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Class DEMOSPONGIAE Sollas, 1885


Compiler and date details

2012 - John N.A. Hooper, Queensland Museum, Brisbane, Queensland, Australia(1994, updated 1999, 2004, 2011); Felix Wiedenmayer Naturhistorisches Museum Basel, Basel, Switzerland (1994)

Introduction

2012 - INTRODUCTION
Rather than laboriously revising the previous Introductions published in the 1994 book the Zoological Catalogue of Australia. Volume 12. Porifera (Hooper & Wiedenmayer 1994), and the revised 2004 online version of the Australian Faunal Directory, Porifera section, these are left intact here. Instead, an updated list is provided of higher taxonomic groups now recognised, reflecting still significant changes being undertaken to the systematics of Porifera over the nearly 20 years, due in particular to molecular genetics.

In January 2012 there were 7,187 valid species of Demospongiae recognised (from 14,368 nominal species) (Van Soest et al. 2011), including both marine and freshwater species. The subdivision of class Demospongiae into three subclasses (Tetractinomorpha, Ceractinomorpha and Homoscleromorpha) was abandoned, being clearly paraphyletic, and the latter elevated to a class of its own, no longer considered part of the Demospongiae (Gazave et al. 2010). Of the living fauna there are 12 Orders of Demospongiae, one incertae sedis, and one genus incertae sedis, as follows (from Hooper, Van Soest & Pisera 2011):

Class Demospongiae Sollas, 1885
Order Spirophorida Bergquist & Hogg, 1969 (3 families)
Family Tetillidae Sollas, 1886 (8 genera, 160 species)
Family Samidae Sollas, 1888 (1 genus, 1 species)
Family Spirasigmidae Hallmann, 1912 (2 genera, 2 species)
Order Astrophorida Sollas, 1888 (6 families)
Family Ancorinidae Schmidt, 1870 (15 genera, 303 species)
Family Calthropellidae Lendenfeld, 1906 (1 genus, 12 species)
Family Geodiidae Gray, 1867 (6 genera, 249 species)
Family Pachastrellidae Carter, 1875 (12 genera, 148 species)
Family Thoosidae Cockerell, 1925 (4 genera, 30 species)
Family Thrombidae Sollas, 1888 (2 genera, 7 species)
Order Hadromerida Topsent, 1894 (12 families)
Family Acanthochaetetidae Fischer, 1970 (2 genera, 5 species)
Family Clionaidae d’Orbigny, 1851 (11 genera, 194 species)
Family Hemiasterellidae Lendenfeld, 1889 (6 genera, 37 species)
Family Placospongiidae Gray, 1867 (3 genera, 10 species)
Family Polymastiidae Gray, 1867 (15 genera, 122 species)
Family Spirastrellidae Ridley & Dendy, 1886 (2 genera, 21 species)
Family Stylocordylidae Topsent, 1928 (1 genus, 8 species)
Family Suberitidae Schmidt, 1870 (11 genera, 218 species)
Family Tethyidae Gray, 1867 (14 genera, 119 species)
Family Timeidae Topsent, 1928 (1 genus, 53 species)
Family Trachycladidae Hallmann, 1917 (2 genera, 8 species)
Order Chondrosida Boury-Esnault & Lopès, 1985 (2 families)
Family Chondrillidae Gray, 1872 (4 genera, 32 species)
Family Halisarcidae Schmidt, 1862 (1 genus, 21 species)
‘Order Lithistida’ Schmidt, 1870 (polyphyletic, 13 families + 1 incertae sedis)
Family Azoricidae Sollas, 1888 (3 genera, 12 species)
Family Corallistidae Sollas, 1888 (5 genera, 33 species)
Family Desmanthidae Topsent, 1894 (4 genera, 14 species)
Family Isoraphinidae Schrammen, 1924 (1 genus, 2 species)
Family Macandrewiidae Schrammen, 1924 (1 genus, 8 species)
Family Neopeltidae Sollas, 1888 (4 genera, 15 species)
Family Phymaraphiniidae Schrammen, 1924 (3 genera, 5 species)
Family Phymatellidae Schrammen, 1910 (3 genera, 9 species)
Family Pleromidae Sollas, 1888 (2 genera, 5 species)
Family Scleritodermidae Sollas, 1888 (5 genera, 25 species)
Family Siphonidiidae Lendenfeld, 1903 (3 genera, 11 species)
Family Theonellidae Lendenfeld, 1903 (5 genera, 56 species)
Family Vetulinidae Lendenfeld, 1903 (1 genus, 1 species)
‘Lithistida’ incertae sedis (3 genera, 6 species)
Order Poecilosclerida Topsent, 1928 (4 suborders)
Suborder Microcionina Hajdu, Van Soest & Hooper, 1994 (4 families)
Family Acarnidae Dendy, 1922 (13 genera, 121 species)
Family Microcionidae Carter, 1875 (9 genera (1 incertae sedis), 493 species)
Family Raspailiidae Nardo, 1833 4 (23 genera (2 incertae sedis), 246 species)
Family Rhabderemiidae Topsent, 1928 (1 genus, 30 species)
Suborder Myxillina Hajdu, Van Soest & Hooper, 1994 (11 families)
Family Chondropsidae Carter, 1886 (5 genera, 80 species)
Family Coelosphaeridae Dendy, 1922 (8 genera, 208 species)
Family Crambeidae Lévi, 1963 (4 genera, 28 species)
Family Crellidae Dendy, 1922 (5 genera, 77 species)
Family Dendoricellidae Hentchel, 1923 (3 genera, 17 species)
Family Desmacididae Schmidt, 1870 (2 genera, 13 species)
Family Hymedesmiidae Topsent, 1928 (10 genera, 284 species)
Family Iotrochotidae Dendy, 1922 (6 genera, 32 species)
Family Myxillidae Topsent, 1928 (8 genera, 136 species)
Family Phellodermidae Van Soest & Hajdu, 2002 (2 genera, 11 species)
Family Tedaniidae Ridley & Dendy, 1886 (3 genera, 84 species)
Suborder Mycalina Hajdu, Van Soest & Hooper, 1994 (9 families)
Family Cladorhizidae Dendy, 1922 (6 genera, 113 species)
Family Desmacellidae Ridley & Dendy, 1886 (6 genera, 110 species)
Family Esperiopsidae Hentschel, 1923 (4 genera, 71 species)
Family Guitarridae Dendy, 1924 (4 genera, 25 species)
Family Hamacanthidae Gray, 1872 (2 genera, 27 species)
Family Isodictyidae Dendy, 1924 (2 genera, 44 species)
Family Merliidae Kirkpatrick, 1908 (1 genus, 4 species)
Family Mycalidae Lundbeck, 1905 (2 genera, 245 species)
Family Podospongiidae de Laubenfels, 1936 (7 genera, 31 species)
Suborder Latrunculina Kelly & Samaai, 2002 (1 family)
Family Latrunculiidae Topsent, 1922 (5 genera, 51 species)
Order Halichondrida Gray, 1867 (5 families)
Family Axinellidae Carter, 1875 (11 genera, 225 species)
Family Bubaridae Topsent, 1894 (4 genera, 27 species)
Family Dictyonellidae Van Soest, Diaz & Pomponi, 1990 (10 genera, 101 species)
Family Halichondriidae Gray, 1867 (14 genera (3 incertae sedis), 290 species)
Family Heteroxyidae Dendy, 19055 (12 genera (1 incertae sedis), 63 species)
Order Agelasida Hartman, 1980 (2 families)
Family Agelasidae Verrill, 1907 (1 genus, 35 species)
Family Astroscleridae Lister, 1900 (5 genera, 7 species)
Order Haplosclerida Topsent, 1928 (3 suborders)
Suborder Haplosclerina Topsent, 1928 (3 families)
Family Callyspongiidae de Laubenfels, 1936 (4 genera, 214 species)
Family Chalinidae Gray, 1867 (6 genera (1 incertae sedis), 478 species).
Family Niphatidae Van Soest, 1980 (9 genera, 157 species)
Suborder Petrosina Boury-Esnault & Van Beveren, 1982 (3 families)
Family Phloeodictyidae Carter, 1882 (5 genera, 122 species)
Family Petrosiidae Van Soest 1980 (4 genera, 125 species)
Family Calcifibrospongiidae Hartman, 1979 (1 genus, 1 species)
Suborder Spongillina Manconi & Pronzato, 2002 (6 families of living sponges, + 1 incertae sedis)
Family Lubomirskiidae Rezvoi, 1936 (4 genera, 11 species)
Family Malawispongiidae Manconi & Pronzato, 2002 (5 genera, 6 species)
Family Metaniidae Volkmer-Ribeiro, 1986 (5 genera, 27 species)
Family Metschnikowiidae Czerniawsky, 1880 (1 genus, 1 species)
Family Potamolepidae Brien, 1967 (6 genera, 31 species)
Family Spongillidae Gray, 1867 (24 genera, 173 species)
Spongillina incertae sedis (4 genera, 4 species)
Order Dictyoceratida Minchin, 1900 (5 families)
Family Ircinidae Gray, 1867 (3 genera, 110 species)
Family Thorectidae Bergquist, 1978 (24 genera, 167 species)
Family Spongiidae Gray, 1867 (7 genera (1 incertae sedis), 182 species).
Family Dysideidae Gray, 1867 (6 genera (1 incertae sedis), 90 species)
Family Verticillitidae Steinmann, 1882 6 (Living fauna has 1 genus, 1 species)
Order Dendroceratida Minchin, 1900 (2 families)
Family Darwinellidae Merejkowsky, 1879 (4 genera, 45 species)
Family Dictyodendrillidae Bergquist, 1980 (4 genera, 26 species)
Order Verongida Bergquist, 1978 (4 families)
Family Aplysinellidae Bergquist, 1980 (3 genera, 14 species)
Family Aplysinidae Carter, 1875 (3 genera, 54 species)
Family Ianthellidae Hyatt, 1875 (3 genera, 19 species)
Family Pseudoceratinidae Carter, 1885 (1 genus, 6 species)
Demospongiae incertae sedis (1 genus, 1 species (Myceliospongia))


