Family RASPAILIIDAE Nardo, 1833

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Family RASPAILIIDAE Nardo, 1833

Compiler and date details

2010 - John N.A. Hooper, Queensland Museum, Brisbane, Queensland, Australia; Felix Wiedenmayer (1994), Naturhistorisches Museum Basel, Basel, Switzerland; updated by John N.A. Hooper (1999)

Raspailidae Nardo, G.D. 1833. Auszug aus einem neuen System der Spongiarien, wornach bereits die Aufstellung in der Universitäts-Sammlung zu Padua gemacht ist. Isis Oken, coll. 519–523.
Sollasellidae Lendenfeld, R. von 1888. Descriptive Catalogue of the Sponges in the Australian Museum, Sydney. London : Taylor & Francis 260 pp. 12 pls.
Secondary source:
van Soest, R.W.M. 2002. Family Sollasellidae Lendenfeld, 1887. pp. 279-280 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.; van Soest, R.W.M., Hooper, J.N.A, Beglinger, E. & Erpenbeck, D. 2006. Affinities of the family Sollasellidae (Porifera, Demospongiae). I. Morphological evidence. Contributions to Zoology 75(3/4): 133-144; Erpenbeck, D., Hooper, J.N.A., List-Armitage, S.E., Degnan, B.M., Wörheide, G. & van Soest, R.W.M. 2007. Affinities of the family Sollasellidae (Porifera, Demospongiae). II. Molecular evidence. Contributions to Zoology 76(2): 95-102.
Euryponidae Topsent, E. 1928. Spongiaires de l'Atlantique et de la Méditerranée provenant des croisières du Prince Albert Ier de Monaco. Résultats des Campagnes Scientifiques accomplies par le Prince Albert I. Monaco 74: 1-376 11 pls.

Introduction

Raspailiidae Nardo, 1833 are sponges with encrusting, massive, lobate, fan-shaped or branching growth forms, usually with a very hispid surface. A specialised ectosomal skeleton is typically present, consisting of brushes of small thin styles (Hooper & Wiedenmayer 1994: fig. 17) or oxeas (Hooper & Wiedenmayer 1994: fig. 5), surrounding individual long thick styles or oxeas. The choanosomal skeleton varies from a compressed axial skeleton, to plumo-reticulate or exclusively reticulate structures. Spongin fibres usually completely enclose coring spicules (choanosomal styles, oxeas or both). A special category of spined styles (Hooper & Wiedenmayer 1994: fig. 22), or modifications to styles (e.g. Figs 22–25, 28), echinate fibres, protrude at right angles from fibres. Microscleres are usually absent, although single raphides (Hooper & Wiedenmayer 1994: fig. 109) or bundles (trichodragmata; Hooper & Wiedenmayer 1994: fig. 110) may occur in some genera. Where known, reproduction is oviparous and this condition is probably typical for this family.

Raspailiids are widely distributed, ranging from shallow waters to at least 2460 m in depth (Hartman 1982). Over 30 genera have been included in the family, but only 18 of these are presently considered to be valid, plus three incertae sedis (see Hooper 1991). Published Australian records are available for twelve genera. In addition, it is predicted that with dredging of softer substrates in deeper waters, many species of Hymeraphia, not presently recorded for the Australian region, will probably be discovered. In other parts of the world, these sponges are apparently important in binding soft sediments (Shirley Stone, pers. comm.).

Raspailiidae (including Euryponidae) and the poecilosclerid Microcionidae both have echinating acanthostyle megascleres. Furthermore, some raspailiid taxa (e.g. Raspailia, sensu stricto) show identical construction to some microcionids (e.g. Axociella, sensu stricto). It has been suggested (Hooper 1991) that a compressed axial skeleton, and differentiated axial and extra-axial skeletons (used to define 'axinellids'), have been acquired independently several times in the Porifera, de-emphasising the primary importance of this feature in systematics. This, together with chemotaxonomic evidence (Hooper et al. 1992), was used to justify the inclusion of Raspailiidae in Poecilosclerida (Hooper 1991).

The classification of the Raspailiidae centres around three principal diagnostic features: skeletal architecture (e.g. forms with axially compressed skeletons typical of 'axinellids' versus those with reticulate, plumo-reticulate or plumose skeletons), presence or absence of a specialized ectosomal skeleton (apomorphic for the raspailiids, but secondarily lost in some taxa), and modifications to echinating megascleres (with about 15 distinct morphologies, apomorphic for various genera).

