Professional Documents
Culture Documents
John N.A. Hooper. Queensland Museum, PO Box 3300, SOUTH BRISBANE, QLD, 4101, Australia
(email: JohnH@qm.qld.gov.au).
Two versions of this ‘Sponguide’ appear on the internet in Adobe Acrobat format:
http://www.qmuseum.qld.gov.au/organisation/sections/SessileMarineInvertebrates/index.asp
[“Download the Guide to Sponge Collection and Identification.”]
FOREWARD
This sponge identification guide is an ongoing project of the Queensland Museum. It was originally
conceived as a response to an overload of requests for sponge identifications, particularly from the
professional community. Diagnoses of many genera are still incomplete, which would take several more
years to accomplish. In any case, this guide will be superceded eventually by a more comprehensive
‘Systema Porifera’, a collaborative project amongst 30 sponge taxonomists to publish the supraspecific
classification of living and fossil sponges, providing a sound baseline for higher systematics debate and a
bench document for routine classification (Hooper & Van Soest (eds), in preparation, expected date of
publication 2002). The main intention in distributing the current Sponguide is to encourage other
scientists to undertake at least some level of standard preparation and identification (i.e. taxonomic
sorting) of their material prior to sending it to us or others for further identification, to make our lives a
little bit easier.
There are no original illustrations included in this version, and for the time being the Sponguide should
be used in conjunction with the recommended references listed at the end of the document which contain
some relevant illustrations.
CONTENTS
0. List of sponge higher taxa.
1. Introduction to sponges
2. Methods of dealing with sponges in the laboratory and preparation for their identification.
3. Outline of characters used for Demospongiae identification.
4. Major characters used to describe the Demospongiae, based on the 'DELTA' (CSIRO computerised
descriptions) format.
5. Key to the extant orders of Porifera.
7. The sponge classification (extant taxa only).
8. Preferred format for sponge samples sent for identification.
9. Recommended reading
10. Glossary.
11. Illustrations.
12. Index to genera of extant Porifera.
Order Verticillitida.
Family Cryptocoeliidae Steinmann, 1982. Class Calcarea.
Order Agelasida.
Family Agelasidae Verril, 1907. Subclass Calcinea.
Family Astroscleridae Lister, 1900. Order Clathrinida.
Order Poecilosclerida. Family Clathrinidae Minchin, 1900.
Suborder Microcionina . Family Soleniscidae Borojevic et al., 1990.
Family Iophonidae Burton, 1929. Family Levinellidae Borejevic et al., 1986.
Family Microcionidae Carter, 1875. Family Leucaltidae Dendy & Row, 1913.
Family Raspailiidae Hentschel, 1923. Family Leucascidae Dendy, 1893.
Family Rhabderemiidae Topsent, 1928. Family Leucettidae de Laubenfels, 1936.
Suborder Myxillina. Order Murrayonida.
Family Anchinoidae Topsent, 1928. Family Murrayonidae Kirkpatrick, 1910.
Family Coelosphaeridae Hentschel, 1923. Family Paramurrayonidae Vacelet, 1967.
Family Crambidae Lévi, 1963. Family Lelapiellidae Borojevic, Boury-Esnault
Family Crellidae Hentschel, 1923. & Vacelet, 1990 .
Family Hymedesmiidae Topsent, 1928. Subclass Calcaronea.
Family Myxillidae Topsent, 1928. Order Leucosoleniida.
Family Phoriospongiidae Lendenfeld, 1888. Family Leucosoleniidae Minchin, 1898.
Family Tedaniidae Ridley & Dendy, 1886. Family Amphoriscidae Dendy, 1892.
Suborder Mycalina. Family Grantiidae Dendy, 1892.
Family Cladorhizidae de Laubenfels, 1936. Family Heteropiidae Dendy, 1893.
Family Guitarridae Burton, 1929. Family Lepidoleuconidae Vacelet, 1967.
Family Desmacellidae Ridley & Dendy, 1886. Family Sycettidae Dendy, 1892.
Family Hamacanthidae Gray, 1872. Family Staurorrhaphidae Jenkin, 1908.
Family Mycalidae Lundbeck, 1905. Order Lithonida.
Order Halichondrida. Family Lelapiidae Dendy & Row, 1913.
Family Axinellidae Carter, 1875. Family Minchinellidae Dendy & Row, 1913.
Family Desmoxyidae Hallmann, 1917. Family Petrobionidae Borojevic, 1979.
Family Dictyonellidae van Soest,Diaz &
Pomponi, 1990. Subphylum Symplasia
Family Halichondriidae Vosmaer, 1887.
Order Haplosclerida. Class Hexactinellida.
[Marine haplosclerida families]:
Family Callyspongiidae de Laubenfels, 1936. Subclass Amphidiscophora.
Family Chalinidae Gray, 1867. Order Amphidiscosida.
Family Niphatidae Van Soest, 1980. Family Hyalonematidae Gray, 1857.
Family Phloeodictyidae Carter, 1882. Family Monorhaphididae Ijima, 1927.
Family Petrosiidae van Soest 1980. Family Pheronematidae Gray, 1870.
[Freshwater haplosclerid families]. Subclass Hexasterophora.
Family Spongillidae Gray, 1867. Order Hexactinosida.
Family Metaniidae Volkmer-Ribeiro, 1986. Family Aphrocallistidae Gray, 1867.
Family Potamolepidae Brien, 1967. Family Aulocalycidae Ijima, 1927.
Family Lubomirskiidae Brien, 1969. Family Craticulariidae Rauff, 1893.
Order Dictyoceratida. Family Euretidae Zittel, 1877.
Family Ircinidae Gray, 1867. Family Farreidae Gray, 1872.
Family Thorectidae Bergquist, 1978 Family Tretodictyidae Schulze, 1886.
Family Spongiidae Gray, 1867. Order Lychniscosida.
Order Dendroceratida. Family Aulocystidae Schulze, 1886.
Family Dysideidae Gray, 1867. Family Dactylocalycidae Gray, 1867.
Family Darwinellidae Merejkowsky, 1879. Order Lyssacinosida.
Family Dictyodendrillidae Bergquist, 1980. Family Caulophacidae Schulze, 1886.
Family Halisarcidae Vosmaer, 1885. Family Euplectellidae Gray, 1867.
Order Verongida. Family Leucopsacasidae Ijima, 1903.
Family Aplysinidae Carter, 1875. Family Rossellidae Gray, 1872.
Family Druinellidae Lendenfeld, 1889.
Family Ianthellidae Hyatt, 1875.
1. INTRODUCTION TO SPONGES
qWHAT ARE SPONGES ?
l Sponges are the most primitive of multicellular animals (metazoa).
l They have a cellular grade of construction without true tissues. Body plans range from simple
(asconoid, syconoid) through to complex (leuconoid) produced by varying degrees of infolding of the
body wall and complexity of water canals throughout the sponge.
l Adults are asymmetrical or radially symmetrical.
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 3
l Sponges are exclusively aquatic (water dwelling), most marine, found from deepest oceans to the
edge of the sea.
l Sponges play such important roles in so many marine habitats but we still know very little about their
diversity, biology and ecology as compared with most other animal groups. In many benthic (sea bottom)
habitats sponges are often the dominant animals.
l Sponges have evolved an amazing range of growth forms, best described as highly irregular and
sometimes completely plastic, frequently altered by prevailing external conditions (currents, turbidity,
salinity etc.). Sponges also have evolved an amazing array of colours (possibly linked to photoprotection
?)
l Adult sponges are sedentary (sessile), attached to the seabed or other substrate for most of their lives,
although many have larvae that motile, swimming or crawling away from their parent.
l Sponges have sexes that are separate, or sequencially hermaphroditic, although most population
dispersal and recruitment is asexual (through budding, fragmentation from storm events, etc).
l Larvae are motile, incubated within the parent or broadcast into the seawater: parenchymella (solid,
ciliated), amphiblastula (central cavity).
l Sponges filter sea water to eat, breath and excrete waste products. Sponges often have complex water
canal systems running throughout the body, with smaller inhalant (ostia) and larger exhalant pores
(oscules). Sponges are able to actively pump up to 10 times their body volume each hour, making them
the most efficient vacuum cleaners of the sea.
l Sponges appear to be very stable, long-lived animals, although growth rates vary enormously between
different groups. Some sponges, like haplosclerids can grow centimetres in weeks, and may have shorter
life spans. Others sponges, like the living fossil 'sclerosponges' are VERY slow growing, with the largest
known individuals (up to 30cm diameter) thought to be around 5,000 years old (which makes them the
oldest living individuals on the planet, if this is true !).
l Sponges are a unique group of animals because ....
l Sponges have unique collar-cells (choanocytes) which are surrounded by cilia with a central
flagellum that moves to actively create a current pulling water in and out of the sponge. These collar
cells line the walls of small chambers throughout the water vascular system. There may be 7,000-18,000
of these chambers per cubic millimeter of sponge, and each chamber may pump approximately 1,200
times its own volume of water per day !
l Sponges have no tissues or sensory organs but they do have MANY different types of cells with
MANY different functions that carry out normal bodily routines, including a primitive cell type (called
an archaeocyte, an amoeboid-like cell) that is totipotent (able to change functions as required by the
sponge [e.g. secrete the skeleton, form the epidermis, become feeding and reproductive cells etc.]).
l Outer and inner layers of cells (exopinacocytes, basipinacocytes) (="the skin") lack a basement
membrane; middle layer.(mesohyl) is variable but always includes motile cells and usually some skeletal
material.
l Mineral skeleton is present in most (but not all) groups of sponges composed of calcium carbonate,
silicon dioxide, and/or collagen fibres.
l Skeletal elements (spicules) are diverse in their geometry and size.
l Sponges are individuals, having a continuous "skin" (epithelium) that contains roving cells inside;
they are not colonies (like corals and sea-squirts in which individuals animals group together).
l Sponges catch, eat, digest their food and excrete their waste products within cells, not within any
common body cavity, unlike most multicellular animals.
l Some sponges (particularly those growing on coral reefs) have a unique symbiosis with
cyanobacteria not found in any multicellular animal. These cyanobacteria (or blue-green algae) provide
the sponge with nutrients from photosynthesis to supplement those obtained by the sponge from normal
filter feeding activities. These extra nutrients greatly augments sponge growth rate and competitive
ability in coral reef systems.
4 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
l Sponges are morphological conservative, with a VERY ancient geological history they were already
well established during the Lower Cambrian (>550 million years ago), and were major reef builders
during the Devonian (>370 million years ago) before the reef-building corals "took over" reef
formation; most modern genera and species similar to those around today appeared at the beginning of
the Cretaceous (>150 million years ago).
qWHAT DO SPONGES DO ?
l Sponges don't appear to do much at all, but in fact they are very active ! Sponges sit on the bottom of
the sea actively pumping seawater through their bodies. This water, containing nutrients, is filtered
through a series of sieve-like pores (diminishing in size), finally ending up at the collar cells. Nutrients
are actively carried across the cell wall, engulfed by archaeocytes, and subsequently transferred
throughout the mesohyl.
l In addition to “regular food”, this seawater also contains the toxic chemicals excreted by other plants
and animals, such as corals growing on the coral reefs above. Sponges feed on these chemicals, modify
and reuse them for their own purposes.
l Many of these sponge-modified and sponge-produced chemicals have potent toxicity against human
pathogens, cancer cells etc., or are useful against certain human ailments (e.g. anti-inflammatory,
cardiovascular, respiratory, analgesic etc. properties).
l Why ..... ?
l Sponges are not mobile, and as such they cannot escape from predators (such as fishes, turtles,
gastropods, echinoderms, flatworms). They are generally unable to physically defend themselves.
Perhaps they use these chemicals to defend themselves ?
l Sponges do not have arms or legs and so they cannot physically remove other animals and plants
settling on their exterior surfaces and from within water canals. Perhaps they use these chemicals to
repel parasites ?
l Sponges are generally slow-growing, easily out-competed for space and living resources by other
faster growing animals and plants (such as corals and ascidians). Perhaps they use this arsenal of
chemicals in a chemical warfare against these faster-growing species ?
l Sponges are often full of small animals and microbes, particularly embedded within body cavities and
throughout the water canals. Perhaps many of these chemicals are antibiotic to these microbes ?
l Some sponges burrow into corals and use chemicals to eat away the calcium, eventually occupying the
entire interior surface of the coral (with breathing tubes or fistules poking through the surface). Other
sponges grow on top of corals, smothering and eventually eroding the dead coral, and some that bore
into oysters and pearl oyster shells killing the animal inside. There are also many sponges that live in
between coral branches, at the base of corals and in the substrata surrounding corals that bind the corals
together. Perhaps some of these chemicals are important in the continual process of bioerosion of coral
reefs and releasing calcium back into the system, and consolidation of dead coral producing stable reef
structures ?
l Some sponges have intimate, symbiotic relationships with other animals (such as gastropods, hermit
crabs, shrimps), or plants (blue-green algae or cyanobacteria), which is frequently species specific.
Perhaps these chemicals produce recognition signals between the symbiotic partners ?
l Or maybe all of these factors are important ?
years or so. Each of these regions probably has between 400-600 species, with about 100 shared species
at most.
l Other places have poorly known sponge faunas, and (worse still) most of our knowledge is from old
expeditions (pre-1900s; which makes "data" even more dubious). These areas include Australia,
Indonesia, Papua New Guinea and other parts of the Indo-Malay archipelago. However, it is well known
from studies on other groups of animals that this region has the highest proportion of the world's marine
biodiversity, and there is no reason to show us that this is different for sponges.
l Worldwide: In the literature there are about 7,000 "valid" species published worldwide, BUT we
estimate that there are at least 15,000 living species in all the world's seas and lakes.
l Australia: In Australia there are about 1,400 species described in the scientific literature, but we
estimate that there are probably at least 5,000 species living in continental and territorial waters. We
know this because currently our collections contain about 4,000 species of sponges, mostly Australian,
with most (about 2,000) new to science and requiring descriptions and naming in the scientific
literature. We haven’t yet begun to look comprehensively at the cryptic and encrusting fauna which is
probably as equally diverse as the macrobenthic fauna. Documenting and describing biodiversity is a
long, time consuming process that requires accuracy and patience, but it is an essential prerequiste to
conservation and management of our marine resources.
l Indo-Malay Archipelago and South China Sea: There are approximately 1200 described species of
sponges known from the South China Sea region (extending from Cocos, Nicobar and Andaman Islands,
the Andaman Sea, islands and coast of Burma, west coasts of Thailand and the Malay Peninsula,
Singapore, islands and waters of the Straits of Malacca, all the islands of the Indonesian Archipelago, the
Gulf of Siam, east coast of the Malay peninsula, Thailand, Cambodia, Vietnam; Borneo, Sarawak, Sabah,
Brunei; southern China, Hong Kong; Philippines, Palawan. (This excludes Papua New Guinea and
northern Australia) which have largely different sponge faunas. However, it is thought (with some
justification) that this region may contain a very high diversity, perhaps in the order of 4000-6000
species, but it is probably also the least comprehensively documented fauna by modern standards given
that most of our knowledge of the fauna derives from pre- and early 1900 literature (plus generally
unreliable identifications made by collectors for chemical studies).
l Organic basis for toxicity (a new molecule in the class 'thio sugar') found in Clathria pyramida from
southern New South Wales (extremely effective antibiotic and anti-spermaticide (???) without toxicity
towards mammalian tissues).
l Organic basis for toxicity (Amphimedon terpenensis from the Great Barrier Reef) effective against
human tumours and some microbes, but also toxic towards mammalian tissues.
l Other animals that feed on sponges can concentrate and modify (sequester) the sponge chemicals for
their own purposes (presumably for chemical defence), either making the sponge molecule less toxic or
more toxic (Cacospongia mollior from the Mediterranean with the nudibranch gastropods Glossodoris
and Chromodoris).
l Most celebrated example is Dysidea avara from the Mediterranean producing an effective anti-AIDS
molecule.
lA more recent example is a new genus and species of sponge from the Great Barrier Reef (Cymbastela
hooperi and C. coralliophila) with anti-malarial compounds of considerable potential for
pharmaceutical use.
q 2.1. Collection.