2004 - INTRODUCTION
Demospongiae contains about 85% of all living sponges, with about 6000 'valid' living species of demosponges already described in the literature. There are potentially double this number of species, with the extant Poriferan fauna estimated to comprise at least 15,000 species worldwide (Hooper & Lévi, 1994; based on surveys of unpublished museum collections). Furthermore, this estimate is probably conservative as it largely neglects the grossly under-sampled and under-studied encrusting, cryptic, sciaphilic and other small taxa that pervade the many crowded marine communities, such as the coral reefs (Hooper et al. 1998). Most demosponges are marine, but several dozens of species occur in freshwater habitats all over the world (so far excluding Antarctica).

Within Demospongiae three subclasses are recognized (Homoscleromorpha, Tetractinomorpha, Ceractinomorpha), although there is an increasing number of anomalies and exceptions between otherwise closely allied family groups - based on putative morphological similarities and their differing reproductive strategies - signaling that some subclass taxa require tighter definition based on new data. Nevertheless, a rough division is possible based on the mutually exclusive presence of aster microscleres and reticulate skeletal elements. These might be equated with Tetractinomorpha and Ceractinomorpha, respectively (although the original contents of these subclasses were different, e.g., based on reproductive strategies, larval morphology). Homoscleromorpha is homogeneous, with a single order and family (Homosclerophorida, Plakinidae). Thus, although some taxa do not appear to fit this model this subclass system offers an hypothesis to evaluate the diverse demosponge orders and families.

A nominal supraordinal group, 'Keratosa', was proposed by early authors for sponges lacking a siliceous skeleton. Such a group, if it had any plausibility today, would contain the orders Dictyoceratida, Dendroceratida, Verongida and Halisarcida, but these groups in reality are quite disparate, and hence the taxon has no real basis for support (Bergquist et al. 1998).

Several species of 'living fossils' previously assigned to 'Sphinctozoa' (now included in the order Verticillitida) and 'Sclerospongiae' are also undoubted Demospongiae, possessing a viviparous reproductive strategy and producing parenchymella larvae, in Recent species at least (Vacelet 1979). These species presently sit uneasily within a homogeneous concept of Demospongiae, based on a poriferan bauplan.

Lévi (1953a, 1956a, 1957b, 1973) provides an outline and discussion of the various proposals subdividing the Demospongiae at suprafamily levels, and he was also the first to provide a comprehensive synthesis of the 'modern' sponge classification.

Database Notes

Three subclasses of demosponges with extant representatives are recognised, distributed amongst 15 orders, 88 families and ca. 1000 nominal genera (although only about 500 genera are presently considered valid). Most of these genera are marine but there are also about 40 genera confined to freshwater. Orders included at this time are: (1) Homoscleromorpha: Homosclerophorida; (2) Tetractinomorpha: Astrophorida, Chondrosida, Hadromerida, most 'lithistids' (polyphyletic), Spirophorida; (3) Ceractinomorpha: Agelasida, Dendroceratida, Dictyoceratida, Halichondrida, Halisarcida, Haplosclerida, Poecilosclerida (which includes some 'lithistids'), Verongida, and Verticillitida (the latter a fossil order to which a single Recent genus is currently assigned). Several other widely employed ordinal taxa are allocated to existing orders, following contemporary revisions of these groups, although not presently universally accepted (e.g., Axinellida (see Halichondrida), Ceratoporellida (see Agelasida), Choristida (see Astrophorida), Petrosida, also known previously as Nepheliospongida (see Haplosclerida)).

1994 - INTRODUCTION
The siliceous sponges, class Demospongiae, are those sponges with the skeleton composed of spongin fibres alone or together with siliceous spicules. Some ‘relict sclerosponge’ forms, however, have both a basal calcareous and aragonitic skeleton as well as free siliceous spicules, and the orders Dictyoceratida, Dendroceratida and Verongida (and some members of the Sub-class Homoscleromorpha) lack a mineral skeleton entirely. Collagenous filaments or fibrils (forming the ground substance of the intercellular mesohyl) are ubiquitous and spongin fibres (also composed of collagen) occur in most families; the histological organisation is always cellular (as opposed to syncytial in the Hexactinellida). Choanocytes occupy chambers that are spherical, hemispherical, elongate or branched.

