Australian Biological Resources Study

Australian Faunal Directory

Calcarea: Leucosoleniidae

Calcarea: Leucosoleniidae


Regional Maps

Class CALCAREA Bowerbank, 1864

Compiler and date details

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


The Calcarea were most recently revised by Manuel et al. (2002), with recent studies on ultrastructure and molecular genetics (e.g. Manuel 2006) supporting the higher subclass classification of Calcinea and Calcaronea, but their internal relationships still largely unresolved whereby some orders and families are recognised as not monophyletic (Dohrmann et al. 2006).

Recently the name Calcispongia (common in the old literature prior to 1864) was proposed to be resurrected for calcareans to distinguish Recent representatives with calcitic spicules from the exclusively fossil Heteractinida (with polyactine spicules) (Manuel 2006, Van Soest et al. 2012), but most authors continue to follow Bowerbank's (1864) nomenclature.

In January 2012 there were 672 valid species of Calcarea recognised (from 1,259 nominal species) (Van Soest et al. 2011), exclusively marine, with the list of higher taxa and their current global diversity summarised below (from Hooper, Van Soest & Pisera 2011).

Class Calcarea Bowerbank, 1864 (2 subclasses)
Subclass Calcinea Bidder, 1898 (2 orders)
Order Clathrinida Hartman, 1958 (6 families, +1 incertae sedis)
Family Clathrinidae Minchin, 1900 (2 genera, 86 species)
Family Leucaltidae Dendy & Row, 1913 (4 genera, 18 species)
Family Leucascidae Dendy, 1893 (2 genera, 16 species)
Family Leucettidae de Laubenfels, 1936 (2 genera, 24 species)
Family Soleneiscidae Borojevic, Boury-Esnault, Manuel & Vacelet, 2002 (2 genera, 13 species)
Family Levinellidae Borojevic & Boury-Esnault, 1986 (3 genera, 4 species)
Clathrinida incertae sedis (1 genus (Leucomalthe), 1 species)
Order Murrayonida Vacelet, 1981 (3 families)
Family Lelapiellidae Borojevic, Boury-Esnault & Vacelet, 1990 (1 genus, 2 species)
Family Murrayonidae Dendy & Row, 1913 (1 genus, 1 species)
Family Paramurrayonidae Vacelet, 1967 (1 genus, 1 species)
Subclass Calcaronea Bidder, 1898 (3 orders)
Order Leucosolenida Hartman, 1958 (9 families)
Family Leucosoleniidae Minchin, 1900 (3 genera, 47 species)
Family Amphoriscidae Dendy, 1892 (3 genera, 33 species)
Family Grantiidae Dendy, 1892 (11 genera, 208 species)
Family Heteropiidae Dendy, 1893 (7 genera, 65 species)
Family Lelapiidae Dendy & Row, 1913 (5 genera, 8 species)
Family Sycettidae Dendy, 1892 (2 genera, 94 species)
Family Jenkinidae Borojevic, Boury-Esnault & Vacelet, 2000 (6 genera, 11 species)
Family Achramorphidae Borojevic, Boury-Esnault, Manuel & Vacelet, 2002 9 (2 genera, 10 species)
Family Sycanthidae Lendenfeld, 1891 (2 genera, 4 species)
Order Lithonida Vacelet, 1981 (2 families)
Family Minchinellidae Dendy & Row, 1913 (5 genera, 19 species)
Family Petrobionidae Borojevic, 1979 (1 genus, 1 species)
Order Baerida Borojevic, Boury-Esnault & Vacelet, 2000 (3 families)
Family Baeriidae Borojevic, Boury-Esnault & Vacelet, 2000 (4 genera, 12 species)
Family Trichogypsiidae Borojevic, Boury-Esnault & Vacelet, 2000 (3 genera, 5 species)
Family Lepidoleuconidae Vacelet, 1967 (1 genus, 1 species)


The Calcarea, or calcareous sponges, have the skeleton composed exclusively of calcium carbonate spicules. The structure ranges from discrete monactinal, diactinal, triactinal or tetractinal spicules (Hooper & Wiedenmayer 1994: figs 147–182), to reticulate skeletons composed of fused crystalline calcite spicules. The distinction between megascleres and microscleres is not recognised for this class. Calcarea are always viviparous.