Sponges are often soft bodied, many are fragile and colours are generally unstable (e.g. aerophobic,
soluble). Many sponges are also harmful to humans, producing physical damage (e.g. from sharp
spicules protruding through the surface) and/or with an irritating mucus and other chemicals, sometimes
causing severe dermitis. Consequently, special care must be taken when collecting to minimise damage
to both the sponge and collector. Sponges may be removed from the substrate with a knife or chisel,
preferably using protective gloves and protective clothing.
Collections of sponges intended for identification should be accompanied by in situ photographs and
adequate documentation (locality, habitat, surface features, colour notes etc.). In many species both
colouration and morphology may change dramatically following collection and preservation, and
identifications, even by specialists, are often greatly facilitated if there are adequate colour photographs
of live material.
Sponges may also be air dried in the sun, although many may lose their shape, most lose their
colouration (but few lose their noxious smell!). For several groups of sponges (i.e. those which have
strong fibre skeletons such as the commercial 'bath sponges'), specimens may be rotted in freshwater and
subsequently washed in solutions of potassium permanganate and then sodium metabisulphide. This
technique softens and cleans the fibrous skeleton from incorporated sand particles.
2.3.1. Spicule preparations. For spicule preparations several simple methods are available, none of
which requires extensive experience or sophisticated equipment.
2.3.1.1. Bleach digestion. This technique is useful for rapid surveys of spicules within a sponge,
although preparations are not as clean as those obtained through an acid digestion process. Sponges with
calcareous spicules are routinely prepared in this manner. Small fragments of 'tissue', including
fragments from both the surface and deeper parts of the sponge, are placed in small Ehrlenmeyer flasks
or directly on microscope glass slides. A small quantity of active bleach (sodium hypochlorite) is added
to the fragment, and after a short period the organic components dissolve leaving only the mineral
skeleton. The bleach must then be carefully diluted and eventually washed out of tissues several times,
replaced firstly with water and then with ethanol. If bleach is not completely removed preparations
become crystalline. Finally, clean spicule suspensions are aspirated and pipetted onto a glass slide, the
ethanol allowed to evaporate, and mounted. It is important to note that during each stage of pipette wash
the suspension should be left to settle for about 10-15 minutes, prior to decanting the supernatant, to
avoid accidental decanting of smaller spicules. Using flasks for the actual digestion process, instead of
slides, has the advantage that a centrifuge can be used to eliminate the settling time of the supernatant.
Conversely, preparations made directly on slides have the advantage that spicules do not have to be
pipetted, and hence minimising the potential for losing the smaller spicules.
2.3.1.2. Acid digestion. This technique provides cleaner, permanent preparations, but the process
involves noxious chemicals and should be undertaken only with suitable facilities (e.g. protective
clothing, fume extraction). This process uses nitric acid instead of bleach. Fragments of sponge are
placed in flasks, directly on glass slides, or directly on electron microscope stubs. Several drops of acid
are placed on the fragment, gently heated over a flame until bubbling, and repeated until all organic
matter is digested (this is easily ascertained by eye). The heat-accelerated digestion process produces
various oxides, including nitrous oxide, and it is cautioned that these are noxious. It should also be
noted that the acid is evaporated rather than burnt, so low heat is preferrable (e.g. using an alcohol flame
rather than gas). Once dry and cool, preparations can be mounted immediately without washing.
Siliceous spicules are bonded directly onto the substrate by this technique, which makes it useful for
both light and scanning electron microscopy.
Spicule preparations obtained from both techniques are now ready for covering using a suitable
mounting medium (e.g. Depex, Canada balsam, Euparol, Durcupon, etc.).
8 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
2.3.2. Section preparations. For sponge sections there are more complex procedures involved, using
microtome-sectioning or at least thick, hand-cut sections. The object of these techniques is to observe
skeletal structures and cytological characteristics as much as they appear in the live animal, so wax
embedding techniques, staining and/or simple clearing agents are required. Several techniques are
available, most requiring specialist histological facilities.
2.3.2.1. Simple clearing. The easiest method to determine the structure of the mineral skeleton is to
using thick hand-cut sections cleared in a clearing agent (e.g. toluene, xylene, phenol-xylene, Histosol,
Histoclear, lactophenol creosote, etc.). A perpendicular section through the surface and deeper skeleton
is cut from a larger, preserved fragment of sponge by hand, using a new, clean scalpel. Relatively even,
thick sections (between 50-100µm thickness) are possible using hand-cutting techniques, but success is
certainly linked with practice. Cut sections are placed directly in a saturated mixture of phenol and
xylene (matured for at least 1 week) to clear the section, which eliminates the need for an alcohol
dehydration series. Clearing may take between 4-24 hours, depending on the development of collagen in
the species. Cleared sections can be mounted directly on slides, but cover glasses should be supported
with glass slivers or card on thick sections.
2.3.2.2. Wax embedding. To produce a perfectly uniform section thickness, and for thin sections, wax
embedding and microtome techniques are required. Fragments of preserved sponges should be passed
through a dehydration series, cleared in toluene, and wax embedded for at least 2 hours. Alternatively,
fixed samples can be processed directly in an automated paraffin embedding system, such as Tissue Tek
V.I.P. 2000, on a 16-48 hour cycle. Sections should be cut from trimmed wax blocks, cutting from the
centre of the block to the exterior so as to include both the outer surface and inner skeleton relatively
intact. For most species relatively thick sections are required (>50µm), so as to avoid breaking the
spicules in situ, but for 'keratose' (non-spiculous) sponges thinner sections are preferrable. Cut sections
are placed in clearing agent for an adequate period to dissolve wax and clear the 'tissue', then soaked in
ethanol (perhaps clearing and dehydrating several times until perfectly clear, and/or dewaxing on a hot
plate), floated onto slides, orientated and flattened, and mounted.
2.3.2.3. Staining. Staining as a technique for sponge histology is not a widespread procedure given that
taxonomy of most groups is base on the inert silicon (or calcareous) skeleton. For some groups, such as
the 'keratose' sponges, or where histological information is required, staining such as Mallory
Heidenhain's solution is useful. Sections are processed, wax embedded, cut and placed on slides as
described above. Sections placed on albumen coated slides are drained and dried at 60°C, dewaxed in
xylol for at least 5 minutes, hydrated through an alcohol series to water, stained in Mallory Heidenhain
for 7 minutes, dipped in tap water 6 times, dehydrated in an alcohol series, cleared in xylol and mounted.
generally obtained on coated specimens at higher voltage. Typical viewing conditions are 25kV,
at small working distances to provide best depth of field and focus, and at low magnifications.
q Sponge identifications are primarily based on morphology. Some of these morphological characters
vary substantially between widely separated populations, or those living in different habitats, indicating
no more than ecophenotypic variation within the species, whereas other features are much more
consistent between individuals irrespective of their geographic distribution. Unfortunately, however, we
are still not completely sure which of these 'variable characters' indicate population variability within a
single species and which are consistent in the evolution of species, and thus are more important in
sponge taxonomy. Over recent years many new non-morphological characters have been discovered from
genetic, biochemical and ultrastructural studies of sponges. Some of these characters have been useful in
supporting or refuting current ideas on morphological-based sponge taxonomy, but in other instances it
is difficult to find any morphological characters that correspond to these new schemes: ultimately
taxonomy must somehow be related to the morphology to be of practical value.
Consequently, most sponges are not easy to identify, even for experts, requiring specialised techniques.
Some of of these techniques are outlined below, including the preferred methods for collection,
documentation, histological preparation, and a brief explanation of many of the features used to identify
sponges.
The sponge is bounded on the exterior surface by a unicellular layer (exopinacoderm), called the
ectosome, composed of special epithelial cells (pinacocytes). Some of these epithelial cells form small
external pores (ostia) through which the water passes into the sponge, and others form larger pores
10 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
(oscules) through which water is expelled. Internally (called the choanosome) the sponge is excavated
by water current canals, also lined by a single layer of cells (porocytes) forming the endopinacoderm.
'Water pumping stations' (choanocyte chambers) are found at certain locations along the water canals,
lined by special collar cells with a flagellum (choanocytes), unique to the Porifera. A water current is
created by the beating of choanocyte flagella. Water is actively pumped through the water filtration
system, through a series of sieves or filters of diminishing size, which serve to extract nutrients and
oxygen from the incoming water. Similarly, waste products are expelled into the excurrent canals and
jettisoned through the oscules. For the Calcarea the development of the water current system, from
simple to complex (asconoid, syconoid, leuconoid), is important in its taxonomy, whereas in the
Demospongiae most are of the complex (leuconoid) construction.
The living 'tissue' bounded on all sides by the pinacoderm is called the mesohyl. This contains a matrix
or ground substance composed of a striated protein called collagen, an organic skeleton composed of
spongin fibres, and/or an inorganic skeleton composed of mineral spicules. Within this mesohyl are
found roving totipotent cells, capable of changing function as required. These include generalised
amoebocytic 'stock' cells (archaeocytes), as well as many other types that have become specialised to
carry out particular functions within the sponge, such as producing fibres (collencytes), secreting
spicules (sclerocytes), contractile cells around excurrent pores or oscules (myocytes), and so on. There
are many sorts of cells in sponges and only few so far have a known function. Attempts have been made
to use these cytological characters in a taxonomic framework but with limited success.
There are many morphological characters which can be used to aid in sponge identification including
shape, distribution of surface pores, colour, ornamentation of the surface, texture, structure and
composition of the organic skeleton and water canal system, and the structure, composition, size and
geometry of the inorganic skeleton. In addition, several non-morphological features have proven useful
practical tools in sponge taxonomy.
q 3.2. Major characters used in sponge identification [see illustrations at end of ‘Sponguide’].
3.2.1. Shape. Many sponges are thought to be morphologically plastic, with individuals and populations
potentially differing widely in shape and colouration depending on a complex series of local
environmental conditions. Intraspecific genetic differences (clines) are also associated with geographic
range and populations, and thus shape (or habit) is not a particularly reliable absolute descriptive
character. However, this 'problem' is perhaps overemphasised in the literature, and in only few instances
have species been shown to be truly polymorphic. Generally species' growth forms can be defined within
reasonable limits, used with a certain degree of caution shape may be informative for particular species
determinations. The range of possible shapes seen in sponges is enormous, extenmding from thin
encrustations to massive volcano shapes, finger-like or whip-shapes, 'golf balls', fans and so on.
3.2.2. Size. The size to which particular specimens may grow may be influenced by several factors, such
as the individual's age, the prevailing environmental conditions (current, sedimentation, light
availability, etc.) and of course particular species' genetic potentials. Some species are capable of
growing into huge volcano shapes (e.g. Xestospongia) whereas other related species are merely thinly
encrusting on dead coral (e.g. Petrosia). Size is more important as a descriptive taxonomic character,
such as when comparing populations of particular species or comparing closely related (sibling) species,
and is less important as an absolute taxonomic descriptor.
3.2.3. Colour. Certain groups of sponges (such as the Verongida), have peculiar pigments that darken
upon contact with air (aerophobic pigments), and others (such as Mycalidae and Tedaniidae, order
Poecilosclerida) produce a pigmented mucus that stains or irritates human skin. Some groups of sponges
are characteristically brightly coloured (e.g. Microcionidae, order Poecilosclerida) whereas others are
typically drab (e.g. Halichondriidae, order Halichondrida). These characters are often useful for field
identications, and therefore colour notes and/or colour photographs are now considered to be essential
for accurate identification.
The range of sponge pigments is enormous, varying from drab, colourless forms (black, beige or white)
to very colourful species (vibrant reds, greens, yellows and blues, etc.). Sponge colouration can often be
attributed to the presence of particular carotenoid pigments, and because a large proportion of these
pigments is obtained and modified from the diet, mainly from the plankton, there may be some slight
variability between populations of particular species from different localities. In contrast, a few species
are truly polychromatic, with individuals, sometimes growing side by side, showing dramatic differences
in colouration. By and large, however, colouration is a useful descriptor for species identifications, and
when used cautiously colour illustrations, as presented in this book, can be useful tools for field
identifications.
As noted above, colour may be fixed to a certain extent in live material by freezing specimens prior to
preserving them, but most sponge pigments are alcohol soluble and colouration will be leached out into
the preserving fluid to a greater or lesser extent. Thus, care should be taken when preserving several
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 11
species of sponges in the same container, particularly with the aerophobic verongids that tend to dye all
other sponges a dark purple colour.
3.2.4. Texture. To an experienced field biologist sponge texture often provides good clues as to the
nature of the skeleton and water-canal system inside. A sponge which is rubbery, compressible but
difficult to tear or cut may contain no or few spicules but a well developed spongin fibre system (e.g.
Ircinia); a sponge that is soft, friable and easily torn probably has both fibres and spicules reduced (e.g.
Haliclona); one that has a hard, stony but easily crumpled texture may lack spongin fibres altogether but
have a closely compacted spicule skeleton (e.g. Petrosia); sponges incorporating sand into the skeleton
are also to a large extent brittle, easily crumbled and incompressible (e.g. Chondropsis); and sponges
that are hard, incompressible, difficult to cut or break may lack spongin fibres but have interlocking
spicules (desmas), and/or a dense surface crust of spicules (e.g. Desmanthus, Geodia). The
permuatations are diverse.
Similarly, the texture of a sponge, the degree to which it can be compressed, and whether it retains its
shape after it has been removed from the water may provide a good indication of the histology and
water-canal system (the size of choanocyte chambers, the development of the skeleton and mesohyl in
relation to the size of water-canals and choanocyte chambers, and the density of the roving cell
populations). These features are particularly useful as both field and laboratory characters for the orders
Dictyoceratida, Dendroceratida and Verongida (all of which lack a mineral skeleton).
3.2.5. Mucus production and smell. Many sponges produce mucus: usually clear, sometimes
pigmented, and in many cases toxic or irritating to the human skin. This feature is certainly characteristic
for particular species (e.g. Aplysilla sulphurea), sometimes characteristic for a particular genus (e.g.
Thorectandra), but rarely consistent at the family level (e.g. Desmacellidae, with the well known toxic
sponges Neofibularia and Biemna). Mucus production is particularly common in intertidal tropical
species and may serve a physiological role in protecting (e.g. cooling) the sponge when exposed to the
sun and air. Certainly some sponges literally drip mucus when exposed to the sun during low tides (e.g.
Clathria), but probably more importantly mucus production may protect or even repel competing
species, preditors and parasites.
With experience a field biologist may also be able to recognise particular chemical smells emitted by
particular species of sponges. Not many of these aromas have yet been documented, nor has this feature
yet been quantified, but there are several groups of species that do have unique aromas (e.g. acrid smell
of Ircinia, pungent smell of Xestospongia).
3.2.6. Surface ornamentation. The presence and distribution of surface pores, ridges, microconules,
stalks, digits, protruding spicules and other processes are often important descriptive characters, and
sometimes useful features in recognising particular genera. Small inhalant surface pores (ostia) may be
confined to one side of the sponge (inhalant surface), with the larger exhalant pores (oscules) only on
the other side (exhalant surface). This is sometimes seen in vase- or cup-shaped species (e.g.
Xestospongia). Similarly, the oscules may be scattered or aggregated into clusters (sieve-plates or
porocalyces), raised on stalks (fistules) or flat against the surface. Exhalant pores often have a
surrounding membraneous lip, which may or may not be contractile, with or without subsurface
drainage canals radiating away from the pores (astrorhizae).