Demosponges are found at tidal (0 m) to hadal (>6000 m) depths. About 5000 species have been described, and there are probably at least three times as many yet to be described. The deeper water fauna is possibly better known than that of shallow waters.

Three subclasses of Demospongiae have been distinguished on the basis of larval morphology and life cycle strategy based on the work of Lévi (1953) and subsequently developed by Bergquist (1978), Hartman (1982) and others. The basis for this system has been questioned by more recent authors (e.g. Van Soest 1991) indicating that vivipary/ovipary (as adaptive strategies, the former predominant in ephemeral habitats such as the inter-tidal) as well as blastula versus parenchymella larvae, are not confined to any one class. Nevertheless, this subclass division persists at the moment and each subclass is divided into several orders on the basis of geometry and spicule types. A fourth (polyphyletic) group, the ‘sclerosponges’ with calcified basal skeletons, has now been distributed amongst various established demosponge families.

The subclass Homoscleromorpha is characterised by amphiblastula larvae and viviparous reproduction. The skeleton consists of tetraxonid siliceous spicules (Hooper & Wiedenmayer 1994: fig. 35) and derivatives with equal rays. Megascleres and microscleres are not differentiated, although there may be size differences between types of spicule. Only one order is known. Until recently, two families were recognised, the Plakinidae and the Oscarellidae, differentiated by the presence and absence of spicules, respectively. However, recent genetic evidence (Solé-Cava et al. 1992) from a Corticium lacking spicules, suggests that this division between families is artificial. Thus only one family is recognised here.

Members of the subclass Tetractinomorpha have parenchymella larvae (one has a creeping blastula larva), and are predominantly oviparous, although in some genera young sponges are incubated by the parent and released as small adults. Megascleres are tetraxonid (Hooper & Wiedenmayer 1994: figs 36–46) and monaxonid (Hooper & Wiedenmayer 1994: figs 5–10, 12, 13, 17–21), occurring together or separately; microscleres include astrose forms and derivatives (Hooper & Wiedenmayer 1994: figs 117–131). Skeletal structure is usually radial or axially compressed.

Three orders of Tetractinomorpha are well established. These are Astrophorida (better known as Choristida), Hadromerida and Spirophorida. Spirophorida and Astrophorida have triaenes, which are not found in Hadromerida. A fourth order, Lithistida is clearly polyphyletic.

Members of the order Spirophorida are typically spherical, with tetraxonid and monaxonid megascleres (triaenes, Figs 36–43; oxeas, Fig. 5), in radiate pattern; protriaenes (Hooper & Wiedenmayer 1994: fig. 38) are most common and often protrude from the surface; monocrepidial desmas may be present (Hooper & Wiedenmayer 1994: figs 47, 52–57, 59); microscleres are contorted microspined sigmaspires (Hooper & Wiedenmayer 1994: fig. 103). Reproduction is oviparous without a larval stage, or viviparous with production of young adults within the parent. Two families are now included in this order (Tetillidae and Scleritodermatidae), both with sigmaspire microscleres, but only one is well represented in Australian waters (see Rützler’s (1987) revision).

The order Astrophorida typically has astrose microscleres (Hooper & Wiedenmayer 1994: figs 117–131) (but sometimes lost), microxeas and microrhabds (Hooper & Wiedenmayer 1994: figs 105, 106); with tetractinal megascleres, usually triaenes (Figs 35–46), calthrops (Figs 29–34), or short-shafted triaenes (Fig. 35), together with oxeas (Fig. 5); with radial skeletal architecture obvious at least at the surface. Reproduction is oviparous although gametes have been described for very few species; larval stages are not known.

Six of the seven families included in this order are recorded in the Australian fauna. No Australian representatives of the Calthropellidae Lendenfeld, 1906 have yet been described, although specimens resembling Pachastrissa have now been collected (Hooper, unpublished data). Maldonado (1993) recently revised the order.

The order Hadromerida is a relatively cohesive group of sponges, with uniform spiculation of monaxonid megascleres (monactinal or diactinal; Hooper & Wiedenmayer 1994: figs 59, 17–21); with radially arranged skeleton always obvious at the surface if not in the choanosome; spongin fibres poorly developed if present; ectosomal spicules typically smaller than choanosomal spicules, and usually standing perpendicular to the surface and protruding through the ectosome; microscleres, if present, consist of euasters, spinispiras, spirasters and derivatives (Figs 99–103). All groups are oviparous with development of a parenchymella larva (in one case a blastula larva) directly in sea-water.

Fourteen families of living sponges are included in this order. One found exclusively in fresh water, the Potamolepidae Brien, 1967, is apparently most closely related to the Suberitidae and Placospongiidae (Volkmer-Ribeiro & de Rosa-Barbosa 1979). This is the only family not represented in Australia; it is known only from certain rivers and lakes in Africa and South America (Hartman 1982).

The order Lithistida (in synonymy with Desmophorida Hentschel, Triaenosa Sollas, Tetracladina Zittel) is a problematic, polyphyletic assemblage of sponges, abundant in the period from the Cambrian to Quaternary, with many Recent relatives all of which retain articulated siliceous desma spicules (Figs 47–63), producing a rigid skeletal structure. Desmas are classified according to the number of secondarily silicified rays (crepis), from one (monocrepidial; Hooper & Wiedenmayer 1994: figs 47, 52–57, 59) to four (tetracrepidial; Hooper & Wiedenmayer 1994: figs 48–51, 58, 60–63). Many species also have secondary skeletons composed of free spicules, and it is these (if present) which indicate phylogenetic relationships of particular species. In this sense most orders of living sponges have desma-bearing representatives (?living relicts), and the possession of desmas is interpreted as being a primitive feature. Some ‘lithistids’ lack free spicules, so their relationships with other demosponge orders are more difficult to determine. The existing classification of desma-bearing sponges is based on the types of free spicules present, or if absent on desma morphology.

Three suborders, Triaenosina, Rhabdosina, and Anoplina, are recognised by Hartman (1982) with eight families, for only one of which there are published records for the Australian fauna (included here in Hadromerida). Among those not yet known from Australia are the Corallistidae Sollas, 1888 which is distributed in tropical and warm temperate waters; Pleromidae Sollas, 1888 and Neopeltidae Sollas, 1888, in tropical Atlantic waters; Cladopeltidae Sollas, 1888 also in tropical waters; Azoricidae Sollas, 1888, living in tropical or warm temperate waters; and Vetulinidae Lendenfeld, 1903, known only from the West Indies. The Desmanthidae Topsent, 1893 is another tropical or warm temperate group. One species, very similar to but possibly not con-specific with Desmanthus incrustans, is known to be particularly common in the Indo-west Pacific, including the coast of northwestern Australia, although it has not been formally recorded from this region (Hooper, unpublished data). Scleritodermatidae Sollas, 1888 is included in Spirophorida, based on the presence of sigmaspires.