The skeleton and aquiferous system of the Calcarea occurs in three grades of construction. The asconoid system consists of a simple tubular construction, without folding of the body wall, with thin walls pierced externally by ostia, leading to tubular water canals (porocyte canals) opening onto a central choanocyte-lined cavity (choanoderm), connected to the exterior by a single osculum at the apex of the sponge. Syconoid construction is produced by folding of both the exterior (pinacoderm) and interior (choanoderm) walls, producing choanocyte chambers which lie within the body wall rather than only lining the central atrium as in simpler asconoid structures, but these chambers open directly onto the atrium. The most complex grade of construction is leuconoid, found in most sponges (including the Demospongiae), in which the choanocyte chambers are ovoid and isolated in a maze of canals within the body wall, with chambers opening onto branching and complex excurrent canals.

Calcareous sponges are most common in shallow, relatively sheltered waters (<1000 m), and are predominant in temperate regions, or in the tropics where they are mainly associated with coral reefs. Two subclasses are recognised, differentiated by larval morphology and the grade of organisation of the aquiferous system, Calcinea (with two orders Clathrinida and Murrayonida), and Calcaronea (with two orders Leucocoleniida and Lithonida). There are about 400 described species.

Sponges in the subclass Calcinea have entirely ciliated hollow blastula (coeloblastula) larvae. Most species have regular triradiate free spicules (Hooper & Wiedenmayer 1994: fig. 147), which are equiangular and equiradiate, or exceptionally, parasagittal or sagittal triradiate spicules (Hooper & Wiedenmayer 1994: figs 149–156). Choanocytes are basinucleate; nuclei are spherical, and the basal body of the flagellum is not adjacent to the nucleus. Some species also have monactinal or diactinal free spicules (Hooper & Wiedenmayer 1994: figs 164–174) as well as, or instead of, triradiates. Recent revisions are given by Borojevic & Boury-Esnault (1986) and Borojevic et al. (1990).

Members of the Clathrinida are simple calcareous sponges with a skeleton composed exclusively of free spicules, without hypercalcified non-spicular reinforcements or spicule tracts. Six families are known, five with published records for the Australian fauna.

The second order, Murrayonida, has a reinforcement of the skeleton composed of either spicule tracts, calcareous plates or a rigid monospicular skeleton; the canal system is leuconid in the three known species; diapasons or tuning-fork shaped triradiates (Hooper & Wiedenmayer 1994: figs 179–180, 182), or modified biradiates are present, and are generally fasciculated (Vacelet 1981; Borojevic et al. 1990). Three families are represented but none yet recorded from the Australian fauna. Extral-limital families are Murrayonidae, with a single Recent genus and species that is widely distributed on reefs in the Pacific Ocean and in the eastern Pacific Ocean; Paramurrayonidae with a single genus and species from Madagascar; and Lelapiellidae with a single genus in cryptic habitats of shallow coral reefs of New Caledonia (Vacelet 1981; Borojevic et al. 1990).

Sponges of the subclass Calcaronea have incubated amphiblastula larvae which are flagellated only on the anterior half. The nuclei of choanocytes are apical in position, and the flagellum arises directly from the nucleus. Spicules are triradiate (Hooper & Wiedenmayer 1994: fig. 44) and sagittal (two rays are paired and the third ray is longer than the others; Hooper & Wiedenmayer 1994: figs 149, 154, 155), as well as free monaxonic (monactinal or diactinal) forms (Hooper & Wiedenmayer 1994: figs 164–178), and there is usually a system of sagittal tetractines, exceptionally regular, forming a basal skeleton. The aquiferous system ranges from asconoid to leuconoid grades of construction.