Surface microconules, ridges and undulations are common features in many groups, whereas some
species have characteristic, more specialised surface processes (e.g. Myrmekioderma with polygonal
plates, producing a pineapple-like texture, and apical pore sieve-plates; Aka with large oscules on the
ends of long fistules poking through the substrate; many Clathria with astrorhizae radiating from
oscules; Callyspongia and Dysidea with a cobweb-like surface ornamentation composed of spicules or
sand, respectively).
3.2.7. Organic and inorganic skeletons. To provide a structure for the roving cell populations inside
the sponge, the small choanocyte water pumps, and the water-canals themselves there are often two type
of skeleton present, both of which are secreted by specialised sponge cells:
3.2.7.1. An organic (spongin fibre) skeleton, composed of collagen, usually forming strands. The
construction of the fibres themselves, the patterns they form, and the material contained within the fibres
are important characters used in classification.
3.2.7.2. An inorganic (spicule) mineral skeleton, found within and outside spongin fibres. Spicules are
constructed of either opaline silica or calcite, and the shape, ornamentation, size, origin and arrangement
of these spicules inside the sponge are also important characters used for classification.
3.2.8. Foreign particles. Many groups of sponges incorporate foreign particles into their mesohyl,
particularly sand particles and spicules from other sponges, but also including shell debris from
Foraminifera, Mollusca, Bryozoa, and filamentous algae. Foreign debris may be found inside spongin
12 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
fibres, actively taken into fibres by a curious exchange process whereby in some species there is a
complete loss of native spicules, replaced by debris. In other species foreign particles are found within
the proteinaceous mesohyl of the sponge but only outside fibres, or they may be restricted to the exterior
surface of the sponge only (sand cortex). There are several groups of sponges that are notorious in
being able to organise foreign particles into a 'foreign skeleton', artially or completely replacing the
'native skeleton' (e.g. Dysidea, Hyrtios, Phoriospongia, Psammoclemma, Clathriopsamma). These so-
called arenaceous sponges are usually easily detected in the field by their harsh, sandy texture.
3.2.9.Skeletal structure. Structurally the sponge may be divided into two major skeletal regions:
3.2.9.1. The outer surface of the sponge (ectosome, dermis or cortex), bounded by single (epithelial)
cells on the external surface. In some groups there may be a specialised skeleton on the surface (the
ectosomal skeleton), composed of both or either spongin fibres and spicules.
3.2.9.2. The inner region of the sponge (choanosome) includes all organic portions of the sponge inside
the epithelial cells (mesohyl, comparable to the mesenchyme of higher multicellular animals), including
the water current system. Both spicules and spongin fibres may be present in the choanosome, although
one or both may be lost in some groups. There are no true tissues, organs or coordinated nervous
systems in the Porifera, although there are documented instances of limited locomotion and coordinated
contractile responses. Traditionally the choanosomal region near the periphery is called the
subectosome.
The patterns in which the organic and inorganic skeletons grow are informative at all levels of sponge
taxonomy and generally useful in their identification. A special terminology has been produced to define
this range of skeletal structures, with several categories of skeletal architecture recognised (although
combinations and intermediate forms of these may also occur)
3.2.9.3. Branching and rejoining network (reticulate), producing regular triangular meshes (isodictyal
reticulate) or quadrangular meshes (myxillid reticulate).
3.2.9.4. Repeatedly branching but not rejoining (dendritic).
3.2.9.5. Diverging, expanding, but not branching (plumose).
3.2.9.6. Diverging, simply concentric (radial).
3.2.9.7. Disorganised criss-crossed spicule (halichondroid).
3.2.10. Spongin fibres. In several orders of sponges the mineral skeleton has been lost completely, and
for these groups fibre characteristics are important in their classification. In other groups, where there is
both spongin fibres and spicules, the latter may be partially or fully contained inside the former, and thus
the skeletal architecture is predominantly dictated by the form of the organic skeleton. In some groups
(e.g. some Haplosclerida) there are no fibres but spicules are cemented together with granular collagen.
Mostly, though, spongin fibres are useful in identification.
Spongin fibres vary both in a hierarchy of size and construction. Three size categories of fibres are
generally recognised (primary, secondary, tertiary fibres), sometimes differentiated by both size,
construction, and the material contained within each type of fibre. In addition to these fibres some
groups have collagen filaments (e.g. Ircinia), which are minute, convoluted, terminally swollen
collagenous structures dispersed within the mesohyl.
Several other classes of fibre construction are recognised, based on the amount of spongin protein
deposited when the fibre was secreted, and whether or not this spongin was deposited evenly
(homogenous fibres) or periodically (statified fibres).
Sponges with heavy spongin fibres are often termed 'horny' or 'keratose' sponges. The most simple fibres
are homogeneous in cross section without a central core (or visible pith) (e.g. Spongia), whereas the
most 'complex' fibres are stratified in cross section, composed of concentric rings of protein ('bark'),
with an optically diffuse pith in the centre of each fibre (e.g. Aplysina). Intermediate forms are also
common, such as found in species of Thorecta with slightly stratified (laminated) fibres (not bark-like),
with a granular pith.
3.2.11. Mineral skeleton. The inorganic or mineral skeleton is traditionally the most important feature
for identifying sponges. This skeleton may consist of a fused, coral-like basal skeleton and/or individual
components called spicules.
3.2.11.1. Basal skeleton. Some groups of sponges secrete a secondary, calcareous (hypercalcified),
spicular basal skeleton, in addition to free siliceous or calcitic spicules. This feature was once
considered diagnostic for a class of sponges known as 'sclerosponges', but is now interpreted as a grade
of construction found within both Calcarea and Demospongiae. The species concerned (e.g. Astrosclera)
usually live in coral reefs and their calcareous skeletons contribute to the overall accretion of these
reefs.
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 13
Megascleres:
3.2.11.2.5.A. Number of central axes (axons): monaxonic spicules with no more than two rays (points
of growth); triaxonic spicules with three perpendicular axes; tetraxonic spicules with four rays, each
with a central axis.
3.2.11.2.5.B. Number of rays (actines): monactinal spicules have one ray with asymmetrical ends (i.e.
the spicule is secreted by one or more cells commencing at one end and finishing at another); diactinal
spicules have two rays, with symmetrical ends (i.e. the spicule is secreted in both directions by one or
more cells, commencing at the centre); tetractinal spicules have more than two rays.
Microscleres:
3.2.11.2.5.C. Meniscoid or sigmoid microscleres include a diversity of curved, symmetrical and
asymmetrical spicules (chelae and sigmas).
3.2.11.2.5.D. Monaxonic microscleres include spicules with only a single axis and one or two rays.
3.2.11.2.5.E. Asterose microscleres are tetraxonic, with more than one axis and more than two rays.
3.2.12. Cytology. Several cytological characters have been instrumental in providing further
understanding of the relationships within the Porifera, particularly at higher taxonomic levels. Foremost
amongst these are choanocyte ultrastructure (including the presence, absence and position of the nucleus
within the choanocyte), and the distribution of choanocytes and shape of choanocyte chambers (e.g.
spherical, sac-like or elongate and branching). The characters have been particularly useful in describing
relationships between the 'keratose' or aspicular sponges. Other cytological characters, such as the
possession of particular cell types (e.g. cells with inclusions), have so far been found to be of limited
value, possibly because they are still documented for only few species and still poorly understood.
3.2.13. Larvae and reproductive strategy. Larval morphology is known for only relatively few
species, but in these cases it appears to be a consistent character useful for sponge taxonomy. Larval
shape (e.g. solid parenchymella, hollow amphiblastula), pattern of ciliation (e.g. partial, complete) and
mode of locomotion (e.g. swimming versus creeping) are all useful descriptive features. Mode of
reproduction, including sexual and asexual modes, has been particularly useful in developing taxonomic
hypotheses for sponges. Several reproductive characters been important in the detection of cryptic sibling
species, such as whether larvae are brooded within the parent or gametes are broadcast into seawater, and
the periodicity of spawning events. For example, sympatric populations of closely related species of
Xestospongia were found to have markedly different reproductive strategies, possibly serving as a
mechanism for niche separation (sympatric speciation). Some of these characters have also been used at
higher levels of classification, particularly oviparity versus viviparity, although our knowledge of such
strategies is still incomplete.
3.2.14. Ecology. Ecological data are virtually essential in modern sponge taxonomic descriptions,
although sadly they are lacking from most of the earlier literature that described many the known species.
These data are most useful at the species level, with proven success in being able to clearly differentiate
living populations of closely related (morphologically similar) species when it is not always possible to
do so from preserved specimens. However, it is still difficult, or impossible in some cases, to reconcile
14 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
many of the older nineteenth century species descriptions with living populations, and clearly this is one
of the major challenges facing sponge biologists for years to come.
q Note: In applying objective criteria to sponge taxonomy we have tried to quantify perhaps the
unquantifiable. Much of the process in current sponge taxonomy involves subjective interpretation of
characters and their character states, such that there are rarely hard boundaries between one character
state and the next one. The characters and states listed here are applicable mainly to demosponges, and
they are
certainly far from complete. Nevertheless, this list does provide a useful recipe to follow in describing a
sponge.
*SHOW Note that this character list is still in the 19. hollow, several tubular digits attached to
developmental stage and may be frustrating to use common
base/
*SHOW Sponge character list. Revised JNAH 28- 20. lamellate, plate-like/
NOV-1993 21. fan-shaped (flabelliform), with lamellae in one
plane/
*CHARACTER LIST 22. fan-shaped (flabelliform), with lamellae in more
than one plane/
#1. <EXTERNAL MORPHOLOGY> 23. capitate/
<COMMUNITY> sponge/ 24. club-shaped/
1. macrobenthic, fixed to the substrate/ 25. cup-shaped/
2. macrobenthic, fixed to rolling over substrate/ 26. vase-shaped/
3. endolithic, burrowing into soft substrate/ 27. spherical (golf ball)/
4. excavating, bioeroding, boring into substrate/
5. encrusting, photophilic, exposed/ #3. <EXTERNAL MORPHOLOGY> <POINT OF
6. encrusting, sciaphilic, sheltered in overhangs or ATTACHMENT> attached/
caves/ 1. directly to substrate/
7. zoophytic, overgrowing live organic substrates/ 2. directly to substrate, with enlarged basal
holdfast/
#2. <EXTERNAL MORPHOLOGY> <GROWTH 3. to substrate with basal portion burrowed into
FORM>/ sediment/
1. thinly encrusting/ 4. to substrate, insinuating into cavities/
2. thickly encrusting/
3. insinuating, boring calcitic substrates/ #4. <EXTERNAL MORPHOLOGY> <HUE>/
4. enlarged basal portion below substrate, fistules 1. /
protruding through substrate/ 1. pale/
5. lobate, spherical-bulbous/ 2. light/
6. lobate, massive/ 3. bright/
7. lobate, stoloniferous, spreading over substrate/ 4. dark/
8. simple whip-like (flagelliform), unbranched/ 5. olive/
9. arborescent, simple branching, cylindrical 6. drab/
digitate 7. mottled/
branches, few bifurcatations/ 8. deep/
10. arborescent, cylindrical digitate branches, 9. speckled/
complex 10. reddish/
branching, repeatedly bifurcate/ 11. greyish/
11. arborescent, flattened digitate branches, 12. brownish/
complex
branching, repeatedly bifurcate/ #5. <EXTERNAL MORPHOLOGY> <LIVE
12. arborescent, flattened digitate branches, COLOURATION>/
complex 1. unknown/
reticulate branching in one plane/ 2. white/
13. arborescent, flattened digitate branches, 3. beige/
complex 4. yellow/
reticulate branching in more than one plane/ 5. blue/
14. arborescent, bushy, irregular branches, thickly 6. turquiose/
branching in more than one plane/ 7. green/
15. arborescent, bushy, flattened branches, thickly 8. blue-green/
branching in more than one plane/ 9. orange/
16. tubulo-digitate, solid construction/ 10. pink/
17. hollow, single tubular digit/ 11. red/
18. hollow, bifurcate tubular digits/ 12. red-orange/
13. red-brown/
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 15
7. membraneous, hispid, with erect spicules from 8. compressed, with more-or-less disorganized
choanosomal skeleton protruding through surface/ paratangential tracts of spicules lying below
8. unispicular, isotropic, single spicules lying ectosome/
tangential to surface/ 9. scattered tangential tracts of spicules lying below
9. unispicular, isodictyal, with single spicules lying ectosome/
tangential to surface/ 10. single or paucispicular isodictyal tracts/
10. uni- or paucispicular, isodictyal tracts of 11. cavernous, with wide-meshed reticulate tracts
spicules lying tangential to surface/ of
11. uni- or paucispicular, radial tracts of spicules spicules lying below ectosome/
perpendicular to surface/ 12. plumo-reticulate, with diverging and
12. paucispicular, with sparse bundles of spicules anastomosing
paratangential or perpendicular to surface/ tracts of spicules supporting ectosome/
13. pauci- or multispicular, with discrete bundles of 13. plumose, with diverging brushes of spicules
spicules standing perpendicular spicules/ supporting ectosome/
14. multispicular, with a continuous palisade of 14. radial, with perpendicular bundles of spicules
spicules perpendicular to surface/ supporting ectosome/
15. multispicular, with a thick crust with spicules 15. radial, with single spicules perpendicular to axis
tangential or paratangential to surface/ protruding through ectosome/
16. membraneous, but with a felt of microscleres/
#16. <SKELETON> <SUBECTOSOMAL
#14. <SKELETON> <ECTOSOMAL SPECIALISATION>/
SPECIALISATION>/ 1. /
1. / 2. composed of undifferentiated choanosomal
2. composed of undifferentiated choanosomal spicules/
spicules/ 3. composed of undifferentiated ectosomal
3. composed of undifferentiated subectosomal spicules/
spicules/ 4. composed of special category of auxiliary
4. composed of special category of ectosomal spicules,
spicules geometrically different from choanosomal spicules/
scattered individually on surface/ 5. composed of special category of auxiliary
5. composed of special category of ectosomal spicules,
spicules differing from ectosomal spicules only in
forming erect bundles on surface/ dimensions/
6. composed of special category of ectosomal 6. composed of special category of auxiliary
spicules spicules,
forming continuous palisade on surface/ differing from choanosomal spicules only in
7. composed of dense crust of subectosomal dimensions/
spicules, 7. composed of echinating spicules concentrated in
tangential or paratangential to surface/ peripheral skeleton/
8. composed of acanthostyles in plumose brushes 8. supplemented by sand grains and other foreign
around particles dispersed throughout peripheral skeleton/
protruding subectosomal spicules/
9. raspailiid, with bundles of ectosomal auxiliary #17. <SKELETON> <CHOANOSOMAL
spicules surrounding larger protruding ARCHITECTURE>/
subectosomal 1. collagenous/
spicules/ 2. hymedesmoid/
10. raspailiid, with plumose bundles of 3. microcionid/
choanosomal 4. plumose/
spicules surrounding larger protruding spicules/ 5. plumo-reticulate/
11. composed of bundles of raphides dispersed 6. irregularly reticulate/
over 7. regularly reticulate/
ectosome and surrounding larger protruding 8. renieroid-subisodictyal reticulate/
spicules/ 9. isodictyal reticulate/
10. isotropic reticulate/
#15. <SKELETON> <SUBECTOSOMAL 11. disorganised halichondroid reticulate/
SKELETON>/
1. / #18. <SKELETON> <CHOANOSOMAL AXIAL
2. undifferentiated from choanosomal skeleton/ AND EXTRA-AXIAL
3. collagenous, lacking any region skeleton/ DIFFERENTIATION> with/
4. compressed, with fibres forming a more close- 1. /
meshed 2. undifferentiated axial or extra-axial regions/