For the third subclass, Ceractinomorpha, eight orders are differentiated in this work. Some authors also recognise a ninth but this is not universally accepted (Petrosida is combined here with the Haplosclerida), although there is biochemical evidence that contradicts this interpretation (e.g. Bergquist 1980a). Members have parenchymella larvae and are usually viviparous, although now several oviparous orders and families may also be included in this group. Generally the spicule skeleton is associated with a system of well-developed spongin fibres, forming hymedesmoid, plumose, plumo-reticulate, reticulate or condensed axial architecture, but three orders have lost their siliceous spicules altogether and several genera have also lost spongin fibres. Spicules are monaxonic, either monactinal (styles; Hooper & Wiedenmayer 1994: figs 17–26) or diactinal (oxeas–strongyles; Hooper & Wiedenmayer 1994: figs 5–15), but never tetractinal (although modifications to the ends of some of these spicules do occur e.g. Figs 16, 28). Microscleres are diverse (meniscoid, oxeote, toxote, spheres; Hooper & Wiedenmayer 1994: figs 66–116) but never astrose forms.

The well known fossil group, ‘Sphinctozoa’, sometimes placed in its own class (e.g. Hartman 1982), is now included in the Demospongiae Ceractinomorpha based on their larval morphology (e.g. Vacelet 1979, 1985). Sponges in this small group are found in shallow water environments associated with coral reefs. They have a solid aragonitic cortex producing a series of chambers on top of each other. Living ‘sphinctozoans’ lack free spicules. The Recent genus Vaceletia lacks spicular elements in its skeleton but has cells and larvae that resemble those of the Demospongiae. The current, popular theory suggests that the ‘Sphinctozoa’ are living relict species of the ceractinomorph Demospongiae.

The alternative view, that ‘Sphinctozoa’ are Calcarea with affinities with the Late Mesozoic genus Barroisia (with monaxonid and triradiate calcareous spicules enclosed within the walls of aragonitic chambers) is rejected by Vacelet (1985), as is a third possibility that they represent a completely independent class of sponges (Hartman 1982). Vacelet (1979, 1985) suggests that the two living species of Vaceletia are merely convergent on the Calcinea line of the Calcarea, with the alleged affinities inferred from the presence of a coeloblastula larval stage in their development being highly speculative.

Only one extant order, the Verticillitida, and one family are known. The Recent family, the Cryptocoeliidae, is characterised by a solid, cortical aragonitic skeleton consisting of a series of solitary or colonial chambers, one on top of the other. The lowest (oldest) chambers are usually partially filled with secondary secretions of aragonite, whereas the youngest chambers contain living ‘tissue’. Walls to these chambers are perforate, with perforations corresponding to the sites of inhalant pores (ostia) in the living ‘tissue’. Larger openings on the apex of chambers house the exhalant oscule. Aragonitic chambers contain a reinforcement of radially disposed pillars. Internal struts, or vesiculae (thin curved internal partitions) may be present or absent. The lining of the atrial cavity may or may not be calcified. The atrial cavity (spongocoel) may be lacking (=asiphonate, in Cryptocoelia, partly developed from the apex down, with annular constrictions produced by the bulging inner chamber walls (=retrosiphonate, in Stylothalamia), or uninterrupted, with a continuous cylindrical wall (=ambisiphonate in the fossil Vaceletia progenitor; or prosiphonate in the living Vaceletia n.sp. (Vacelet et al. 1992). The larvae are parenchymellae that develop from coeloblastulae.

‘Sphinctozoans’ were the main reef constructors in the Middle Triassic. The Cryptocoeliidae (or Verticillitidae) dates from the Middle Cambrian (see Pickett & Jell 1983) and is well represented in the geological record during the remainder of the Palaeozoic Era and the entire Mesozoic Era. The two monotypic fossil genera, Cryptocoelia Steinmann and Stylothalamia Ott, are known only from the alpine Triassic (Ott 1967; Pickett 1982). Only a single genus and two species have been recorded so far from Recent seas (Vaceletia crypta and Vaceletia n.sp. (Vacelet et al. 1992). Vaceletia is widely distributed on Indo-west Pacific reefs, occurring in shaded caves, tunnels and crevices from depths of 5–40 m (Hartman 1982). Specimens are known from several reef systems on the Great Barrier Reef (including Lizard Is.) although these have not yet been reported formally for the Australian reefs.

The order Poecilosclerida comprises more species than all other extant Porifera orders. It includes marine and some freshwater species; species with strictly siliceous spicules as well as those with both calcitic and siliceous skeletons; and species with free spicules and fused (desmoid) skeletons. The main skeleton is composed of megascleres (monactinal, diactinal or both; Hooper & Wiedenmayer 1994: figs 5–28) and spongin fibres in various stages of development; megascleres are frequently localised to distinct regions; microscleres include meniscoid forms such as chelae (unique to the order; Hooper & Wiedenmayer 1994: figs 66–78, 84–88) and sigmas (Figs 79–82), and other diverse forms (toxas, raphides, microxeas; Hooper & Wiedenmayer 1994: figs 89–91, 94, 105). Most families are viviparous, with uniformly ciliated parenchymellae with bare posterior poles. Sixteen families are included here, but three are atypical: two completely lacking chelae microscleres (Desmacellidae, Hamacanthidae); and one lacking chelae and presumed to be oviparous (Raspailiidae).

A recent proposal (Hajdu et al. 1994) to reorganise the Poecilosclerida recognises three suborders (Microcionina, Myxillina, Mycalina) based on the presence, respectively, of (a) palmate chelae, toxas and predominantly monactinal and terminally microspined ectosomal megascleres; (b) tridentate-derived chelae, no toxas, and predominantly diactinal ectosomal spicules (although aniso-terminations are widespread); and (c) sigmancistra-derived meniscoid microscleres and subtylostyles of a single smooth category only. This reorganisation accepts 19 valid families. Many of these, however, are largely incongruent with some of the existing family concepts, given the new emphasis placed on characters different from those considered to be important in the present classification (i.e. a substantial number of generic reassignments are required), and this scheme remains controversial. Nevertheless, Hajdu et als. (1994) proposal is followed here, with genera reorganised from the original published version of the ZCA Porifera volume.

Published records are available for the Australian fauna for 16 of these poecilosclerid families. Only the Hamacanthidae Gray, 1872 is not formally recorded in the Australian fauna, although several species have now been collected (Hooper, unpublished data).

The order Halichondrida is defined by the choanosomal skeleton composed of styles (Figs 17–19), oxeas (Figs 5, 8, 10, 12, 13), strongyles (Figs 6, 9) or intermediate spicules, of widely diverging sizes; spicule categories not usually functionally localised to any particular region of the skeleton although distinct skeletal structures do exist (e.g. differentiated ectosomal and choanosomal regions); skeletal structure ranging from a disorganised plumoreticulate, criss-crossed ‘halichondroid skeleton’ to one which has a distinctly compressed axis (or basal) region and a differentiated extra-axial (radial, plumose or plumoreticulate) region; spongin fibres usually poorly developed or absent. The ectosomal skeleton may be organised into a tangential layer of spicules or erect spicule bundles, with minimal collagenous spongin, typically with large subectosomal cavities; microscleres may include raphides (Figs 109–110), microxeas (Fig. 105), spined microxeas with central bend (similar to Fig. 107a) and discorhabds in a few (Figs 92, 93).