Two orders are recognised. The order Leucosoleniida includes sponges with asconoid grade of construction, consisting of long branching tubes, each with a terminal osculum, in which choanocytes line the central atrium; spicules are triradiate (Hooper & Wiedenmayer 1994: fig. 147), sagittal with unequal angles (Hooper & Wiedenmayer 1994: fig. 155), and there are non-spicular reinforcements in the skeleton. Seven families are known of which six are represented in Australian waters.

The order Lithonida are sponges that generally occur in shaded habitats such as caves and tunnels. They have a massive reinforced calcitic skeleton, together with randomly distributed tuning-fork spicules (Hooper & Wiedenmayer 1994: figs 179–182) and an armature of quadriradiate spicules (Hooper & Wiedenmayer 1994: figs 149, 154, 155) around each osculum. Larvae are blastulae and amphiblastulae.

There are three families, only one reported so far from Australian waters, Minchinellidae, which is dispersed across the Indo-Pacific, with records from Sagami Bay, Japan, Vanuatu, Tulear, western Indian Ocean and other Indo-Pacific reefs (Vacelet 1981; Reitner 1992). Some of these other groups are undoubtedly present in the Australian coral reef fauna, but very few sponge workers are familiar with this highly specialised calcareous fauna.

Several recent studies on the calcareous sponges (e.g. Vacelet 1981, 1991; Borojevic 1979; Borojevic & Boury-Esnault 1987; Borojevic et al. 1990) have produced dramatic changes in the taxonomy and phylogenetic interpretation of this group since a synthesis of the class was presented by Hartman (1982). These revisions are still incomplete (particularly for the order Calcaronea); they are, nevertheless, currently in progress (Boury-Esnault, personal communication). The recognition of only two lines of Calcarea evolution, first proposed by Bidder (1898), is largely supported by recent revisions (with abandonment of ‘subclass Pharetronida’ by Vacelet (1991) and removal of ‘subclass Sphinctozoa’ to the ceractinomorph Demospongiae by Vacelet (1979, 1985)), although the distribution of orders and families within these subclasses is now founded on very different characters from those originally proposed.

The classification of Burton (1963) is completely rejected, being based on external shape of the sponge, and presence or absence of certain spicule types whereas anatomical, cytological and embryological differences indicate that the water current system has developed in parallel in these two subclasses irrespective of grade of organisation (asconoid, syconoid or leuconoid) (Borojevic 1979; Vacelet 1991). Major characters used to differentiate subclasses are: choanocyte ultrastructure (i.e. having an apical nucleus from which the flagellum arises, or a basal nucleus without connection with the flagellum), larval morphology (amphiblastula larva derived by eversion of a stomoblastula, versus a coeloblastula which becomes a solid, free swimming blastula). Adult spicule morphology is diagnostic with triradiate spicules mostly sagittal (e.g. Hooper & Wiedenmayer 1994: figs 155, 157, 159, 160), versus triradiates usually regular, rays of equal length at equal angles (e.g. Hooper & Wiedenmayer 1994: figs 147, 152, 153, 156), and in larvae, diactines versus triactines, for Calcaronea and Calcinea, respectively (Vacelet 1991).

Note that in conventional zoological nomenclature, subclass endings ‘ia’ are usual. However, this convention has been ignored by most sponge workers. As the ICZN does not govern superfamilial nomenclature, we are not obliged to alter the current usage of ‘ea’ endings for subclasses Calcaronea and Calcinea in the interests of stability of nomenclature, although this usage is at odds with other animal phyla. {reitnerj1992a}



Marine Porifera in which the mineral skeleton is composed entirely of calcium carbonate. The skeleton is composed of free diactine, triactine, tetractine and/or polyactine spicules, to which can be added a solid basal calcitic skeleton with basal spicules either cemented together or completely embedded in an enveloping calcareous cement. The aquiferous system can be asconoid, syconoid, sylleibid or leuconoid. Members of the Calcarea are viviparous and their larvae are blastulae.