reticulation in periphery than in axial skeleton/ 3. wide-meshed relatively homogeneous fibres/
5. cavernous, with peripheral fibres less 4. wide-meshed skeletal tracts/
compressed 4. compressed basal fibres, and radial extra-axial
than in axial skeleton/ skeleton/
6. vestigial, with spicules sparsely dispersed 5. compressed basal fibres and plumose extra-axial
throughout peripheral skeleton/ skeleton/
7. compressed, with more-or-less disorganized 6. compressed basal skeleton composed of short
tangential tracts of spicules lying below ectosome/ bulbous
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 17
fibre nodes, cored by plumose tufts of radiating 13. only primary (ascending) skeletal tracts present,
extra- cored by principal spicules, without connecting
axial spicules/ tracts/
7. differentiated, compressed, reticulate axis, and 14. skeletal tracts in axial skeleton cored by
plumo-reticulate extra-axis/ auxiliary spicules, whereas tracts in peripheral region
8. differentiated, compressed, reticulate axis, and wholly arenaceous/
plumose extra-axis/ 15. primary spongin fibres interconnected by
9. differentiated, compressed, reticulate axis, and smaller
radial extra-axis/ secondary fibres/
10. differentiated plumose axis, and radial extra- 16. without spongin fibres or spicule tracts,
axis/ although
11. mostly non-anastomosing fibres and spicule collagen fibrils present/
tracts/
12. differentiated primary (plumose-dendritic) and #20. <SKELETON> <CHOANOSOMAL
secondary (renieroid-subrenieroid) skeletons/ SPICULES> choanosomal
13. differentiated primary multispicular skeleton and spicules/
secondary renieroid skeleton/ 1. /
14. radially dispersed single spicules or 2. absent/
paucispicular 3. completely enclosed in spongin fibres/
tracts of spicules throughout/ 4. core spongin fibres as well as echinate fibre
15. a criss-cross of spicules, vaguely recognisable endings, protruding through fibres in "spicate"
in arrangement/
structure only at the periphery/ 5. echinates spongin fibres as well as form plumose
16. -out apparent skeletal organization/ ascending brushes in peripheral skeleton/
6. echinate fibre nodes/
#19. <SKELETON> <CHOANOSOMAL 7. strewn in loosely aggregated tracts within
SKELETAL TRACTS>/ mesohyl/
1. / 8. strewn in halichondroid tracts within mesohyl/
2. basal fibres aspiculose, although bases of 9. form secondary renieroid skeleton, without a
spicules fibre
embedded in spongin fibres, standing perpendicular component, bound at nodes by collagen/
to 10. form a rigid, interlocking skeleton/
substrate/
3. homogeneous unispicular skeletal tracts cored by #21. <SKELETON> <CHOANOSOMAL
choanosomal spicules/ ECHINATING SPICULES>
4. homogeneous uni- or paucispicular skeletal tracts echinating spicules/
cored by choanosomal spicules/ 1. /
5. homogeneous multispicular skeletal tracts cored 2. absent (presumed lost)/
by 3. absent, although second category of larger,
choanosomal spicules, occupying only proportion acanthose choanosomal spicule present/
of fibre 4. absent, although cladotylotes echinate skeletal
diameter/ tracts/
6. homogeneous multispicular skeletal tracts cored 5. clumped on basal spongin/
by 6. concentrated in tufts at fibre nodes/
choanosomal spicules, occupying entire fibre 7. concentrated on exterior edges of skeletal tracts/
diameter/ 8. concentrated on primary skeletal tracts/
7. homogeneous multispicular skeletal tracts cored 9. confined to peripheral skeleton, forming plumose
by brushes on ectosome, surrounding protruding
auxiliary spicules identical to those in peripheral subectosomal
skeleton/ spicules/
8. homogeneous multispicular skeletal tracts cored 10. clumped at junction of axial and extra-axial
by skeletons, around bases of subectosomal spicules/
choanosomal spicules with echinating spicules 11. secondarily incorporated into fibres/
secondarily incorporated into fibres/ 12. form a dense, rigid, interlocking secondary
9. larger primary (ascending) and smaller secondary skeleton/
(transverse) skeletal tracts cored by principal 13. forming plumose or subrenieroid tracts only/
spicules/ 14. sparse, evenly distributed over skeletal tracts/
10. larger primary (ascending) skeletal tracts cored 15. heavy, evenly distributed over skeletal tracts/
by 16. supplemented by cladotylotes also echinating
principal spicules, smaller secondary (connecting) skeletal tracts/
tracts cored by auxiliary spicules/
11. larger primary (ascending) skeletal tracts cored #22. <SKELETON> <CHOANOSOMAL
by SPONGIN FIBRES> spongin
principal spicules, smaller secondary (connecting) fibres/
tracts aspiculose/ 1. /
12. larger primary (ascending) skeletal tracts cored 2. absent/
by 3. absent, spicule tracts bound together at their
auxiliary spicules, smaller secondary (connecting) nodes by collagen/
tracts aspiculose/ 4. poorly developed/
18 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
2. With at least some free triradiate spicules ........................................................ [Subclass Calcinea] ....
(3)
Spicules are free and sagittal tetracts and monaxonic forms ......................... [Subclass Calcaronea] ...
(4)
Most spicules are fused ('hypercalcified'), often with tuning fork spicules included (polyphyletic
character) .................................................. [Orders Murrayonida (Calcinea) and Lithonida
(Calcaronea)].
3. Spicules may include triradiate and quadriradiate forms periphery of skeleton has a distinct cortex
..................................................................................................... [Order Clathrinida, family Leucettidae].
Only triradiate spicules present, body plan asconoid ....................... [Order Clathrinida, other
families].
5. Birotulate microscleres present, hexaster microscleres absent, sponges not attached to substrate but
embedded within it on 1 or more long basal spicules …………… ………………………… ………..
………………………………………..………….. [Subclass Amphidiscophora, Order
Amphidiscosida].
Hexaster microscleres present, birotulate microscleres absent, sponges usually fixed to substrate
................................................................................................................. [Subclass Hexasterophora] .....
(6)
6. Parenchymal skeleton consists of fused hexactine spicules forming rigid skeleton ………. ………..
……….. ……………………………………………………………………..……... [Order
Hexactinosida].
Parenchymal spicules are lychniscs united together, with centre of each spicule surrounded by 12 struts
................................................................................................................................... [Order Lychniscosida].
Parenchymal spicules consist of hexactines usually free within syncytial (acellular) matrix, and with
specialized ectosomal hexactines or pentactines with longest ray pointing inwards ……….. ……….
………. …………………………………………………………………………….. [Order
Lyssacinosida].
7. Skeleton composed of tetraxonid spicules and derivatives with equal rays, megascleres and
microscleres undifferentiated, (sometimes spicules are lost completely and sponge may be superficially
confused with compound ascidians) ................................... [Subclass Homoscleromorpha, Order
Homosclerophorida].
Tetraxonid and monaxonid megascleres often occur together, asterose microscleres common, skeleton
is usually radial or axially compressed .................................................. [Subclass Tetractinomorpha] .....
(8).
Monaxonic megascleres, with a diversity of microscleres but never asterose forms, (two order lacking
free spicules altogether) ...................................................................... [Subclass Ceractinomorpha] .....
(9).
8. Spherical growth form usual, radial pattern of triaenes and oxeas, microscleres sigmaspires
.................................................................................................................................... [Order Spirophorida].
Large oxeas always present, sometimes with triaenes, radial at surface only, microscleres asterose
forms ................................................................................................................................... [Order
Astrophorida].
Monaxonic spicules only (styles, oxeas, never tetractinal forms), radial at least at surface, microscleres
may be absent or may include asterose and monaxonic forms (microxeas, spirasters) .………… ……….
………………………………………………………………………………......... [Order
Hadromerida].
Articulated siliceous desma megascleres, with or without free spicules ………….. ………. ……….
……………………………………………………………………….... [Order Lithistida
(polyphyletic)].
11. Lacking mineral skeleton completely (although detritus and contaminating spicules often occur,
confusing these with poecilosclerids), with well developed relatively homogeneous spongin fibres
forming reticulate skeleton, typically with 2 or 3 different sized networks, consistency not collagenous
.................................................................................................................................. [Order Dictyoceratida].
Spongin fibres forming reticulate skeleton, with laminated spongin fibres, with distinct pith of fine
fibrils, forming reticulate skeleton without differentiation of primary and secondary elements,
collagenous consistency, frequently with a live yellow colouration which darkens in contact with the air
......................................................................................................................................... [Order Verongida].
With strongly lamellated spongin fibres forming dendritic skeleton arising from basal attachment
................................................................................................................................ [Order Dendroceratida].
Without free spicules but with a solid aragonitic cortex producing a series of chambers on top of each
other, the youngest (uppermost) chambers lined with living tissue ……………….... [Order
Verticillitida].
q THE SUBPHYLA
Two subphyla are now recognised: Cellularia including the classes Demospongiae and Calcarea, and
Symplasia including Hexactinellida. Although not explicit by any author the extinct Archaeocyatha
would also be grouped within the Cellularia.
q THE CLASSES
The Demospongiae contain about 95% of living species, with a described fauna already consisting of
about 4500-5000 species and an estimated total extant fauna of between 14000-15000 species
worldwide. Within this class there are three subclasses, 13 orders (1 dubious), 71 families and 1005
nominal genera included, although only 507 genera are presently considered to be valid. 481 genera
include marine species and 26 genera concern freshwater species. The number of genera recognised as
valid varies between different authors and the whole classification is not yet stable.
The Calcarea contains two subclasses, 5 orders, 18 families, 98 nominal genera (63 of which are
apparently valid), and an estimated fauna of between 400-500 species worldwide. All species are marine.
The Hexactinellida are divided into 2 subclasses, 4 orders, 19 families, 113 nominal genera [of which
101 are currently recognised but many may be synonyms], and an estimated 450-500 species worldwide.
The Archaeocyatha (extinct) were an important group of marine sessile organisms, limited to the
Cambrian. They have been attributed to various phyla and even considered as an independent one. Recent
studies of comparative anatomy show similarities in growth pattern, structure of the skeleton (primary
or secondary), functional morphology and trends of evolution clearly demonstrate that they have a
sponge grade of organization and form a Class within the Phylum Porifera. Recent studies of immune
responses and peculiar budding types further suggest affinities with Demosponges. Archaeocytha contain
6 orders (based on the number of walls and types of development), 12 suborders (based on the intervallar
elements of the first order), 50 superfamilies (based on the outer wall primary elements), 124 families
(based on the inner wall primary elements, and 306 valid genera.
q PHYLUM PORIFERA.
q SUBPHYLUM CELLULARIA.
q CLASS DEMOSPONGIAE (Siliceous sponges).
DEFINITION: Sponges with the skeleton composed of spongin fibres alone or together with siliceous
spicules (although some "relict sclerosponge" forms have both a basal calcitic skeleton as well as free
siliceous spicules). Some groups lack a mineral skeleton entirely (some Homoscleromorpha and others).
Collagenous filaments or fibrils (forming the ground substance of the intercellular mesohyl) are
ubiquitous, spongin fibres (also composed of collagen) occur in most families, and histological
organisation is always cellular (as opposed to syncytial in the Hexactinellida). Choanocytes occupy
chambers that are spherical, hemispherical, elongate or branched.
REMARKS. Three subclasses of Demospongiae are distinguished on the basis of larval morphology and
life cycle strategy (Homoscleromorpha, Tetractinomorpha, Ceractinomorpha), and a fourth
24 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
(polyphyletic) group, the "sclerosponges" with calcified basal skeletons, is now distributed amongst the
existing various families of demosponges. The `sphinctozoa' are also now included in Demospongiae,
subclass Ceractinomorpha. There are 13 orders (1 dubious), 71 families and 1005 nominal genera
included, although only 507 genera are presently considered to be valid. 481 genera include marine
species and 26 genera concern freshwater species (the latter not included in this work).
Demospongiae contain about 95% of living species, with a described fauna already consisting
of about 4500-5000 species and an estimated total extant fauna of between 14000-15000 species
worldwide.
q SUBCLASS HOMOSCLEROMORPHA
DEFINITION: Demospongiae with secondarily derived amphiblastula larvae and viviparous
reproduction; skeleton composed of tetraxonid siliceous spicules and derivatives with equal rays (diods,
triods, lophate spicules), arranged around choanocyte chambers reflecting the canal structure; no
differentiation between megascleres and microscleres although size differences do occur between types
of spicules; spicules usually small (100µm or less), not localised to any particular region; choanocyte
chambers with large numbers of choanocytes. One order, one family.
q ORDER HOMOSCLEROPHORIDA
DEFINITION: As for subclass.
varying abundance; deformed calthrops in some species and some with diactine-derived microscleres
dispersed regularly throughout the mesohyl (Diaz & van Soest, 1994).
Oscarella Vosmaer, 1884 (type species: Halisarca lobularis Schmidt, 1862) (syn. Oscaria,
sensu Vosmaer, 1880; Octavella Tuzet & Paris, 1963) - encrusting, lobulate smooth surface; no
spicules, no fibres; choanocyte chambers eurypylous (Boury-Esnault et al., 1984, 1992; Sole-Cava et al.,
1992; Diaz & van Soest, 1994).
q SUBCLASS TETRACTINOMORPHA
DEFINITION: Demospongiae with parenchymellae or creeping blastula larvae, predominantly oviparous
reproduction although in some genera young sponges are incubated within parent and set free as small
adults; megascleres tetraxonid and monaxonid, occurring together or separately; microscleres asterose
forms and derivatives; skeletal structure usually radial or axially compressed.
REMARKS. Three orders of Tetractinomorpha are well established (Astrophorida (also known as
Choristida), Hadromerida, and Spirophorida), and a fourth polyphyletic order ("Lithistida") shows major
affinities to, and will probably be merged eventually in, Hadromerida.
q ORDER SPIROPHORIDA
DEFINITION: Typically with spherical growth form, with tetraxonid and monaxonid megascleres
(triaenes, oxeas), in radiate pattern; protriaenes most common and often protrude from surface;
monocrepidial desmas may be present; microscleres contorted microspined sigmaspires (an apomorphy
for the group); reproduction oviparous without a larval stage, or viviparous with production of young
adults within parent.
A single family (Tetillidae) is traditionally included in this order, but the "lithistid" family
Scleritodermidae also has the apomorphic sigmaspire microsclere and is also best included here.
Tetilla Schmidt, 1868 (type species: Tetilla euplocamos Schmidt, 1868) (syn. Chrotella
Sollas, 1886; Psetalia Gray) - without porocalices, without cortex, without unusual accessory
megascleres.
Ancorina Schmidt, 1862 (type species: Ancorina cerebrum Schmidt, 1862) (syn.
Stellettinopsis Carter, 1879; Algol Sollas, 1888; Thalassomora Lendenfeld, 1888; Sanidastrella
Topsent, 1892) - surface microrhabds and sanidasters.
Cryptosyringa Vacelet, 1979.
Ecionemia Bowerbank, 1863 (type species: Ecionemia acervus Bowerbank, 1863) -
microstrongyles and euasters, without oxyasters.
Holoxea Topsent, 1892 (type species: Holoxea furtiva Topsent, 1892a) - massive or encrusting
growth forms; choanosome consists of an irregular halichondroid main skeleton of horizontally
disposed, larger oxeas, without axial and extra-axial differentiation, although extra-axial skeleton may
consist of larger oxeas in loose tracts, protruding through the surface; ectosome with a smaller category
of oxeas erect on the surface. Microscleres are minute sanidaster-like microxeas, and raphides occurring
singly or in trichodragmata.