The traditional concept of Halichondrida included only two families (Hymeniacidonidae and Halichondriidae), differentiated by the presence of monactinal versus diactinal main megascleres, respectively, both with viviparous reproduction and with larvae which are completely ciliated. More recently, three (probably) oviparous families from the polyphyletic order Axinellida were also referred here (Axinellidae (including Bubaridae), and Desmoxyidae). Hymeniacidonidae was combined in Halichondriidae and a new family, Dictyonellidae, was created for species with dendritic spicule tracts (Van Soest et al. 1990). This scheme was not readily accepted by subsequent workers, not because it lacked intrinsic merit over the pre-existing system, but because there was considerable disagreement over the genera contained in the families (Axinellidae, Desmoxyidae and Dictyonellidae in particular, see Hooper & Bergquist 1992; Hooper et al. 1992). The families Axinellidae (including Bubaridae), Desmoxyidae, Dictyonellidae, and Halichondriidae (including Hymeniacidonidae) are retained here, but further revision of these groups is required, especially of their generic composition.

In the order Haplosclerida the main skeleton is partially or entirely composed of an isodictyal reticulation of spongin fibres and/or spicules, with uni- to multispicular tracts of diactinal spicules forming triangular, rectangular or polygonal meshes. Megascleres are exclusively oxeote (Figs 5, 8) or strongylote (Fig. 6), bonded together with collagenous spongin or enclosed within spongin fibres; microscleres, if present, may include sigmas (frequently centrangulate; Hooper & Wiedenmayer 1994: figs 79, 80, 83), smooth toxas (Fig. 89) or microxeas (Fig. 105).

Seven families of sponges, six with published Australian records, are included in the Haplosclerida. Five families are viviparous, with parenchymellae bearing various patterns of ciliation, one group is oviparous (Petrosiidae), and reproductive patterns of one are uncertain (Lubomirskiidae). The order includes two of the four families of ‘freshwater sponges’, Spongillidae Gray, 1867 and Lubomirskiidae Brien, 1969, with their own peculiar microsclere geometries and special spicules surrounding the gemmule (gemmoscleres, Figs 132–145). (The other ‘freshwater sponges’ are referred to Poecilosclerida (Metaniidae) and Hadromerida (Potamolepidae) following Volkmer-Ribeiro & de Rosa-Barbosa (1979) and Volkmer-Ribeiro (1986), although the alleged homologies in microscleres between these spongillids, poecilosclerids and hadromerids are still contentious). The Lubomirskiidae are known only from lakes of Eurasia.

There is disagreement as to the level of divergence between the ‘petrosiids’ and the ‘haplosclerids’, and to which families belong in each group. Bergquist (1980a) proposed that the two groups be differentiated at the ordinal level, whereby Petrosiida (as Nepheliospongiida) included Petrosia and allied genera (now in Petrosiidae) as well as Oceanapia and similar genera (now referred to Phloeodictyidae), and all other families be retained in Haplosclerida. Several recent authors (e.g. Desqueyroux-Faundez 1984, 1987; Fromont 1991, 1993) have used this arrangement, although recognising Petrosiidae and Phloeodictyidae as distinct.

Hartman (1982) modified Bergquist’s scheme to include only Petrosiidae in the Petrosiida, with all the other families in the Haplosclerida. Van Soest (1980) and others suggest that all those families belong to a single order. This latter scheme has wide usage amongst contemporary authors although it should be recognised that biochemical and morphological evidence are not entirely congruent. Fromont (1991, 1993) provides a summary of the arguments and literature of this debate. In the present work we recognise five marine families in the order Haplosclerida on the basis of recent empirical support from molecular data (Lafay et al. 1992).

Haplosclerids are probably the most abundant and diverse of all the true coral reef sponges, and are also relatively common in other habitats (patch reefs, rocky reefs, deeper water soft sediments, etc.). Family and genus level designations are relatively easily made, whereas species identifications are nearly impossible for anyone except a specialist. Recent revisions include Griessinger (1971), Bergquist & Warne (1980), Van Soest (1980), Desqueyroux-Faundez (1984, 1987) and de Weerdt (1986, 1987).

Members of the order Agelasida include both oviparous and viviparous species. Two families are includedin this order, Agelasidae and Astroscleridae, united on the basis that both have verticillate spined acanthostyles (Fig. 27; see Van Soest 1984). This affinity is now confirmed by ultrastructural studies (Boury-Esnault et al. 1990), although there is still uncertainty as to whether recognition of one or two families is appropriate. Growth forms in Agelas are branching, tubular, fan-shaped or massive, there is a well developed spongin fibre skeleton forming a regular or irregular reticulation, and fibres are echinated or cored (or both) by short styles or oxeas with verticillate spines (Fig. 27). Microscleres are absent. Hypercalcified ‘sclerosponge’ genera also included in Agelasida (Astrosclera, Ceratoporella, Goreauiella, Hispidopetra and Stromatospongia) are massive or encrusting, typically living in cryptic cavities in coral reefs. These ‘sclerosponges’ are coral-like organisms having both a fused scleritic basal skeleton composed of aragonite, and free spicules in the form of verticillate acanthostyles composed of silica (the latter may be secondarily lost in some species). Agelasids were previously assigned to the Tetractinomorpha (Bergquist 1978) and although now included in the Ceractinomorpha (Hartman 1982; Van Soest 1984; Boury-Esnault et al. 1990), their true affinities are still enigmatic.

Dictyoceratida is one of the so-called ‘keratose sponge’ orders, lacking mineral spicules, although detritus and foreign spicules may be acquired; sponges are usually tough, difficult to tear, and frequently with differences in pigmentation between the surface and subectosomal regions; the main skeleton consists of a reticulation of spongin fibres, often organised into primary, secondary and sometimes tertiary networks; fibres are usually homogeneous or lightly laminated in cross-section, with or without central pith, and collagenous spongin filaments may be scattered within the mesohyl. Larvae are large incubated parenchymellae, evenly covered with short cilia except at one pole where tufts of large flagella occur, and both poles have rings of pigmented cilia-free cells.

Sponges in the order Dictyoceratida are particularly abundant on coral reefs, and together with the Haplosclerida comprise the greater part of this fauna as presently known. Two families of dictyoderatids are now included, differentiated by their fibre characteristics; both are represented in the Australian fauna. A third family (Dysideidae) is now placed in the Dendroceratida, although this relationship is not completely straightforward. Recent revisions are given by Bergquist (1980b) and Bergquist et al. (1988).

