of some new genera. Annals and Magazine of Natural History 1 16: 400-410 pls 13-14

Bowerbank, J.S. 1859. On the anatomy and physiology of the Spongiadae. Part I. On the spicula. Philosophical Transactions of the Royal Society of London 148: 279-332 pls 23-26

Bowerbank, J.S. 1861. List of British sponges. In McAndrew, R., List of the British marine invertebrate fauna. Report of the British Association for the Advancement of Science (Oxford) Oxford 30: 235-236

Bowerbank, J.S. 1863a. On the anatomy and physiology of the Spongiadae. Part II. Philosophical Transactions of the Royal Society of London 152: 747-829 pls 27-35

Bowerbank, J.S. 1863b. Supplement to Part I. ‘On the anatomy and physiology of the Spongiadae’. Philosophical Transactions of the Royal Society of London 152: 830-836 pl. 36

Bowerbank, J.S. 1863c. A monograph of the Spongillidae. Proceedings of the Zoological Society of London 1863: 440-472

Bowerbank, J.S. 1864. A Monograph of the British Spongiadae. London : Ray Society Vol. 1 290 pp. 37 pls.

Bowerbank, J.S. 1866. A Monograph of the British Spongiadae. London : Ray Society Vol. 2 388 pp.

Bowerbank, J.S. 1869. On the generic name Alcyoncellum, and in reply to Dr. Gray's ‘Observations on sponges and on their arrangement and nomenclature’. Annals and Magazine of Natural History 4 3: 84-87

Bowerbank, J.S. 1872. Contributions to a general history of the Spongiadae. Part I. Proceedings of the Zoological Society of London 1872: 115-129 pls 5-6

Bowerbank, J.S. 1873a. Contributions to a general history of the Spongiadae. Part IV. Proceedings of the Zoological Society of London 1873: 3-25 pls 1-4

Bowerbank, J.S. 1873b. Report on a collection of sponges found at Ceylon by E.W.H. Holdsworth, Esq. Proceedings of the Zoological Society of London 1873: 25-32 pls 5-7

Bowerbank, J.S. 1873c. Contributions to a general history of the Spongiadae. Part V. Proceedings of the Zoological Society of London 1873: 319-333 pls 28-31

Bowerbank, J.S. 1874a. A Monograph of the British Spongiadae. London : Ray Society Vol. 3 367 pp. 92 pls.

Bowerbank, J.S. 1874b. Contributions to a general history of the Spongiadae. Part vi. Proceedings of the Zoological Society of London 1874: 298-305 pls 46-47

Bowerbank, J.S. 1875. Contributions to a general history of the Spongiadae. Part vii. Proceedings of the Zoological Society of London 1875: 281-296

Bowerbank, J.S. 1876. Contributions to a general history of the Spongiadae. Part VIII. Proceedings of the Zoological Society of London 1876: 768-775 pls 78-81

Bowerbank, J.S. 1877. Description of five new species of sponges discovered by Dr. A.B. Meyer on the Philippine Islands and New Guinea. Proceedings of the Zoological Society of London 1877: 456-464

Bowerbank, J.S. & Norman, A.M. 1882. A Monograph of the British Spongiadae. (supplementary). London : Ray Society Vol. 4 250 pp. 17 pls.

Brøndsted, H.V. 1924. Papers from Dr. Th. Mortensen's Pacific Expedition, 1914–16. Sponges from the Auckland and Campbell Islands. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjøbenhavn 75: 117-167

Brøndsted, H.V. 1927. Antarctic and subantarctic sponges collected by S. Wallin 1924. Arkiv för Zoologi 19(A,6): 1-6

Burton, M. 1927. A revision of the genera and species contained in Lendenfeld's ‘Die Chalineen des australischen Gebietes’. Annals and Magazine of Natural History 9 20: 289-296, 502-512

Burton, M. 1929. Porifera. Part 2. Antarctic sponges. In, British Antarctic (`Terra Nova') Expedition, 1910. Natural History Report. Zoology. 6(4): 393–458 pls 1–5. 393-458 pls 1-5

Burton, M. 1932. Sponges. Discovery Reports 6: 237-392 56 figs pls 48-57

Burton, M. 1934a. Sponges. Scientific Reports of the Great Barrier Reef Expedition 1928-1929 4: 513-621 pls 1-2

Burton, M. 1934b. Sponges pp. 1–58, 8 pls in Bock, S. (ed.) Further Zoological Results of the Swedish Antarctic Expedition 1901–1903 Vol. 3(2).

Burton, M. 1938. Non-calcareous sponges. Scientific Reports of the Australasian Antarctic Research Expedition 1911-1914 C 9(5): 1-22 fig. 1

Burton, M. 1963. A Revision of the Classification of the Calcareous Sponges. London : British Museum pp. 1-693 figs 1-375.

Cannon, L.R.G., Goeden, G.B. & Campbell, P. 1987. Community patterns revealed by trawling in the inter-reef regions of the Great Barrier Reef. Memoirs of the Queensland Museum 25(1): 45-70

Carter, H.J. 1881. Supplementary report on specimens dredged up from the Gulf of Manaar, together with others from the sea in the vicinity of the Basse Rocks and from Bass's Straits respectively, presented to the Liverpool Free Museum by Capt. H. Cawne Warren. Annals and Magazine of Natural History 5 7: 361-385 pl. 18

Carter, H.J. 1885a. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 15: 107-117

Carter, H.J. 1885b. Mode of circulation in the Spongida. Annals and Magazine of Natural History 5 15: 117-122 pl. 4 figs 5, 7

Carter, H.J. 1885c. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 15: 196-222

Carter, H.J. 1885d. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 15: 301-321

Carter, H.J. 1885e. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 16: 277-294

Carter, H.J. 1885f. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 16: 347-368

Carter, H.J. 1886a. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 17: 40-53, 112-127

Carter, H.J. 1886b. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 17: 431-441, 502-516

Carter, H.J. 1886c. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 18: 34-55

Carter, H.J. 1886d. Description of sponges from the neighbourhood of Port Phillip Heads, South Australia. Annals and Magazine of Natural History 5 18: 126-149

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