The most recent revision of the family is that of Hooper (1991). Other reviews are available in Topsent (1928), Brien et al. (1973), and Bergquist (1970). Bergquist (1978) synonymised Euryponidae Topsent, 1928 with Raspailiidae. Van Soest et al. (2006) synonymised Sollassellidae Lendenfeld, 1887 with Raspailiidae.

Sollasellidae (long known only from a single species Sollasella digitata Lendenfeld, 1888), was clearly demonstrated to belong to Raspailiidae from morphological and molecular evidence (Van Soest et al. 2006; Erpenbeck et al. 2007a). Prior to that it had a multitude of ordinal and even subclass assignments. Hallmann (1914) maintained the family predominantly because of the incompatibility of S. digitata with established families. Topsent (1904) also assigned Trachya hystrixto Sollasella, but Hallmann (1914) erected for that species the genus Pseudotrachya and referred this to Axinellidae (now considered to belong to Polymastiidae). De Laubenfels (1936) assigned the family to his order Epipolasida (intended to receive families and genera with astrophorid affinities but lacking triaenes - which is no longer recognized and aster-bearing members are assigned to Ancorinidae and Hemiasterellidae. De Laubenfels (1936) assigned a number of other genera to this family, in addition to Sollasella. Van Soest (2002) rejected this proposal noting that most of these taxa belong to Halichondriidae. In the first edition of the Zoological Catalogue of Australia (Hooper & Wiedenmayer 1994), Sollasella was assigned to Coppatiidae (a dustbin family for putative Epipolasida), but these authors gave insufficient grounds for this. Van Soest (2002) maintained Sollasellidae solely for Sollasella within Hadromerida, on the basis that it shared with Polymastiidae the presence of a cortex, most similar to Pseudotrachya, with which it also shared the combination of choanosomal styles and ectosomal oxeas. It also showed similarities with some Suberitidae (Homaxinella, Plicatellopsis, Rhizaxinella) with which it shared the stalked habit and axially condensed skeleton.

That Sollasella is confirmed a member of Raspailiidae through molecular data (Erpenbeck et al. 2007a), yet shares these morphological features with some Hadromerida, is indicative of the likely incorrect assignment of Raspailiidae to the Poecilosclerida (e.g. Erpenbeck et al. 2007b).

Diagnosis

Encrusting, massive, lobate, fan-shaped or branching growth forms, usually with a very hispid surface. Specialised ectosomal skeleton typically present, consisting of small thin styles, oxeas or anisoxeas forming bouquets around long styles or oxeas that penetrate the surface. Choanosomal skeletal structure is typically reticulate with a compressed axial region and plumo-reticulate extra-axial (peripheral) skeleton, but may range from hymedesmioid and microcionid in encrusting taxa, to plumose, radial or evenly reticulate (non-compressed) skeletons in other taxa. Spongin fibres usually completely enclose coring spicules (choanosomal styles, oxeas or both). Echinating acanthostyles (or modified styles) echinate fibres, standing perpendicular to them. Microscleres are usually absent, although a few genera have raphides in bundles (trichodragmata). Where known, reproduction is oviparous and probably typical for this family.

ID Keys

Key to Subfamilies
(1) Echinating megascleres present ------------------------------------------------------------------------------------------ 2
Echinating megascleres absent ---------------------------------------------------------------- Thrinacophorinae
Echinating megascleres absent ---------------------------------- Raspailiinae (Raspailia (Parasyringella))

(2) Echinating megascleres club-shaped or rhabdose -------------------------------------------------------------------- 3
Echinating megascleres modified to sagittal acanthoplagiotriaenes -------------------------------- Cyamoninae

(3) Choanosomal skeleton cored by oxeas or styles ---------------------------------------------------------------------- 4
Choanosomal skeleton cored by acanthostrongyles or acanthotylostrongyles (‘peg-top’ spicules) -------- Plocamioninae

(4) Choanosomal skeleton with greater or lesser differentiation between axial and extra-axial regions, ectosome with specialised raspailiid skeleton -------------------------------------------------------------------- Raspailiinae
Choanosomal skeleton regularly reticulate, extra-axial skeleton vesigial or virtually absent, and all but one species lack ectosomal specialization------------------------------------------------------------------------- Echinodictyinae