Another ‘keratose sponge’ order, the Dendroceratida, also lacks mineral spicules. In this order the main skeleton is dendritic or reticulate, and fibres originate from a basal plate, without any obvious differences between primary and secondary spongin fibre elements; fibres are strongly laminated, with distinct pith. Larvae are incubated parenchymellae, evenly ciliated, with or without a posterior tuft of long flagella. Spongin spicules (Figs 64, 65) may be present in a few taxa.

Three families are recognised, distinguished by their respective fibre development and skeletal arrangement. All families are recorded from Australian waters. A recent revision is given by Bergquist (1980b). Boury-Esnault et al. (1990) showed that Dysideidae is heterogeneous amongst the other families of Dictyoceratida (Spongillidae, Irciniidae) with respect to choanosomal ultrastructure, showing apparently closer relationships to the Dendroceratida. Bergquist et al. (1990) similarly showed closer relationships between some members of the Dysideidae and Darwinellidae (Dendroceratida), based on terpenoid chemistry. These two studies do not completely concur on assignment of genera to Dictyoceratida and Dendroceratida, but they do show that the concept of Dysideidae is presently heterogeneous. Both studies also indicate that Halisarcidae is atypical of the Dictyoceratida; the authors retain it incertae sedis. Thus, composition of Dendroceratida and relationships of its components remain uncertain.

Sponges in the order Verongida also lack mineral spicules, and are typically fleshy and soft, with pigment that often oxidises to purple coloration; the skeleton consists of large, widely spaced spongin fibres forming dendritic or reticulate structures, and fibres may be aggregated (fasciculated) into bundles; there is no differentiation of primary and secondary elements, and detritus is only rarely incorporated into fibres; fibres have a laminated cortical (bark) region and a distinct central pith of fine spongin fibrils, but the cortex may be reduced or disappear entirely in some species. Mesohyl contains abundant collagenous fibrils.

Three families are known, all of which are thought to be oviparous and are represented in the Australian fauna. A recent revision is given by Bergquist (1980).

 

Diagnosis

Encrusting, massive, lobate, tubular, branching, flabellate, cup-shaped or excavating sponges. Skeleton composed of spongin fibres alone or together with siliceous spicules which are usually divided into megascleres and microscleres. Megascleres are basically monaxone or tetraxone; microscleres are diverse, polyaxial or monaxone, often quite elaborate in shape and ornamentation. Spongin is almost universally present, forming discrete fibres or binding the skeletal elements. In most cases the spicular skeleton and fibrous skeleton form a combined reinforcement. Fibrils of collagen are ubiquitous. Some groups lack spicular skeletons, but compensate that by building an elaborate fibre skeleton. A few (unrelated) groups have no skeletal elements other than diffuse fibrillar collagen. Other minor groups have developed a hypercalcified basal skeleton in addition to other skeletal elements, or have a solid aragonitic structure lacking free spicules. In total these skeletal variants contribute to a heterogenous morphological concept of Demospongiae, although there are other (non-morphometric) characters that provide more valuable clues as to their phylogenetic affinities (e.g., possession of viviparity, parenchymella larvae). The cellular elements are discrete, never syncytial, and cellular diversity may be considerable. The aquiferous system is of the leucon-type, although one family of deep-water Poecilosclerida (Cladorhizidae) has lost its aquiferous system and has assumed a carnivorous lifestyle, and dependent on fibroblastic flow (archaeocyte/ pinacocyte). Choanocyte chambers may be eurypylous, diplodal or aphodal. Larvae are mostly parenchymella, but in some groups there are cinctoblastula or blastula forms produced. Both oviparous and viviparous reproductive strategies occur.

 

ID Keys

KEYS TO ORDERS OF RECENT DEMOSPONGIAE

This key may not always result in the assignment of each individual sponge specimen to its proper order due to the imperfectness of juvenile or growth stages, phenomena like reduced spiculation in carbonate environments, or deviating species associated with orders only through circumstantial similarity with species showing a full complement of ordinal characters. Consequently, several redundancies are deliberately included in the key, but it is necessary to use it with care.

(1). Skeleton absent ---------------------------------------------------------------------------------------------------------------- 2
–. Skeleton present -------------------------------------------------------------------------------------------------------------------7

(2). Firm sponges with cartilaginous consistency ------------------------------------------------------------------------------- 3
–. Soft sponges --------------------------------------------------------------------------------------------------------------------- 4

(3). With leuconoid aquiferous system and diplodal choanocyte chambers; ectosome thick ------------------------------------------------------------------------ Homosclerophorida (Pseudocorticium)
–. With a well developed cortex made of thick fascicles of fibrillar collagen, numerous spherulous cells, and inhalant apertures localised in special structures -------------------------------- Chondrosida (Chondrosia)

(4). With fibrillar collagen only --------------------------------------------------------------------------------------------------- 5
–. With a nodular spongin fibre skeleton -----------------------------------------------Chondrosida (Thymosia)

(5). Choanocyte chamber eurypylous, simple ----------------------------------------------------------------------------------6
–. Choanocyte chambers tubular and branched, size about 100µm; ectosomal and subectosomal collagen highly organised and structurally diversified ----------------------------------------------------------------------------- Halisarcida

(6). Ectosome thin, with sylleibid-like aquiferous system; choanocyte chambers eurypylous, rounded, less than 60µm diameter --------------------------------------------------------- Homosclerophorida (Oscarella)
–. With a thin cortex enriched with fibrillar collagen parallel to the surface, a superficial cuticle and pore-sieves may be present --------------------------------------------------------------- Chondrosida (Thymosiopsis)
–. Ectosome strongly collagen-reinforced and bounded by a distinct skin, with spherulous cells ~10µm in diameter are common throughout the mesohyl but particularly concentrated in the ectosome; sponge attaining a thickness of only about 5mm; choanocyte chambers large and sac-shaped --------------------------------- Verongida (Hexadella)

(7)Megascleres present ---------------------------------------------------------------------------------------------------------- 8
–. Only asterose microscleres present -------------------------------------------------------- Chondrosida (Chondrilla)
–. Siliceous spicules absent (or secondarily lost) -------------------------------------------------------------------------------- 17

(8)Spicules exclusively verticillate-spined styles or oxeas --------------------------------------------------------- Agelasida
–. Spicules may be spined or smooth but are not exclusively verticillate- spined -------------------------------------------- 9
–. Megascleres always include articulated siliceous desmas, with or without free spicules (Note. Halichondrida and Poecilosclerida also contain species with 'sublithistid' desma skeletons) ------- Demospongiae 'lithistids' (polyphyletic)

(9)Megascleres are all monaxones -------------------------------------------------------------------------------------------- 10
–. Megascleres include diods and/or triods, megascleres and microscleres undifferentiated, sometimes spicules are lost completely and sponge may be superficially confused with compound ascidians --------------------- Homosclerophorida
–. Megascleres include triaenes ----------------------------------------------------------------------------------------------------16

(10)Megascleres exclusively diactines (oxeas and/or strongyles) --------------------------------------------------------- 11
–. Megascleres diverse or exclusively monactinal (tylostyles, styles, strongyloxeas) ------------------------------------- 15

(11)Asterose microscleres ------------------------------------------------------------------- Astrophorida
–. No asterose microscleres --------------------------------------------------------------------------------- 12