Melophlus Thiele, 1899 (type species: Melophlus sarasinorum Thiele, 1899) - with spiny
microrhabds (not true sanidasters)
Monosyringa Brondsted, 1924 (type species: Monosyringa mortenseni Brondsted, 1924).
Myriastra Sollas, 1886 (type species: Myriastra subtilis Sollas, 1886) - 1 category of aster.
Penares Gray, 1867 (type species: Stelletta helleri Schmidt, 1864) - spiculation of
microrhabds abundantly distributed in cortex and external parts of choanosome (Ref. Desqueyroux-
Faundez & Van Soest, 1997).
Psammastra Sollas, 1886 (type species: Psammastra murrayi Sollas, 1886).
Rhabdastrella Thiele, 1903 (type species: Coppatias distinctus Thiele, 1900) (syn. Diastra
Row, 1911; Aurorella de Laubenfels, 1957; Aurora, sensu Sollas, 1888) - triaenes absent or vestigial.
Stelletta Schmidt, 1862 (type species: Stelletta grubii Schmidt, 1862) (syn. Myriastra Sollas,
1886; Pilochrota Sollas, 1886; Anthastra Sollas, 1886) - 2-3 categories of asters distributed on both
surface and choanosome, or only present in choanosome; cortex well-developed (Ref. Desqueyroux-
Faundez & Van Soest, 1997).
Stryphnus Sollas, 1886 (type species: Stryphnus niger Sollas, 1886).
Calthropella Sollas, 1888 (type species: ) (syn. [Corticella] Sollas, 1888 [preocc.];
Corticellopsis Bergquist, 1968) - without oxeas.
Chelotropaena Lendenfeld (type species: ) - with calthrops, long shafted dichotriaenes and
microscleres streptasters and microxeas.
Chelotropella Lendenfeld, 1906 (type species: ) - with oxeas and peripheral dichotriaenes.
Pachastrissa Lendenfeld, 1903 (type species: ) - with oxeas.
Pachataxa de Laubenfels, 1936 (type species: ).
DEFINITION: Massive sponges with diplodal aquiferous systems; megascleres small, minutely spined
triaenes (plagio-, dicho- and trichotriaenes with trifurcate clads), organised in ectosomal region with
long shaft directed inwards and small clads tangential to surface, disorganised in choanosomal skeleton;
microscleres amphiasters.
SCOPE: This group is poorly known, with one established genus and one incertae sedis.
REVIEWS: Wilson (1925), Hallmann (1912).
GENERA:
Thrombus Sollas, 1886 (type species: ).
Trachygellius Topsent, 1894 (type species: Trachya globosa Carter, 1886).
q ORDER HADROMERIDA
DEFINITION: Relatively cohesive order with uniform spiculation of monaxonid megascleres
(monactinal or diactinal;); with radially arranged skeleton always obvious at surface if not within
choanosome; spongin fibres poorly developed (if at all present); ectosomal spicules typically smaller
than choanosomal spicules, usually standing perpendicular to surface and protruding through ectosome;
microscleres, if present, euasters, streptasters and derivatives, spirasters or spiraster-like spirules, or
peculiar asterose-like discorhabds; all groups oviparous (where known), with development of
parenchymella larva (in one case blastula larva) directly in seawater.
Twelve families presently included, although Chondrillidae was recently elevated to full ordinal
status by Boury-Esnault & Lopes, 1985 (change not yet incorporated here).
spined ends that can develop into a more spiraster-like shape; 2. Smooth nodulose amphiasters, which
can occasionally be absent even in specimens of the same species (Rosell & Uriz, 1997).
Dotona Carter, 1880 (type species Dotona pulchella Carter, 1880) – Excavating Clionidae
always in alpha stage, with styles in two size classes, occasionally modified to subtylostyles or
strongyles) and oxeas as megascleres. Two types of microscleres: straight relatively small amphiasters
and spiny microstongyles with blunt spines arranged in a spiral ridge. Small styles arranged in palisade in
the papilllae. Megascleres scattered throughout the choanosome. Amphiasters located in the papillae and
spiral-spined microstrongyles widely distributed through the whole sponge (Rosell & Uriz, 1997).
Pione Gray, 1867 (type species Cliona northumbrica Hancock, 1849) – Excavating Clionidae,
developing in an alpha or beta stage, with 2 spicule types of megascleres – tylostyles and oxeas. Spined,
occasionally smooth oxeas showing intraspecific variability in abundance. Oxeas often with central
swelling (centrotylote). Tylostyles in palisade make the skeleton papillae. Scarse tylostyles and abundant
oxeas form the confused skeletal arrangement of the choanosome. Microscleres are present as straight or
undulated, occasionally centrotylote, spined microrhabds. The latter are characterised by the uniform
distribution of spines along the whole shaft, which is truncated at the ends (Rosell & Uriz, 1997).
Scantilletta de Laubenfels, 1936 (type species: Scantilla spiralis Johnson, 1899) (syn.
[Scantilla] Johnson, 1899; Donotella de Laubenfels, 1936) –Excavating Clionidae exclusively
displaying the alpha stage of growth. Smooth oxea in 2 separated size classes as megascleres. The
shortest size class of oxea is always arranged in the palisade in the papillae. Sometimes the palisade is
reinforced by some large choanosomal oxeas. In the choanosome megascleres not arranged in any clear
pattern. Microscleres are amphiasters and spiny or smooth vermiform microstrongyles (Rosell & Uriz,
1997).
Spheciospongia Marshall, 1892 (type species Alcyonium vesparium Lamarck, 1814) (syn.
Heterocliona Verrill, 1907)– Massive Clionidae with robust tylostyles in great abundance forming a
tangential network of indistinct strands or felted without orientation and minute spirasters in small
numbers concentrated in the ectosome and the canal linings; tylostyles commonly modified to styles,
tylostrongyles and strongyles; excavating limestone substrates during early stage of life history (Vicente,
Ruetzler & Carballeira, 1991).
Thoosa Hancock, 1849 (type species: Thoosa cactoides Hancock, 1849) (syn. Thoosa de
Laubenfels, 1936) - megascleres with centrotylote swelling from which sometimes 2-3 pointed shafts
radiate; microscleres peculiar astrose forms.
Volzia Rosell & Uriz, 1997 (type species Cliona albicans Volz, 1939) – Excavating Clionidae
exclusively displaying the alpha stage of growth. Smooth oxeas and tylostyles as megascleres. In the
papillae, oxeas are always arranged in palisade. Tylostyles confusedly arranged are exlusively present in
the choanosome. Microscleres have not been observed.
separate (i.e. the junior name Chondrosiidae has not necessarily 'won general acceptance'), so Article
40b cannot be used in this case. Recommendation 40a recommends that when citing an 'established'
family name (i.e. the name may not necessarily be the most senior but comes under Article 40b) then the
revised author name and date should be included after the name too. Thus, family Chondrillidae Gray,
1872 (Schmidt, 1862) is the senior-most available name for this taxon.
GENERA:
Chondrilla Schmidt, 1862 (type species: Chondrilla nucula Schmidt, 1862) (syn.
Chondrillastra Topsent, 1918) - with ectosomal euasters.
Chondrosia Nardo, 1847 (type species: Chondrosia reniformis Nardo, 1847) (Gummina
Schmidt, 1862; Cellulophana Schmidt) - without spicules.
Thymosia Topsent, 1895 (type species: Thymosia guernei Topsent, 1895).
Thymosiopsis Vacelet & Perez, 1997 (type species Thymosiopsis cuticulatus Vacelet & Perez,
1997) -
cortex of considerable depth. In some species the base of the cortex, just above the cortex-choanosome
boundary, can have heavy collagen deposition. In some species long whip-like subtylostyles or digitate
umbrelliform exotyles echinate the sponge surface. Megascleres are strongyloxeas, subtylostyles, and
tylostyles to which smooth centrotylote oxeas can be added. <I></I>Polymastia<I></I> species
reproduce asexually by the production of stalked bead-likee buds and sexually by the production of
parenchymella larvae following extrusion of eggs and sperm..
Proteleia Dendy & Ridley, 1886 (type species: Proteleia sollasi Dendy and Ridley, 1886).
Quasillina Norman, 1869 (type species: Euplectella brevis Bowerbank, 1861) - massive, with
or without stalk; cortex present or absent; choanosomal skeleton consists only of a subectosomal system
of longitudinal and circumferential tracts; megascleres subtylostyles and strongyloxeas in 2-3 sizes
(Ref. Desqueyroux-Faundez & Van Soest, 1997).
Radiella Schmidt, 1870 (type species: Radiella sol Schmidt, 1870) (syn. Trichostemma Sars,
1872) - disk-shaped with a fringe of spicules.
Sphaerotylus Topsent, 1898 (type species: Polymastia capitata Vosmaer, 1885).
Spinularia Gray, 1868 (type species: ) (syn. Rhaphidorus Topsent, 1898).
Tentorina Burton, 1959 (type species: Tentorina sigmatophora Burton, 1959) - skeleton of
strongyloxeas sparsely present in choanosome and forming continuous tangential ectosomal layer;
microscleres acanthose 'sigmaspirule-like' microxeas.
Tentorium Vosmaer, 1885 (type species: Thecophora semisuberites Schmidt, 1870) (syn.
Thecophora sensu Schmidt, 1870 [preocc.]).
Tylexocladus Topsent, 1898 (type species: ).
Weberella Vosmaer, 1885 (type species: ).
Stylocordyla Thomson, 1873 (type species: Hyalonema boreale Loven, 1868) (syn.
Oxycordyla Topsent, 1904; Microcordyla Zirpolo; Stylorhiza Schmidt) - abyssal, on stalk.
Tethycordyla de Laubenfels, 1934 (type species: Tethycordyla thyris de Laubenfels, 1934) -
bathyal, on stalk, with asters.
predominantly in surface brushes; tylostyles with irregularly shaped heads, wrinkled, lumpy, constricted,
flattened or lobed due to swelling or branching of axial filament; commonly associated with symbiotic
bacteria or algae.
q Suborder Triaenosina
DEFINITION: Peripheral skeleton of radially arranged triaene megascleres, with amphiaster, spiraster or
microrhabd microscleres; with obvious affinities to Hadromerida.
Three families are included.
Theonella Gray, 1868 (type species: Theonella swinhoei Gray, 1868) - phyllotriaenes and
microrhabds.
q Suborder Rhabdosina.
DEFINITION: Ectosomal megascleres absent but ectosome contains minutely spined microstrongyles,
microspined sigma-like microscleres or monocrepidial (one-rayed) disks; affinities with Hadromerida.
Two families are included.
q Suborder Anoplina
DEFINITION: Both ectosomal megascleres and microscleres absent; affinities uncertain.
Three families are included.
q SUBCLASS CERACTINOMORPHA
DEFINITION: Sponges with parenchymella larvae and viviparous sexual reproduction (although with
several oviparous `enclaves': Agelasida, Petrosiidae, Axinellidae, Desmoxyidae); generally with both
spicule skeleton with well developed spongin fibres forming a diversity of skeletal structures (although
siliceous spicules lost altogether in 3 orders, and spongin fibres lost in several genera); spicules
monaxonic (either monactinal (styles) or diactinal (oxeas-strongyles)), never tetractinal (although
modifications to the ends of some monaxonic spicules occur); microscleres diverse (meniscoid, oxeote,
toxote, spheres) but never asterose.
Eight orders are differentiated here, although some authors also recognise a ninth (Petrosida),
based on the possession of oviparous sexual reproductive strategy, now widely included in the
Haplosclerida.
q ORDER VERTICILLITIDA
DEFINITION: DEFINITION: Demospongiae with `sphinctozoan' grade of construction (solid aragonitic
cortex producing a series of chambers on top of each other); living `sphinctozoans' lack free spicules but
have cells and larvae resembling those of other Demospongiae.
One fossil genus, previously associated with the 'sphinctozoa', has calcareous monaxon and
triradiate spicules enclosed within the walls of the chambers. This late Mesozoic genus (Barroisia) bears
monaxonid and triradiate calcareous spicules enclosed within the walls of the chambers, but its inclusion
in this group is now debatable. Conversely, clues on the phylogeny of this group can be seen from the
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 39
Recent genus Vaceletia, which 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"
probably represent independent but convergent lines of the two classes Calcarea and Demospongiae, for
fossil and living species respectively. The alternative hypothesis, that all "Sphinctozoa" are Calcarea -
with their identity revealed by the calcareous spicules of the genus Barroisia - whereby the line
containing the Recent genus Vaceletia is merely convergent upon the Demosponges in cellular and
developmental characters - is rejected, as is a third possibility is that the "Sphinctozoa" represent a
completely independent class of sponges.
Only one extant order, family and genus (Vaceletia) are known, although many more fossil taxa
described.
q ORDER AGELASIDA
DEFINITION: Oviparous sponges, showing (perhaps superficial) resemblance to commercial bath
sponges (Spongiidae) and biochemical similarities to Axinellidae; growth forms branching, tubular, fan-
shaped or massive; well developed spongin-fibre skeleton, forming regular or irregular reticulation;
fibres echinated by short styles or oxeas with verticillate spines; microscleres absent.
Represented by two Recent families.
DEFINITION: Bulbous, encrusting or massive subspherical growth forms; basal skeleton composed of
aragonite, spherulitic in form (with each spherulite laid down in a cell at the surface of the sponge and is
eventually led to a position where it contributes to the general reticulate skeleton of aragonite);
intracellular secretion of sclerodermites only found in Astroscelera, whereas in other genera the skeleton
is secreted extracellularly; living tissue penetrates reticulation to a depth of about 1cm, but no tabulae
seperate the tissue-filled external parts of the skeleton from the interior that is of free living tissue;
interior interskeletal spaces generally fill in with secondary deposits of aragonite; siliceous spicules
verticillately spined acanthostyles, sometimes secondarily lost.
SCOPE: Five genera are included.
REVIEWS: Hartman (1969), Vacelet (1981), Wood (1990)
GENERA:
Astrosclera Lister, 1900 - massive or columnar; basal calcareous skeleton with spherulitic
microstructure, together with free spicules consisting of verticillate acanthostyles.
Ceratoporella Hickson, 1912 - massive, dome shaped; basal aragonitic skeleton with
clinogonal microstructure, together with free spicules consisting of verticillate acanthostyles.
Goreauiella Hartman, 1969 - pedunculate and auriculiform;; basal calcareous skeleton
together with free spicules consisting of verticillate acanthostyles.
Hispidopetra Hartman, 1969 - encrusting or massive; basal calcareous skeleton together with
free spicules consisting of verticillate acanthostyles and free styles.
Stromatospongia Hartman, 1969 - encrusting on serpulid tubes or massive mammilliform;
basal calcareous skeleton together with free spicules consisting of verticillate acanthostyles.
q ORDER POECILOSCLERIDA
DEFINITION: Skeleton with discrete siliceous spicules, although some primitive groups retain a fused
basal calcitic skeleton or a fused siliceous (desmoid) skeleton, along with free siliceous skeletons; main
skeleton composed of megascleres (monactinal, diactinal or both) and spongin fibres in various stages of
development; megascleres frequently localised to distinct regions; microscleres include meniscoid forms
such as chelae (unique to the order) and sigmas, and other diverse forms (toxas, raphides, microxeas);
most families are viviparous, with uniformly ciliated parenchymella having bare posterior poles
(although Raspailiidae is oviparous).