ID Keys

(1) Regular (equiangular and equiradiate) triactines and tetractines, choanocytes basinucleate with spherical nuclei------------------------------------------------------------------------------------(Calcinea)--------2
Sagittal triactines & tetractines, choanocytes apinucleate-------------------(Calcaronea)---------3

(2) Skeleton composed exclusively of free spicules------------------------------------------------Clathrinida
Reinforcement of the skeleton composed of either spicule tracts, calcareous plates or a rigid nonspicular skeleton. Diapasons or modified diactines present and generally fasciculated------------------------------------------Murrayonida

(3) Skeleton composed exclusively of free spicules--------------------------------------------------------------4
Reinforcement of the skeleton consisting either of linked or cemented basal actines of tetractines, or of a rigid basal mass of calcite. Diapason spicules generally present. Aquiferous system leuconoid----------------------------------Lithonida

(4) Skeleton either composed exclusively of microdiactines or in which microdiactines constitute exclusively or predominantly a specific sector of the skeleton. Large or giant spicules frequently present in the cortical skeleton. Dagger-shaped small tetractines (pugioles) frequently the sole skeleton of the exhalant aquiferous system. Aquiferous system leuconoid------------------------------------------------------------------------Baerida
Free spicules diactines, sagittal triactines/tetractines. Aquiferous system asconoid, syconoid, sylleibid or leuconoid------------------------------------------------------------Leucosolenida


General References

Borojevic, R. 1979. Evolution des Spongiaires Calcarea. pp. 527–530. in Lévi, C. & Boury-Esnault, N. (eds) Biologie des Spongiaires/Sponge Biology. Colloques Int. Cent. Natl Rech. Scient. Vol. 291.

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. & Boury-Esnault, N. 1986. Une nouvelle voie d'évolution chez les éponges Calcinea: description des genres Burtonulla n.g. et Levinella n.g. Bulletin du Muséum d'Histoire Naturelle. Paris 4 8(A,3): 443-455

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

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

Dohrmann, M., Voigt, O. & Wörheide, G. 2006. Non-monophyly of most supraspecific taxa of calcareous sponges (Porifera, Calcarea) revealed by increased taxon sampling and partitioned Bayesian analysis of ribosomal DNA. Molecular Phylogenetics and Evolution 40(3): 830-843

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., 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

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]

Manuel, M. 2006. Phylogeny and evolution of calcareous sponges. Canadian Journal of Zoology 84: 225–241

Manuel, M., Borojevic, R., Boury-Esnault, N. & Vacelet, J. 2002. Class Calcarea Bowerbank, 1864. pp. 1103-1110 in Hooper, J.N.A. & Soest, R.W.M. Van (eds). Systema Porifera. A guide to the classification of sponges. New York : Kluwer Academic/Plenum Publishers Vol. 2. [1103]

Reitner, J. 1992. "Coralline Spongien" Der Versuch einer phylogenetisch-taxonomischen Analyse. Berlin, Germany : Selbstverlag Fachbereich Geowissenschaften, Freie Universitaat Berlin 352 pp.

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. 1981. Eponges hypercalcifiées ('Pharétronides', ‘Sclérosponges') des cavités des récifs coralliens de Nouvelle-Calédonie. Bulletin du Muséum National d'Histoire Naturelle. Paris 4,A 3: 313-351

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. 1991. Recent Calcarea with a reinforced skeleton ('Pharetronids'). pp. 252-265 in Reitner, J. & Keupp, H. (eds). Fossil and Recent Sponges. Berlin, Heidelberg : Springer-Verlag xviii 595 pp.

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. Calcarea. 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 on 2012-01-27. (checked on 2012-01-27.)

Van Soest, R.W.M., Boury-Esnault, N., Vacelet, J., Dohrmann, M., Erpenbeck, D., de Voogd, N.J., Santodomingo, N., Vanhoorne, B., Kelly, M. & Hooper, J.N.A. 2012. Global Diversity of Sponges (Porifera). PLoS ONE (Public Library of Science) PONE-S-11-32233


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)