(12)Megascleres arranged in an isodictyal or anisodictyal reticulation ---------------------------------------------------- 13
–. Megascleres arranged in a confused manner or plumose or plumoreticulate --------------------------------------------- 14

(13)Microscleres include chelae, megascleres often localized to distinct regions (e.g., inside fibres), sand/detritus may replace megascleres completely --------------------------------------------------------------Poecilosclerida
–. No chelae; microscleres absent or restricted to sigmas, toxas, raphides, amphidiscs or microspined oxeas, megascleres diactinal usually producing well-formed structures such as triangular, rectangular or polygonal meshes ------------------------------------------------------------------------------------- Haplosclerida

(14)Microscleres include chelae and or sigmas or toxas --------------------------------------- Poecilosclerida
–. No chelae, sigmas or toxas ---------------------------------------------------------------------- Halichondrida

(15)Microscleres may be absent or may include asterose and monaxonic forms (microxeas, spirasters); skeleton peripherally radiate forming palisades of spicules at the surface --------------------------------------------- Hadromerida
–. No asters, and no other microscleres other than trichodragmas (or raphides); skeleton peripherally tangential or undifferentiated, main skeleton composed of a criss-cross of spicules, or compressed into a distinct axis, or with plumose, plumo-reticulate or dendritic mineral skeleton, fibre system poorly developed or absent ---- Halichondrida

(16)Microscleres sigmaspires (rugose c- or s-shaped), spherical growth form usual, radial pattern of triaenes and oxeas ---------------------------------------------------------- Spirophorida
–. Microscleres asters or streptoscleres, large oxeas always present, sometimes with triaenes, skeleton only obviously radial at the surface ------------------------------------------ Astrophorida

(17)Solid carbonate skeleton, lacking free spicules, with a solid cortex producing a series of chambers on top of each other, the youngest (uppermost) chambers lined with living tissue ------------------------------------- Verticillitida
of discrete spongin fibres ------------------------------------------------------------------------------------------------------ 18
–. Fibres generally well laminated, containing a cellular mass visible as a dark pith in transmitted light, without differentiation of primary or secondary elements, many taxa aerophobic (darken in contact with air) ---------------- Verongida
–. Fibres contain a core of sand or spicule fragments or are entirely free of inclusions-----------------------------------19

(19)Skeleton an anastomosing system of interconnected fibres, often well developed and relatively homogeneous fibre construction with 2-3 different sized networks, consistency not collagenous ---------------------- Dictyoceratida
–. Skeleton consists of dendritic fibres arising from basal attachment, with fibres strongly lamellated -- Dendroceratida
–. Skeleton with reticulate, plumoreticulate or plumose fibres containing sand or spicule fragments, with vestigial spicules (check for microscleres or echinating spicules) or occasionally no spicules at all --------- Poecilosclerida
–. Fibre skeleton well-developed, more-or-less regularly reticulate, and also with a tangential ectosomal (tertiary) network of fine aspicular fibres and foreign material, whereas choanosomal fibres are aspicular and with only foreign material (or sometimes extremely vestigial oxeas ----------------------------------------- Haplosclerida (Dactylia)

 

General References

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

Bergquist, P.R. 1980. 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. 1980. 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., 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., Karuso, P. & Cambie, R.C. 1990. Taxonomic relationships within the Dendroceratida: a biological and chemotaxonomic appraisal. pp. 72-78 in Rützler, K. (ed.). New Perspectives in Sponge Biology. Washington : Smithsonian Institution Press.

Bergquist, P.R., Walsh, D. & Gray, R.D. 1998. Relationships Within and Between the Orders of Demospongiae that Lack a Mineral Skeleton. In, Watanabe, Y. & Fusetani, N. (eds). Sponge Sciences: Multidisciplinary Perspectives. Tokyo, Berlin, Heidelberg, New York : Springer-Verlag 458 pp.

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

Boury-Esnault, N., de Vos, L., Donadey, C. & Vacelet, J. 1990. Ultrastructure of choanosome and sponge classification. pp. 237-244 in Rützler, K. (ed.). New Perspectives in Sponge Biology. Washington : Smithsonian Institution Press.

Desqueyroux-Faundez, R. 1984. Description de la faune des Haplosclerida (Porifera) de la Nouvelle-Calédonie I. Niphatidae-Callyspongiidae. Revue Suisse de Zoologie 91(3): 765-827

Desqueyroux-Faundez, R. 1987. Description de la faune des Petrosida (Porifera) de la Nouvelle-Calédonie. II. Petrosiidae-Oceanapiidae. Revue Suisse de Zoologie 94(1): 177-243

Fromont, J. 1991. Descriptions of species of the Petrosida (Porifera: Demospongiae) occurring in the tropical waters of the Great Barrier Reef. The Beagle, Records of the Museums and Art Galleries of the Northern Territory 8(1): 73-96

Fromont, J. 1993. Descriptions of species of the Haplosclerida (Porifera: Demospongiae) occurring in the tropical waters of the Great Barrier Reef. The Beagle, Records of the Northern Territory Museum of Arts and Sciences 10: 7-40

Gazave, E., Lapébie, P., Renard, E., Vacelet, J., Rocher, C., Ereskovsky, A.V., Lavrov, D.V. & Borchiellini, C. 2010. Molecular Phylogeny Restores the Supra-Generic Subdivision of Homoscleromorph Sponges (Porifera, Homoscleromorpha). PLoS ONE (Public Library of Science) 5(12): 1-15 [e14290. doi:10.1371/journal.pone.0014290]

Griessinger, J.M. 1971. Etude des Réniérides de Méditerranée (Démosponges, Haplosclérides). Bulletin du Muséum National d'Histoire Naturelle. Paris 3 (Zool.) 3: 97-182

Hajdu, E., Soest, R.W.M. Van & Hooper, J.N.A. 1994. Proposal of a phylogenetic subordinal classification of poecilosclerid sponges (Demospongiae, Porifera). pp. 123-139 in Soest, R.W.M. Van, Kempen, T.M.G. van & Braekman, J.-C. (eds). Sponges in Time and Space. Rotterdam : Balkema.

Hartman, W.D. 1982. Porifera. pp. 640-666 in Parker, S.P. (ed.). Synopsis and Classification of Living Organisms. New York : McGraw-Hill Vol. 1.

Hooper, J.N.A., Capon, R.J., Keenan, C.P., Parry, D.L. & Smit, N. 1992. Chemotaxonomy of marine sponges: families Microcionidae, Raspailiidae and Axinellidae, and their relationships with other families in the orders Poecilosclerida and Axinellida (Porifera: Demospongiae). Invertebrate Taxonomy 6(2): 261-301

Hooper, J.N.A., Quinn, R.J. & Murphy, P.T. 1998. Bioprospecting for marine invertebrates. In, Van Keulen, M. & Borowitzka, M.A. (eds). Biodiversity, Biotechnology & Biobusiness. 2nd Asia-Pacific Conference on Biotechnology. Perth, WA, 23-27 November 1998. Perth : Murdoch University.