Key to Genera
(1) Echinating spicules are microcionid-like, club-shaped, with rounded or sharp points, subtylote bases, and with evenly or unevenly distributed spines ------------------------------------------------------- 2
Echinating spicules are club-shaped with clavulate points; axial and extra-axial skeletons are composed of a single category of undifferentiated choanosomal megascleres ------------------------ Ectyoplasia
Echinating spicules are acanthose, club-shaped or strongylote, with strongly curved hooks on the base and shaft (cladotylote), and these spicules are usually confined to a particular region within the skeleton ------ Endectyon
Echinating spicules are acanthostyles with smooth rhabdose bases, and large recurved spines are distributed over the shaft --------------------------------------------------------------------------------- Aulospongus
Echinating spicules are smooth rhabdostyles (rare and possibly contaminants) --- Cantabrina (incertae sedis)
Echinating spicules are acanthostyles with bulbous tylote bases, with or without spines on the points and other modifications to the distal portion ------------------------------------------------------------ Hymeraphia
Echinating spicules are absent, but diactinal acanthorhabds form a compact basal mass throughout
the skeleton ------------------------------------------------------------------------------------ Rhabdeurypon
Echinating spicules are sagittal monact-, diact- or tetractinal (acanthoplagiotriaenes) with only one
spined ray--------------------------------------------------------------------------------------- Trikentrion
Echinating spicules are sagittal tetract- or pentactinal (rarely with fewer rays) (acanthoplagiotriaenes) with all or most rays spined ------------------------------------------------------------------------------ Cyamon
Echinating spicules are absent --------------------------------------------------------------------------------------- 3, 9

(2) Choanosomal skeleton consists of strongly to slightly axially compressed reticulation of spongin fibres and/or spicules ---------- Raspailia -------------------------------------------------------------------------- 3
Choanosomal skeleton microcionid, basally compressed fibres lying on the substrate, with or without ascending fibre nodes ------------------------------------------------------------------------------------------------- 7
Choanosomal skeleton reticulate without any trace of axial compression ------------------------------------ 8
Choanosomal skeleton consists of loosely aggregated or plumose axial fibres ---------- Raspaciona

(3) With echinating acanthostyles ----------------------------------------------------------------------------- 4
Without echinating acanthostyles ----------------- Raspailia (Parasyringella) and Sollassella digitata

(4) Choanosomal skeleton consists of an axially compressed reticulation of spongin fibres and/or spicule tracts, with at least some degree of differentiation between axial and extra-axial skeletons ---------------------------- 5
Choanosomal skeleton consists of a reticulation of spongin fibres and/or spicule tracts, without any trace of axial compression, and with reduced differentiation of the axial and extra-axial skeletons --------------- Raspailia (Clathriodendron)

(5) Surface with a characteristic ornamentation of polygonally arranged inhalant pores and the oxeas form a continuous palisade of brushes of oxeas pierced by long styles ------------------ Sollassella
Surface hispid, lacking continuous palisade but with long styles or oxeas surrounded by characteristic bundles of smaller oxeas .----------------------------------- 6

(6) Echinating spicules are club-shaped acanthostyles, with subtylote bases and straight shafts ----------------- Raspailia (Raspailia)
Echinating spicules are acanthose rhabdostyles ------------------------------------------ Raspailia (Raspaxilla)
Echinating spicules have smooth, swollen, tylote bases ------------------- Raspailia (Hymeraphiopsis)

(7) Spicules in the axial or basal skeleton are acanthostrongyles or acanthotylostrongyles ------------ Plocamione
No true choanosomal megascleres occur in the axial skeleton, although large extra-axial spicules and echinating acanthostyles may be embedded in spongin fibres ------------------------------------------ Eurypon

(8) Choanosomal spicules are styles or subtylostyles with spines on both basal and distal ends, forming a renieroid reticulate skeleton ----------------------------------------------------------------- Amphinomia
Choanosomal spicules are exclusively oxeas ------------------------------------------- Echinodictyum
Choanosomal spicules are acanthostrongyles or acanthotylostrongyles forming a secondary (sub)
isodictyal reticulation ------------------------------------------------------------------------ Lithoplocamia

(9) Choanosomal skeleton consists of dense axial compression of criss-crossed spicules, without
axial fibres -------------------------------------------------------------------------------- Thrinacophora
Choanosomal skeleton with a renieroid reticulate skeleton of multispicular primary fibres interconnected by uni- or paucispicular secondary fibres, forming elongate meshes, slightly more compressed at the axis than in the periphery ------------------------------------------------------------------------------------ Waltherarndtia
Choanosomal skeleton consists of axially compressed reticulation of spongin fibres and/or spicule tracts ----------- 10

(10) Spicules in the axial skeleton are sinuous styles or anisoxeas --------------------- Ceratopsion
Spicules in the axial skeleton include both spined oxeas and smooth anisoxeas ------- Axechina