REMARKS. This order contains more living species than all other Recent Porifera, and includes both
marine and some freshwater species. Up to 25 families have been recognised in this order, most being
typical in having chelae microscleres, several atypical in lacking these microscleres, but a recent
proposed reorganisation of the order, based on phylogenetic parsimony analysis, suggests that only 19 of
these may be valid.
tylotes and palmate isochelae, but these were traditionally included in Myxillidae (e.g. Topsent, 1928; de
Laubenfels, 1936), which predominantly contain taxa with ‘tridentate-derived’ chelae (arcuate and
anchorate), sigmas, but lack toxas. Hajdu et al. (1994) proposed to merge these two groups of genera by
extending the scope of Cornulidae to include both fistulose and non-fistulose genera with similar
skeletal arrangements (ectosomal tylotes with spined bases) and spiculation (palmate isochelae, toxas,
lack of sigmas). These authors also correctly noted that the inclusion of Iophon in this group
necessitated recognition of Burton's (1929) higher taxon Iophoneae, which they elevated to full family
status. Iophoneae was initially created only for Iophon and Iophonopsis, as a ‘Section’ under the family
Myxillidae. Under Hadju et al.'s (1994) concept of Iophonidae 35 nominal genera were potentially
allocated here. There are several recent reviews outlining the scope and features of this group, including
comprehensive revisions of particular genera (Van Soest et al., 1991; Hajdu et al., 1993; Van Soest, Zea
& Kielman, 1994; Desqueyroux-Faundez & Van Soest, 1996).
REVIEWS: Berquist & Fromont (1988), Van Soest et al. (1994), Desqueyroux-Faundez & Van Soest
(1996).
GENERA:
Acanthorhabdus Burton, 1929 (type species: Acanthorhabdus fragilis Burton, 1929) -
Massive non-fistulose growth form; ectosomal skeleton composed of a loose halichondroid tangential
layer of acanthoxeas (acanthorhabds) with heavy spines and microspined tylote apices; choanosomal
skeleton with smooth modified styles (anisoxeas) often provided with a terminal mucron, forming
massive multispicular plumose and reticulate columns, with acanthoxeas dispersed between (but not
echinating) fibres; microscleres palmate anisochelae with spurs. References: Burton (1929:432),
Desqueyroux-Faundez & Van Soest (1995).
Acarnus Gray, 1867 (type species: Acarnus innominatus Gray, 1867) (syn. Fonteia Gray,
1867; Trefortia Deszo, 1880; Microtylotella Dendy, 1896; Acanthacarnus Lévi, 1952) - Non-fistulose,
encrusting to massive growth forms; ectosomal tylotes with microspined ends, forming tangential or
paratangential tracts (not brushes); choanosomal skeleton isodictyal, isotropic or anisotropic reticulate,
occassionally plumo-reticulate or plumose or hymedesmioid in encrusting species; skeletal tracts cored
by smooth styles, echinated by 1 or 2 (exceptionally 3) categories of cladotylotes, echinating
acanthostyles present or absent; microscleres are palmate isochelae and toxas of several morphologies.
References: van Soest (1984:63); Lévi (1952); Hooper (1987); van Soest et al. (1991:49); Hiemstra &
Hooper (1991:431).
Acheliderma Topsent, 1892 (type species: Acheliderma lemmiscata Topsent, 1892) (syn.
Astylinifer Topsent, 1927; Fusifer Dendy, 1896) -Fistulose growth forms; ectosomal spicules are tylotes
with microspined bases lying tangential to surface (replaced by sand in one species); choanosomal
skeleton an irregular reticulation of microspined styles and echinating acanthostyles; microscleres
characteristically elongated diamond-shaped microxeas and toxas; chelae absent. Reference: Topsent
(1892:24; 1928:228-9), Dendy (1896:49).
Cornulella Dendy, 1922 (type species: Cornulella lundbecki Dendy, 1922) - Thinly
encrusting, fistular growth forms; ectosomal skeleton a unispicular network of intercrossing tangential
tylotes with spined heads; choanosomal skeleton vestigial, made up of individual tylotes identical to
those in the ectosomal skeleotn, with accessory acanthoxeas distributed singly among the tylotes;
microscleres palmate isochelae and frequently toxas. References: Van Soest, Zea & Kielman, 1994.
Cornulum Carter, 1876 (type species: Cornulum textile Carter, 1876) (syn. Cornulotrocha
Topsent, 1927; Coelosphaerella de Laubenfels, 1936; Melonchela Koltun, 1955; Heterocornulum Lévi
& Lévi, 1983) - Massive or encrusting growth forms, and with erect fistules or pronounced pore areas;
ectosomal skeleton consists of a compact crust of spicule layers orientated tangential to the surface,
composed of smooth tylotes/strongyles with microspined apices; choanosomal skeleton consists of
tracts and scattered tylotes/strongyles as found on the ectosome, together with monactinal spicules
(styles) with microspined bases present in some species; echinating acanthostyles in one species only;
microscleres palmate isochelae, toxas, and microstrongyles in 1 species. References: Lundbeck
(1910:22); Bergquist & Fromont (1988:53); Carter (1876:309), Koltun (1955:17; 1959:187); Hooper
(MS), Lévi & Lévi (1983:966).
Damiria Keller, 1891 (type species: Damiria simplex Keller, 1891) (syn. Anisotylacanthaea
Vacelet, 1969; [Damira]; de Laubenfels (lapsus); Xytopsene de Laubenfels, 1936) -Fistular growth
form; ectosomal skeleton with tangential crust of tylotes with microspined apcies; choanosomal skeleton
irregularly reticulate with tracts of larger tylotes of similar morphology as ectosomal spicules; no other
megascleres; microscleres may include toxas. References: van Soest (1984:73), Topsent (1928:325), de
Laubenfels (1936:54); Carter (1880), Vacelet, 1969, Van Soest, Zea & Kielman, 1994.
Dolichacantha Hentschel, 1914:118 (type species: Dolichacantha macrodon Hentschel,
1914) -Encrusting to massive growth forms; ectosomal skeleton with long acanthostrongyles
intermingled with tylotornotes arranged in bundles on the surface; choanosomal skeleton hymedesmioid
with entirely spined acanthostyles and basally- or vestigially entirely spined subtylostyles embedded in
42 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
the basal skeleton, standing perpendicular to the substrate, and bundles of tylotornotes and/or
acanthostrongyles forming tracts more-or-less ascending through the surface; microscleres include only
uncommon palmate isochelae. References: Hentschel (1914), Koltun (1970).
Iophon Gray, 1867 (type species: Halichondria scandens Bowerbank, 1866) (syn. Alebion,
sensu Gray, 1867; Menyllus Gray, 1867; Ingallia Gray, 1867; Pocillon Topsent, 1893; Iophonopsis
Dendy, 1924; Burtonella de Laubenfels, 1928; Iophonota de Laubenfels, 1936; Hymedesanisochela
Bakus, 1966) -Non-fistulose, massive, branching or encrusting growth forms; ectosomal skeleton
composed of tylotes with microspined bases, lying tangential to surface, intercrossing and in erect
brushes; choanosomal skeleton an isodictyal rounded, triangular or square-meshed reticulation of
smooth or spined choanosomal styles, arranged singly or in 2-3 per row, with (s.l.) or without (s.s.)
echinating acanthostyles; microscleres inequiended bipocilla and palmate anisochelae with spurs, toxas
absent. References: Bowerbank (1866:259); Bergquist & Fromont (1988:90); Bakus (1966:479);
Desqueyroux-Faundez & Van Soest (1996).
Megaciella Hallmann, 1920 (type species: Amphilectus pilosus Ridley and Dendy, 1886) (syn.
Holorodesmia Topsent, 1928; Myxichela de Laubenfels, 1936) - Non-fistulose, branching, lobate,
flabellate or encrusting growth forms; ectosomal skeleton composed of tangential or paratangential layer
of ectosomal tylotes with microspined bases, often sinuous or curved; choanosomal skeleton reticulate
or confused (hymedesmioid in encrusting species), with ascending multi- and paucispicular tracts of
smooth choanosomal styles with spined bases, interconnected by secondary uni- or bispicular tracts
bound together with collagen producing irregular wide meshes; echinating acanthostyles present or
absent; microscleres are palmate isochelae and two morphologies of toxas. References: Ridley & Dendy
(1886:125, 350); Hallmann (1920:772); Desqueyroux-Faundez & Van Soest (1996); Hooper (MS,
from redescription of type), de Laubenfels (1936:85); Wilson (1925:432).
Paracornulum Hallmann, 1920 (type species: Cornulum dubium Hertschel, 1912) (syn.
‘Cornulacantha’ Lévi & Lévi, 1983 [nomen nudum]) -Encrusting-fistular growth form, with ectosomal
skeleton composed of intercrossing halichondroid tracts of larger tylotes with granular or microspined
apices, and choanosomal skeleton composed of similar but slightly smaller spicules forming radial or
plumose tracts arising from a hymedesmioid layer of basal spongin echinated by acanthostyles
perpendicular to the substrate; microscleres are palmate isochelae and toxas. References: Hallmann
(1920:772); Bergquist & Fromont (1988:54); Hentschel (1912:346).
Tedaniphorbas de Laubenfels, 1936 (type species: Amphilectus ceratosus Ridley & Dendy,
1886) - Massive, lobate growth form; ectosomal skeleton detachable, composed of scattered smooth
tylotes, singly or in bundles, forming tangential skeleton; choanosomal skeleton well developed,
reticulate, composed of very horny spongin fibres lacking coring or echinating spicules, but with
ectosomal tylotes scattered between fibres and aggregated into loose wispy tracts near the periphery;
microscleres are palmate isochelae; toxas absent. References: nil.
Zyzzya de Laubenfels, 1936 (type species: Plocamia massalis Dendy, 1922) (syn. Damirina
Burton, 1959; Histodermopsis de Laubenfels, 1936) - Massive, burrowing, fistulose or cryptic
encrusting growth form with solid apical fistules or blind papillae; ectosomal skeleton thick detachable
crust of tangentially orientated tylotes with microspined bases; choanosomal skeleton with distinctive
irregular or plumose multispicular, widely spaced tracts ascending to the surface cored by the same
tylotes, interdispersed with a unispicular, irregular isodictyal reticulation of verticillately-spined
strongyles (or strongyloxeas); microscleres if present are palmate isochelae and toxas. References:
Hooper & Krasochin (1989:); Bergquist & Fromont (1988:56); Dendy (1922:78); de Laubenfels
(1936:72); Lundbeck (1910:16); Van Soest et al. (1994).
these have been merged into well established taxa such that only 8 genera and 14 subgenera are now
recognised.
REVIEWS: Hallmann (1920), Hentschel (1923), Lévi (1960), Simpson (1968), van Soest (1984);
Bergquist & Fromont (1988), Hooper (1988b, 1989a, 1990, 1996), Hooper & Lévi (1993), Hooper et
al. (1990, 1992); Hooper & Wiedenmayer (1994); Howson & Chambers (1999).
GENERA:
Antho Gray, 1867 (type species: Myxilla involvens Schmidt, 1864) (syn. Acarnia Gray, 1867;
Anomoclathria Topsent, 1929; Anthoarcuata Bakus, 1966; Brondstedia Burton, 19??; Clathriella
Burton, 1935; Dictyoclathria Topsent, 1920; Dirrhopalum Ridley, in Ridley and Duncan, 1881;
Echinoplocamia Burton, 1959; Heteroclathria Topsent, 1904; Holoplocamia de Laubenfels, 1936;
Isociona Hallmann, 1920; Isopenectya Hallmann, 1920; Jia de Laubenfels, 1930; Lissoplocamia
Brondsted, 1924; ? Naviculina Gray, 1867; Plocamia Schmidt, 1870; Plocamilla Topsent, 1928;
Plocamiopsis Topsent, 1904; Quizciona de Laubenfels, 1936) - ectosomal skeleton with tangential or
erect-plumose auxiliary styles, choanosomal skeleton a basal or axial renieroid reticulation composed of
acanthostyles, acanthostrongyles or smooth "dumbell spicules", with a secondary dendritic, plumose,
plumo-reticulate or simply echinating skeleton of smooth choanosomal styles arising from fibre nodes
(erect forms), or ascending upwards from basal spongin (encrusting forms), echinating acanthostyles
present or absent, microscleres include palmate isochelae, including modified (cleistochelae) and more
contort forms, pseudoarcuate isochelae, modified sigmoid isochelae (crocae), and smooth or spined
toxas of several forms.
Three subgenera of Antho:
Antho (Antho) (type species: Myxilla involvens Schmidt, 1864) - with basal renieroid skeleton
composed predominantly of acanthostyles, without echinating acanthostyles.
Antho (Acarnia) (type species: Hymeniacidon cliftoni Schmidt, 1870) - with basal renieroid skeleton
composed predominantly of acanthostrongyles or smooth "dumbell spicules", echinating
acanthostyles present.
Antho (Isopenectya) (type species: Clathria ? chartacea Whitelegge, 1907) - axially compressed
renieroid skeleton composed of acanthose choanosomal styles, extra-axially subisodictyal
reticulation composed of smooth choanosomal styles, a plumose subectosomal peripheral
skeleton composed of smooth choanosomal styles, and a tangential ectosomal skeleton
composed of auxiliary styles; echinating acanthostyles absent.
Artemisina Vosmaer, 1885 (type species: Artemisina suberitoides Vosmaer, 1885) (syn. ?
Qasimella Thomas, 1974) - choanosomal skeleton confused halichondroid tracts, vaguely ascending,
composed of principal styles without definite spongin fibres, extra-axial skeleton radially arranged with
1 category of auxiliary style forming paratangtial bundles on the surface, echinating spicules absent,
microscleres are palmate isochelae and toxas with spined points.
Echinochalina Thiele, 1903 (type species: Ophlitaspongia australiensis Ridley, 1884) (syn.
Tablis de Laubenfels, 1936; Protophlitaspongia Burton, 1934) - ectosomal skeleton paratangential,
plumose, choanosomal skeleton more-or-less regularly reticulate, both composed of the same auxiliary
styles, quasidiactinal spicules, or diactinal spicules, and choanosomal fibres echinated by smooth
(sometimes acanthose) principal styles or diactinal spicules, microscleres may include palmate isochelae
and toxas.
Two subgenera of Echinochalina:
Echinoclathria Carter, 1885 (type species: Echinoclathria tenuis Carter, 1885) (syn.
Ophlitaspongia, of authors [not Ophlitaspongia Bowerbank, 1866) - encrusting, arborescent,
branching or fan-shapes, with compressed axial (or basal) skeleton and plumo-reticulate or plumose
extra-axial skeleton.
Holopsamma Carter, 1885 (type species: Holopsamma crassa Carter, 1885) (syn. Plectispa
Lendenfeld, 1888; Aulena, in part (e.g. Lendenfeld, 1888); [Halme] Lendenfeld, 1885 [preocc.],
Echinoclathria of authors) - specialized "honeycomb" reticulate growth forms, with more-or-less
evenly reticulate fibres, without any compression of axial skeleton or any structural differentiation
between axial and extra-axial regions.
REVIEWS: Bergquist (1970), Hooper (1991), Hooper & Lévi (1993), Hooper, Lenhert & Zea (1999).
GENERA:
Amphinomia Hooper, 1991 (type species: Amphinomia sulphurea Hooper, 1991) - massive
and lobate growth forms, choanosomal skeleton renieroid-reticulate, without axial condensation,
composed of light spongin fibres cored by modified styles (with large spines on both base and point, but
smooth in between), echinated by unmodified acanthostyles, with sparse extra-axial smooth styles
scattered throughout the mesohyl, without specialized raspailiid ectosomal skeleton; microscleres
absent. References: Hooper (1991).
Aulospongus Norman, 1878 (type species: Haliphysema tubulatus Bowerbank, 1873) (syn.
Aulospongiella Burton, 1956; Heterectya Hallmann, 1917; Rhaphidectyon Topsent, 1927;
Hemectyonilla Burton, 1959) - cup-shaped, lobate, and cylindrical growth forms, composed of fused,
shaggy fibre bundles, choanosomal axial skeleton not condensed, composed of long, plumose, partially
fused spongin fibres cored by smooth rhabdostyles, diverging in the periphery, ectosome without any
specialized spicules or skeletal structure; echinating acanthostyles have smooth rhabdose bases with
spined points, or sometimes they are entirely spined; raphide microscleres present or absent. References:
Hooper (1991), Hooper et al. (1998).