Hooper, J.N.A., Van Soest, R.W.M. & Pisera, A. 2011. Phylum Porifera Grant, 1826. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness. Zootaxa 3148: 13-18 [13]

Hooper, J.N.A. & Bergquist, P.R. 1992. Cymbastela, a new genus of lamellate coral reef sponges. Memoirs of the Queensland Museum 32(1): 99-137

Hooper, J.N.A. & Lévi, C. 1994. Biogeography of Indo-west Pacific sponges: Microcionidae, Raspailiidae, Axinellidae. pp. 191-212 in Soest, R.W.M. Van, Kempen, T.M.G. van & Braekman, J.-C. (eds). Sponges in Time and Space. Rotterdam : Balkema.

Hooper, J.N.A. & Soest, R.W.M. Van 2002. Class Demospongiae Sollas, 1885. pp. 15-18 in Hooper, J.N.A. & Van Soest, R.W.M. (eds). Systema Porifera: A guide to the classification of sponges. New York : Kluwer Academic/Plenum Publishers Vol. 1.

Hooper, J.N.A. & Wiedenmayer, F. 1994. Porifera. pp. 1-620 in Wells, A. (ed.). Zoological Catalogue of Australia. Melbourne : CSIRO Australia Vol. 12 xiii 624 pp. [Date published 21/Nov/1994]

Lafay, B., Boury-Esnault, N., Vacelet, J. & Christen, R. 1992. An analysis of partial 28s ribosomal RNA sequences suggests early radiations of sponges. Biosystems 28: 139-151

Lévi, C. 1953. Sur une nouvelle classification des Démosponges. Comptes Rendus (Hebdomadaires) des Séances de l'Academie des Sciences. Série D. Sciences Naturelles 236(1): 853-855

Lévi, C. 1956. Etude des Halisarca de Roscoff. Embryologie et systématique des démosponges. Archives de Zoologie Expérimentale et Générale 93: 1-181

Lévi, C. 1957. Ontogeny and systematics in sponges. Systematic Zoology 6: 174-183

Lévi, C. 1973. Systématique de la classe des Demospongiaria (Démosponges). pp. 577-631 in Brien, P., Lévi, C., Sarà, M., Tuzet, O. & Vacelet, J. (eds). Traité de Zoologie, Anatomie, Systématique, Biologie. III. Spongiaires. (Series ed. P.-P. Grassé). Paris : Masson et Cie.

Maldonado, M. 1993. The taxonomic significance of the short-shafted mesotriaene reviewed by parsimony analysis: validation of Pachastrella ovisternata Lendf. (Demospongiae: Astrophorida). Bijdragen tot de Dierkunde 63(3): 129-148

Ott, E. 1967. Segmentierte Kalkschwämme (Sphinctozoa) aus der alpinen Mitteltrias und ihre Bedeutung als Riffbildner im Wettersteinkalk. Abhandlungen der mathematisch-physikalischen Klasse der königlich Bayerischen Akademie der Wissenschaften N.F. 131: 1-96

Pickett, J. & Jell, P.A. 1983. Middle Cambrian Sphinctozoa (Porifera) from New South Wales. Memoirs of the Association of Australasian Palaeontologists 1: 85-92

Pickett, J.W. 1982. Vaceletia progenitor, the first Tertiary sphinctozoan. Alcheringa 6: 241-247

Rützler, K. 1987. Tetillidae (Spirophorida, Porifera): a taxonomic re-evaluation. pp. 187-203 in Vacelet, J. & Boury-Esnault, N. (eds). Taxonomy of Porifera from the N.E. Atlantic and Mediterranean Sea. NATO ASI Series. Berlin : Springer-Verlag Vol. G13.

Senowbari-Daryan, B. 1990. Die systematische Stellung der thalamiden Schwämme und irhe Bedeutung in der Erdgeschichte. Münchner Geowissenschaftliche Abhandlungen 21A: 1–324.

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Soest, R.W.M. Van 1984. Deficient Merlia normani Kirkpatrick, 1908, from the Curaçao reefs, with a discussion on the phylogenetic interpretation of sclerosponges. Bijdragen tot de Dierkunde 54: 211-219

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Solé-Cava, A.M., Boury-Esnault, N., Vacelet, J. & Thorpe, J.P. 1992. Biochemical genetic divergence and systematics in sponges of the genera Corticium and Oscarella (Demospongiae: Homoscleromorpha) in the Mediterranean Sea. Marine Biology, Berlin 113: 299-304

Vacelet, J. 1979. Description et affinités d'une éponge Sphinctozoaire actuelle. (Description and affinities of a living sphinctozoid sponge). pp. 483–493 in Lévi, C. & Boury-Esnault, N. (eds) Biologie des Spongiaires/Sponge biology. Colloques Int. Cent. Natl Rech. Scient. Vol. 291.

Vacelet, J. 1985. 1. Coralline sponges and the evolution of Porifera. pp. 1-13 in Conway-Morris, S., George, J.D., Gibson, R. & Platt, H.M. (eds). The Origin and Relationships of Lower Invertebrates. Systematics Association Special Vol. 28. Oxford : Clarendon Press.

Vacelet, J., Cuif, J.-P., Gautret, P., Massot, M., Richer de Forges, B. & Zibrowius, H. 1992. Un Spongiaire Sphinctozoaire colonial apparenté aux constructeurs de récifs triasiques survivant dans le bathyal de Nouvelle Calédonie. Comptes Rendus (Hebdomadaires) des Séances de l'Academie des Sciences. Série D. Sciences Naturelles 314: 379-385

van Soest, R. 2011. Demospongiae. In: Van Soest, R.W.M, Boury-Esnault, N., Hooper, J.N.A., Rützler, K, de Voogd, N.J., Alvarez de Glasby, B., Hajdu, E., Pisera, A.B., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R., Klautau, M., Picton, B., Kelly, M., Vacelet, J. (2011) World Porifera database. Accessed through: Van Soest, R.W.M, Boury-Esnault, N., Hooper, J.N.A., Rützler, K, de Voogd, N.J., Alvarez de Glasby, B., Hajdu, E., Pisera, A.B., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick, K.R., Klautau, M., Picton, B., Kelly, M., Vacelet, J. (2011) World Porifera database at http://www.marinespecies.org/porifera/porifera.php?p=taxdetails&id=164811. http://www.marinespecies.org/porifera/porifera.php?p=taxdetails&id=164811 (checked on 2012-01-27)

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Weerdt, W.H. de 1987. The marine shallow-water Chalinidae (Haplosclerida, Porifera) of the British Isles. In Jones, W.C. (ed.) European contributions to the taxonomy of sponges. Publications of the Sherkin Island Marine Station 1: 75-108

Wilkinson, C.R. & Cheshire, A.C. 1989. Patterns in the distribution of sponge populations across the central Great Barrier Reef. Coral Reefs 8: 127-134

 

History of changes

Note that this list may be incomplete for dates prior to September 2013.
Published As part of group Action Date Action Type Compiler(s)
28-Feb-2012 28-Feb-2012 MODIFIED
12-Feb-2010 (import)