Axechina Hentschel, 1912 (type species: Axechina raspailioides Hentschel, 1912) - bushy
growth forms, choanosomal skeleton with 2 components - a condensed tightly reticulate axis cored by
oxeas with spined points, and plumose columns of smooth anisoxeas ascending to the surface; extra-
axial skeleton radial, composed of long styles embedded in plumose tracts of oxeas and projecting
through the surface, specialized raspailiid ectosomal skeleton composed of peculiarly curved or sinuous
toxiform styles with spined ends; echinating spicules absent; microscleres absent. References: Hooper
(1991).
? Cantabrina Ferrer-Hernandez, 1914 [incertae sedis] (type species: Cantabrina erecta
Ferrer-Hernandez, 1914) (Ectyonilla Ferrer-Hernandez; de Laubenfels, 1936) - lobate growth form,
choanosomal skeleton halichondroid, without spongin fibres, composed of a criss-cross of long styles,
extra-axial skeleton more-or-less plumose composed of long styles projecting through the surface,
without specialized raspailiid ectosomal skeleton; echinating spicules rare, smooth styles with rhabdose
bases; microscleres absent. References: Hooper (1991).
Ceratopsion Strand, 1924 (type species: Ceratopsis expansa Thiele, 1898) (syn. Ceratopsis,
sensu Thiele, 1898 [preocc.]) - lamellate and branching growth forms, choanosomal skeleton axially
condensed reticulation of spongin fibres cored by sinuous or straight styles or anisoxeas, rarely
strongyles, extra-axial skeleton radial composed of individual long styles or anisoxeas, sinuous or
straight, rarely strongyles, standing perpendicular to axis and projecting through the surface, specialized
ectosomal skeleton composed of thin oxeas or styles; echinating spicules absent; microscleres absent.
References: Hooper (1991).
Cyamon Gray, 1867 (type species: Dictyocylindrus vickersii Bowerbank, 1866) - encrusting or
massive growth forms, choanosome is a basal layer of spongin fibre with microcionid fibre-nodes and
individual or plumose tracts of long thick styles standing erect, projecting through the surface, extra-
axial skeleton with few long thin styles also projecting, without any specialized ectosomal skeleton;
modified acanthostyles (sagittal tetracts or pentacts) usually form a rigid secondary skeleton;
microscleres absent. References: Hooper (1991).
Echinodictyum Ridley, 1881 (type species: Spongia bilamellata Lamarck, 1814, var.ß) (syn.
Dictyocylindrus, in part Carter, 1879; Kalykenteron Lendenfeld, 1888; Kieplitela de Laubenfels, 1954)
- vase, cup, fan shaped and branching growth forms, choanosomal skeleton simply reticulate, without
axial compression or differentiation between the axis and extra-axial skeletons, composed of spongin
fibres cored by oxeas; short or long extra-axial styles are scattered throughout the mesohyl, sometimes
projecting through the surface, typically without a specialized raspailiid ectosomal skeleton (although
present in 1 species, composed of small styles); fibres echinated by unmodified acanthostyles, with
spination varying from sharply pointed erect, sharply pointed recurved, spatuliferous with even margins,
and spatuliferous with serrated margins; microscleres absent. References: Hooper (1991).
Ectyoplasia Topsent, 1930 (type species: Spongia tabula Lamarck, 1814) - fan-shaped or
flattened branching growth forms, choanosomal axial skeleton composed of slightly condensed spongin
fibres cored by styles or anisoxeas, extra-axial skeleton with radially arranged plumose tracts composed
of the same choanosomal spicules, each interconnected by smaller tracts of the same spicules together
producing a regular reticulation, specialized raspailiid ectosomal skeleton composed of small styles or
anisoxeas; spongin fibres echinated by clavulate acanthostyles; microscleres absent. References: Hooper
(1991).
Endectyon Topsent, 1920 (type species: Phakellia tenax Schmidt, 1870) (syn. Hemectyon
Topsent, 1920; Basiectyon Vacelet, 1961) - branching growth form, choanosomal axial skeleton with
46 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
condensed fibres cored by thick styles, extra-axial skeleton radial or plumose tracts of long projecting
styles, specialized raspailiid ectosomal skeleton composed of thin styles; spongin fibres echinated by
acanthostyles or acanthostrongyles bearing long, recurved hooks, and echinating spicules
characteristically localized to particular places within the skeleton (i.e. on outer fibres grouped around
projecting spicules; at the base of the sponge; or along the length of the extra-axial fibres); microscleres
absent. References: Hooper (1991).
Eurypon Gray, 1867 (type species: Hymeraphia clavata Bowerbank, 1866) (syn. Epicles Gray,
1867; Acantheurypon Topsent, 1927; Tricheurypon Topsent, 1928; Protoraspailia Burton & Rao,
1932; Proraspailia Lévi 1958) - encrusting and digitate growth forms, choanosomal skeleton consists
of basally or axially condensed spongin fibres, with swollen fibre nodes (i.e. microcionid skeleton),
cored by erect, individual or plumose bundles of long, smooth or partially spined, extra-axial styles, with
thin styles or oxeas forming a specialized raspailiid ectosomal skeleton; basal fibre-nodes (encrusting
forms) or plumose extra-axial skeletal tracts (branching forms), echinated by unmodified acanthostyles;
microscleres may include raphides. References: Hooper (1991).
Hymeraphia Bowerbank, 1864 (type species: Hymeraphia stellifera Bowerbank, 1864) (syn.
Mesapos Gray, 1867) - encrusting growth form, choanosomal skeleton consists of a basal layer of
spongin lying on the substrate, without specialized choanosomal spicules, but with single or brushes of
echinating acanthostyles and projecting extra-axial styles standing erect on the substrate, without a
specialized raspailiid ectosomal skeleton; echinating acanthostyles modified to forms with bulbous-
swollen, or stellate-swollen heads and spined points; microscleres absent. References: Hooper (1991).
Lithoplocamia Dendy, 1922 (type species: Lithoplocamia lithistoides Dendym 1922) (syn.
Monectyon Lévi & Vacelet, 1958) - encrusting and massive growth forms, choanosomal skeleton is a
dense isodictyal or irregularly subisodictyal reticulation of acanthostrongyles, in 1 or 2 size categories,
without axial condensation, without echinating acanthostyles, with extra-axial radial tracts of smooth
styles, and typically without a specialized raspailiid ectosomal skeleton (although when present
ectosomal spicules are long slender oxeas); microscleres absent. References: Hooper (1991).
Plocamione Topsent, 1927 (type species: Plocamione dirrhopalina Topsent, 1927) (syn.
Raspeloplocamia Burton, 1935; Axoplocamia Burton, 1935; [Plocamia], in part, Schmidt, 1870) -
encrusting and digitate growth forms, choanosomal skeleton consists of a basally condensed (encrusting
specimens) or axially condensed (digitate specimens) reticulation of specialized acanthostrongyles ("peg-
top" spicules), echinated by acanthostyles, with an extra-axial radial or plumose skeleton composed of
long styles in columns, and a specialized raspailiid ectosomal skeleton composed of thin styles or
anisoxeas; echinating acanthostyles are unmodified, although they may vary from entirely smooth to
partially spined, with smooth, partially spined or tuberculate bases; microscleres absent. References:
Hooper (1991).
Raspaciona Topsent, 1936 (type species: Halichondria aculeata Johnston, 1842) - lobate and
branching growth forms, choanosomal skeleton consists of basally condensed spongin fibres with
plumose tracts of long projecting styles, each surrounded by brushes of smaller styles producing the
specialized raspailiid ectosomal skeleton; unmodified echinating acanthostyles, often of more than one
size category, concentrated around the bases of plumose skeletal columns; microscleres absent.
References: Hooper (1991).
Raspailia Nardo, 1833 (type species: Raspailia viminalis Schmidt, 1862) (syn. Abila, sensu
Gray, 1867; Abilana Strand, 1928; Axinectya Hallmann, 1917; Clathriodendron Lendenfeld, 1888;
Dictyocylindrus Bowerbank, 1859; Echinaxia Hallmann, 1916; Parasyringella Topsent, 1928;
Raspailopsis Burton, 1959; Raspaxilla Topsent, 1913; [Raspeila] Nardo, 1833 (nomen nudum);
Syringella, of authors; Valedictyum de Laubenfels, 1936) - branching, lobate and massive growth
forms, choanosomal axial skeleton typically condensed (subgenus Raspailia and others), or simply
open-reticulate (subgenus Clathriodendron), composed of a reticulation of spongin fibres cored by
smaller styles or oxeas, with a plumose, radial or plumo-reticulate extra-axial skeleton cored by long
projecting styles or oxeas, and much smaller styles or oxeas producing specialized raspailiid ectosomal
skeleton; fibres typically echinated by conical, club-shaped acanthostyles (subgenus Raspailia), modified
acanthose rhabdostyles (subgenus Raspaxilla), modified acanthostyles with smooth swollen-tylote bases
(subgenus Hymeraphiopsis), or lacking echinating acanthostyles altogether (subgenus Syringella);
microscleres absent. References: Hooper (1991).
Five subgenera established:
Raspailia (Raspailia), Raspailia (Clathriodendron), Raspailia (Raspaxilla), Raspailia
(Hymeraphiopsis), Raspailia (Syringella).
Rhabdeurypon Vacelet, 1969 (type species: Rhabdeurypon spinosum Vacelet, 1969) -
encrusting growth form, choanosomal skeleton consists of a basally condensed layer of spongin lying on
the substrate (hymedesmoid skeleton), without choanosomal spicules or true echinating spicules, but
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 47
with long extra-axial styles embedded in basal spongin and projecting through the surface, with smaller
oxeas producing a specialized raspailiid ectosomal skeleton; "echinating" acanthostyles modified to
acanthorhabds, scattered throughout the mesohyl; microscleres are microxeas. References: Hooper
(1991).
? Sigmeurypon Topsent, 1928 [incertae sedis] (type species: Microciona fascispiculiferum
Carter, 1880) - encrusting growth form, choanosomal skeleton with basally condensed spongin fibres
lying on the substrate, without choanosomal spicules, with extra-axial, long, smooth styles erect and
projecting through the surface, without specialized raspailiid ectosomal skeleton but with bundles of
raphides lying tangential to the surface; echinating acanthostyles are unmodified and erect on the
substrate; microscleres are raphides. References: Hooper (1991).
Thrinacophora Ridley, 1885 (type species: Thrinacophora funiformis Ridley and Dendy,
1886) - encrusting, massive, and branching growth forms, choanosomal skeleton composed of axially
condensed fibres forming a dense, reticulate core, cored by short, stout oxeas, sometimes anisoxeas or
styles, extra-axial fibres radial or plumose, cored by long, thick styles or anisoxeas, specialized raspailiid
ectosomal skeleton composed of styles, which may be modified; echinating spicules absent; microscleres
raphides. References: Hooper (1991).
Trikentrion Ehlers, 1870 (type species: Spongia muricata Esper, 1793) (syn. Plectronella
Sollas, 1879) - branching growth form, choanosomal axial skeleton a wide-meshed reticulation of
oxeas, with no or little axial compression, extra-axial skeleton consists of plumo-reticulate fibres cored
by the same oxeas with long projecting styles embedded in the peripheral fibres, each surrounded by
brushes of thin styles forming the specialized raspailiid ectosomal skeleton; modified pseudo-tetractinal
spined spicules (sagittal triacts) echinate fibres, usually in light concentrations; microscleres raphides.
References: Hooper (1991).
Acanthodoryx Lévi, 1961 (type species: Acanthodoryx fibrosa Lévi, 1961) - massive;
ectosomal skeleton with tangential or paratangential tylotes and isochelae; choanosomal skeleton
composed of acanthostyles forming plumose skeletal tracts; microscleres arcuate isochelae, no sigmas.
Reference: Lévi, 1961:516.
Anomodoryx Burton, 1934 (type species: Desmacidon dendyi Whitelegge, 1901) - massive;
ectosomal skeleton membraneous, porous, composed of scattered strongyles of the same form as found
in the choanosomal skeleton; choanosomal skeleton plumose or plumoreticulate, composed of whispy
slender primary fibres, cored by strongyles and moderate spongin, and less well developed secondary
fibres, which together branch and anastomose to form a radial fibre system running vertically to the
surface; microscleres are arcuate isochelae and sigmas. References: Burton (1934:554); Whitelegge
(1902:79).
Camptisocale Topsent, 1904 (type species: ) - smaller smooth ectosomal styles, larger
choanosomal smooth styles, microscleres are arcuate isochelae.
Chaetodoryx Topsent, 1927 (type species: ) (syn Coelectys Topsent, 1936) - smooth polytylote
ectosomal tylotes, choanosomal styles with microspined bases, echinating acanthostyles present,
microscleres are arcuate isochelae and rugose microxeas (similar to onychaetes).
Coelocarteria Burton, 1934 (type species: Phloeodictyon singaporense Carter, 1883) (syn.
Ichnodonax de Laubenfels, 1954) - cylindrical burrowing base, erect fistules; ectosomal skeleton
consists of thick compact layers of both sizes of strongyles orientated tangentially; choanosomal
skeleton dense, composed of tracts of long and small strongyles, the larger sometimes modified to
strongyloxeas, and usually with faint microspined bases; microscleres are palmate isochelae. References:
Burton (1934:563); Carter (1883:326); Ridley (1884:421); Bergquist & Fromont (1988:55).
Coelodischela Vacelet, Vasseur & Lévi, 1976 (type species: Coelodischela diatomorpha
Vacelet, Vasseur & Lévi, 1976) - small tubes fixed directly to the substrate on a common base;
ectosomal skeleton reinforced by tangential strongyles, unjoined, single or in bundles, and concentric
spicules surrounding pores on the surface; choanosomal skeleton reticulate composed of weak
multispicular tracts and by fewer tylotes dispersed throughout the skeleton; megascleres are strongyles
and tylotes; microscleres are coelodiscs (bipocilli-like isochelae) and sigmas. References: Vacelet,
Vasseur & Lévi (1976:57); Lévi & Lévi (1983:957).
Coelosphaera Thomson, 1873 (type species: Coelosphaera tubifex Thomson, 1873) (syn.
Coelosphaericon Bakus, 1966; Histoderma Carter, 1874; Histioderma Carter, 1886; Sideroderma,
sensu Ridley & Dendy, 1886; Siderodermella Dendy, 1921; Xytopsoocho de Laubenfels, 1936) -
massive, bladder-like, or encrusting, burrowing growth forms with erect fistules; smooth ectosomal
tylotes form a compact tangential crust; choanosomal skeleton consists of poorly developed tracts and
scattered smooth spicules; megascleres are smooth tylotes or strongyles, of one size only; microscleres
are arcuate isochelae, sigmas and raphides (often in trichodragmata), some of which may be absent.
References: Bergquist & Fromont (1988:47); van Soest (1984:71).
Damiriella Burton, 1935 (type species: ) - smooth ectosomal tylotes, smooth choanosomal
strongyles, microscleres are unguiferous (arcuate) isochelae (with sharp alae); no sigmas (Ref. Hofman
& Van Soest, 1995: 77-103).
Ectyodoryx Lundbeck, 1909 (type species: Hastatus foliatus Fristedt, 1887) - ectosomal
skeleton composed of tornotes or tylotes, lying tangentially and also forming paratangential brushes;
choanosomal skeleton composed of smooth or spined styles forming an isodictyal or square-meshed
reticulation, echinating acanthostyles present; microscleres are arcuate isochelae and sometimes sigmas.
References: Bergquist & Fromont (1988); Hentschel (1911).
Forcepia Carter, 1874 (type species: Forcepia colonensis Carter, 1874) (syn. Ectoforcepia
Cabioch, 1968; Forcepina Vosmaer, 1885; Trachyforcepia Topsent, 1904) - encrusting to massive
growth forms; ectosomal skeleton composed of tangential smooth ectosomal tylotes; choanosomal
skeleton composed of choanosomal styles (Forcepia) or ectosomal tylotes (Ectyoforcepia) forming
hymesmoid structure in encrusting species or reticulate architecture in massive forms; microscleres are
smooth or spined forceps (labis), arcuate isochelae, palmate isochelea, sigmas. References: Carter
(1874); Dendy (1895); van Soest (1984:66).
Histodermella Lundbeck, 1910 (type species: Histodermella ingolfi Lundbeck, 1910) (syn.
Hiltonus de Laubenfels, 1936) - spherical base with erect fistules; ectosomal skeleton consists of
smooth ectosomal tylotes (or strongyles) forming a thick tangential layer; choanosomal skeleton consists
of irregular tracts of scattered tylotes (or strongyles), and acanthoxeas or acanthostrongyles form a layer
at right angles to the ectosomal tylotes and are scattered throughout the choanosome; microscleres are
arcuate isochelae and sigmas, sometimes also raphides in trichodragmata. References: Lundbeck
(1910:13); Bergquist & Fromont (1988:49)
50 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
Histodermion Topsent, 1927 (type species: ) - palmate isochelae, smooth ectosomal tylotes,
hymedesmoid architecture with acanthostyles erect on substrate, microscleres are arcuate isochelae.
Inflatella Schmidt, 1875 (type species: Inflatella pellicula Schmidt, 1875) (syn. Joyeuxia
Topsent, 1890) - massive, spherical with erect fistules; ectosomal skeleton a compact tangential crust of
diactinal spicules; choanosomal skeleton consists of thin tracts of scattered diactinal spicules, the same
as on the surface; diactinal megascleres are of a single sort, being smooth ectosomal strongyles or
anisotylotes; microscleres absent. References: Bergquist & Fromont (1988:51).
Lepidosphaera Lévi & Lévi, 1979 (type species: Lepidosphaera hindei Lévi & Lévi, 1979) -
basal mass burrowing with erect fistules protruding through substrate; ectosomal skeleton consists of a
rigid cortex composed of peripheral layer of platelet-like discs, "ecailles" (? desmas); choanosomal
skeleton pulpy with smooth tylotes dispersed throughout; microscleres absent. References: Lévi & Lévi
(1979:)
Leptolabis Topsent, 1904 (type species: Dendoryx luciensis Topsent, 1888) (syn. Clinolabis
Topsent, 1927; Labisophlita de Laubenfels, 1936; Marcusoldia de Laubenfels, 1936) - encrusting;
ectosomal skeleton with bundles of tylotes; choanosomal skeleton with erect acanthostyles; microscleres
arcuate isochelae, sigmas and forceps. References: Hentschel (1911:71); Topsent (1904:182)
Lissodendoryx Topsent, 1892 (type species: Tedania leptoderma Topsent, 1889) (syn.
Damiriana de Laubenfels, 1950; Jones de Laubenfels, 1936; Paramyxilla Dendy, 1905;
Waldoschmittia de Laubenfels, 1936; Xytopsihis de Laubenfels, 1936; Zetekispongia de Laubenfels,
1936; Zetekopsis de Laubenfels, 1936; Zottea de Laubenfels, 1936) - encrusting to massive with uneven
surface and soft consistency; ectosomal skeleton with smooth ectosomal tylotes forming tangential
tracts and surface brushes; choanosomal skeleton composed of smooth or acanthose choanosomal
monactinal or diactinal spicules (styles, sometimes oxeas or strongyles), forming a renieroid or
isodictyal reticulate architecture of single spicules; no echinating spicules; microscleres are arcuate
isochelae, sigmas, and raphides present in trichodragmata or absent. References: Bergquist & Fromont
(1988:85); van Soest (1984:54); Bakus (1966:491); Zea (1987:160); Hofman & Van Soest (1995:77).
Manawa Bergquist & Fromont, 1988 (type species: Pyloderma demonstrans Dendy, 1924) -
spherical shape with fistulose inhalant and exhalant surface papillae; ectosomal skeleton is a compact
tangential crust of oxeas; choanosomal skeleton consists of coarse irregular, poorly defined tracts of
oxeas, also dispersed throughout the mesoyly; microscleres are arcuate isochelae. References: Bergquist
& Fromont (1988:52); Dendy (1924:370).
Naauna de Laubenfels, 1950 (type species: ) - strongyles scattered individually throughout
skeleton, microscleres are arcuate isochelae and raphides in trichodragmata.
Phelloderma Ridley & Dendy, 1886 (type species: Phelloderma radiatum Ridley and Dendy,
1886) - globular growth form; distinct, thick, detachable cortical ectosome with points of choanosomal
styles protruding; choanosomal skeleton radial, ascending to the surface, composed of smooth polytylote
styles; microscleres are modified arcuate isochelae (thaumatochelae). References: Ridley & Dendy
(1886:113; 1887).
Pyloderma Kirkpatrick, 1908 (type species: Halichondria latrunculioides Ridley and Dendy,
1886) - erect fistules and raised pore areas; ectosomal skeleton parchment-like, detachable, composed of
dense tangential slender oxeas arranged side-by-side; choanosomal skeleton with a loose irregular
reticulation of fibres cored by the same slender oxeas, and also scattered throughout the mesohyl;
megascleres oxeas only; microscleres are tridentate (arcuate) isochelae. References: de Laubenfels
(1936:72); Ridley & Dendy (1886:326; 1887:6); Dendy (1924:370).
Ysila de Laubenfels, 1936 (type species: ) - no ectosomal skeleton, long smooth styles in
choanosomal fibres, microscleres are arcuate isochelae and sigmas.
and ascending tracts of smooth diactinal spicules; megascleres acanthostyles and/or acanthostrongyles,
and smooth strongyles; microscleres absent. References: Boury-Esnault & van Beveren (1982:75).
Crellomima Rezvoj, 1932 (type species: ) (syn. Damonilla de Laubenfels, 1936) - ectosomal
acanthostyles or acanthostrongyles, oxeote tornotes in subectosomal skeleton, microscleres are
unguiferous isochelae.
Grayella Carter, 1869 (type species: ) - no isochelae, no basal acanthostyles.
Naniupi de Laubenfels, 1950 (type species: Naniupi ula de Laubenfels, 1950) - ectosomal
skeleton composed of a crust of acanthostyles or acanthoxeas orientated vertically or tangentially;
choanosomal skeleton with a principal skeleton composed of plumose tracts of smooth styles which may
be echinated by acanthostyles; base of sponge may have an erect layer of acanthostyles also; microscleres
are arcuate isochelae. References: de Laubenfels (1950:19); Bergquist & Fromont (1988:81).
Phlyctaenopora Topsent, 1904 (type species: Phlyctaenopora bitorquis Topsent, 1904) -
massive, with erect fistules; tangential ectosomal skeleton of smooth strongyles; confused compact
choanosomal skeleton of smooth oxeas and scattered bundles of individual strongyles; microscleres are
anisochelae with spurs, sigmas and spined rhabds. References: Topsent (1904:198); Wiedenmayer
(1989:38); Lévi & Lévi (1983:957).
Pseudoclathria Dendy, 1897 (type species: Halichondria ? compressa Carter, 1886) -
massive compact growth form; ectosomal skeleton with tangential layer of acanthostyles; choanosomal
skeleton reticulate composed of smooth styles cemented together by spongin, and many acanthostyles
scattered throughout the mesohyl (but not echinating fibres); microscleres absent. References: Burton
(1929:441); Dendy (1897:258).
Pytheas Topsent, 1890 (type species: Pytheas atra Topsent, 1890) - encrusting; ectosomal
skeleton composed of a dense crust of smaller acanthostyles; choanosomal skeleton composed of criss-
crossed strongyles, with erect basal acanthostyles larger than those on the surface; microscleres are
arcuate isochelae. References: Boury-Esnault & van Beveren (1982:73); Topsent (1928:91); Hentschel
(1914:95).
Yvesia Topsent, 1890 (type species: Halichondria albula Bowerbark, 1864) (syn. Pytheilla de
Laubenfels, 1936; Tisrone de Laubenfels, 1936; Yvesiella de Laubenfels, 1936; Yvesiorbas de
Laubenfels, 1936) - pedunculate, ovoid mass; ectosomal skeleton with a crust of acanthostyles or
acanthostrongyles; choanosomal skeleton composed of smooth tornotes in erect bundles; microscleres
are tridentate (arcuate) isochelae. References: Burton (1929:441); Topsent (1890:10; 1892:102).
birotulate isochelae and sigmas. References: Bergquist & Fromont (1988:50); van Soest (1984:76);
Carter (1886:447)
Amphilectus Vosmaer, 1880 (type species: Isodictya gracilis Bowerbank, 1866) - stalked,
dichotomously branching; ectosome membraneous, without specialized spiculation; choanosomal
skeleton reticulate cored by multispicular ascending primary fibres and uni- or paucispicular transverse
connecting fibres cored by tracts of small styles of one category, echinating spicules absent, microscleres
are palmate isochelae. References: Dendy (1922:58); Bergquist & Fromont (1988:25); Bowerbank
(1866:331).
Burtonispongia de Laubenfels, 1936 (type species: Desmacidon australis Dendy, 1896) -
massive growth form; arenaceous ectosome with a well developed regular reticulation of broken foreign
spicules and sand tangential to the surface; stout radiating columns of sand inside fibres with reduced
spicule skeleton of slender strongyles scattered throughout the mesohyl and occassionally arranged in
loose whisps, especially near the surface; megascleres are strongyles only; microscleres are tridentate
(arcuate) isochelae and sigmas. References: de Laubenfels (1936:52); Wiedenmayer (1989:68); Dendy
(1896:19).
Cercidochela Kirkpatrick, 1907 (type species: Cercidochela lankesteri Kirkpatrick, 1907) -
elongated growth form; choanosomal skeleton with long longitudinal tracts radiating in a plumose
manner to the surface; megascleres oxeas only; microscleres peculiar shuttle-shaped chelae or
canonochelae with the single tooth from each end fused, and with a semicircular vertical lamella
extending inwards from the shaft and from the dental bridge so as to nearly meet. References:
Kirkpatrick (1907:284); Koltun (1964:45).
? Corybas Gray, 1867 (type species: ) .
Damiriopsis Burton, 1928 (type species: Damiriopsis brondstedi Burton, 1928) - massive;
ectosomal skeleton composed of tornotes with microspined bases forming surface brushes;
choanosomal skeleton composed of a loosely aggregated reticulation of strongyles with microspined
bases, singly or in bundles, verging on isodictyal in places; microscleres are palmate isochelae and
spatuliferous (anchorate) isochelae. References: de Laubenfels (1936:53); Burton (1928:124.
Desmacidon Bowerbank, 1861 (type species: Spongia fruticosa Montagu, 1818) - erect
growth forms; ectosomal skeleton composed of smooth oxeote spicules arranged in bouquets;
choanosomal skeleton composed of smooth diactinal megascleres forming an isodictyal reticulation;
microscleres are spatuliferous anchorate isochelae and sigmas. References: Ridley & Dendy (1887:103);
Bergquist & Fromont (1988:37).
Desmapsamma Burton, 1934 (type species: Fibularia anchorata Carter, 1882) - erect, ramose
growth form; ectosomal skeleton tough, arenaceous, with oxeas in bouquets and sand grains forming a
narrow-meshed reticulation sometimes completely replacing oxeas; choanosomal skeleton arenaceous,
also with smooth oxeas forming a paucispicular isotropic reticulation of short spicule tracts composed
of oxeas similar to those of the ectosome; megascleres slender oxeas; microscleres are anchorate
isochelae and sigmas. References: van Soest (1984:35); Hechtel (1965:21); Hartman (1967:20);
Alcolado (1976:5); Zea (1988:147); Desqueyroux-Faundez & Van Soest (1996, 1997).
Echinostylinos Topsent, 1927 (type species: Echinostylinos reticulatus Topsent, 1927) -
ectosomal skeleton consists of a tangential ectosomal subtylostyles, distinct from those in the
choanosome, and occasionally organised into erect brushes; choanosomal skeleton with plumose tracts
of larger choanosomal subtylostyles forming a loose reticulation, and occasionally echinating spicule
tracts; microscleres are arcuate isochelae and sigmas. References: Bergquist & Fromont (1988:45);
Topsent (1928:207).
Ectyonancora Lévi, 1963 (type species: Ectyonancora flabellata Lévi, 1963) - smooth
tornotes with hastate/ mucronate bases and an isodictyal reticulation of thick acanthostrongyles and
echinated by thick acanthostyles, microscleres are robust spatuliferous anchorate isochelae, no sigmas;
Desqueyroux-Faundez & Van Soest (1996).
Fibulia Carter, 1886 (type species: Fibulia carnosa Carter, 1886) (syn. Plumocolumella,
sensu Burton, 1929; Xytopsaga de Laubenfels, 1936) - massive; ectosomal skeleton regularly reticulate
composed of the same spicules as in the choanosomal skeleton, and with brushes of spicules protruding
through the surface; choanosomal skeleton plumose or plumoreticulate, with curved or sinuous primary
spongin interconnected frequently by smaller secondary fibres, both cored by bundles of small thin oxeas
or oxeotes ascending to surface and producing plumose dermal brushes, no echinating spicules;
microscleres are unguiferous (anchorate) isochelae and sigmas. References: Carter (1882:282;
1886:51); de Laubenfels (1936:51,54); Bergquist & Fromont (1988:40); Brondsted (1924:460);
Burton (1929:424).
Homoeodictya Ehlers, 1870 (type species: Spongia digitata Esper) (syn. Neoesperiopsis de
Laubenfels, 1949; Platychalina Ehlers, 1870; Textiliforma de Laubenfels, 1936) - lobate or palmate
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 55
growth forms; megascleres are oxeas; microscleres are modified palmate isochelae with slightly curved
shaft and expanded all the way along from end to end with the two lateral palms remaining united
together. References: Ridley & Dendy (1887:108); Koltun (1959).
Hymenancora Lundbeck, 1910 (type species: Leptosia umbellifera Topsent, 1904) -
encrusting; ectosomal skeleton with smooth ectosomal strongyles with polytylote tyles; choanosomal
skeleton hymedesmoid with erect acanthostyles on the substrate; microscleres are unguiferous isochelae
with multiple-dentate alae. References: Lundbeck (1910:116); Topsent (1904:192); Desqueyroux-
Faundez & Van Soest (1995).
Iotroata de Laubenfels, 1936 (type species: Iotroata acanthostylifera Stephens, 1916) (syn.
Iotaota de Laubenfels, 1936; Hymenotrochota Topsent, 1904; Hymenotrocha Burton, 1930) - massive;
ectosomal skeleton composed of smooth or mucronate ectosomal tylotes, mostly standing erect,
sometimes paratangentially, in bundles on the surface; choanosomal skeleton composed of smooth or
slightly acanthose styles, sometimes oxeote or absent, forming an isotropic reticulation; fibres are poor
but there may be abundant collagen; megascleres are large styles and smaller tylotes; microscleres are
anchorate (unguiferous) isochelae and birotulates, no sigmas. References: de Laubenfels (1936:84,95);
Carter (1874:245); Lundbeck (1905:188); Bergquist & Fromont (1988:94); Stephens (1916:236);
Desqueyroux-Faundez & Van Soest (1995).
Iotrochopsamma de Laubenfels, 1954 (type species: Iotrochota arbuscula Whitelegge, 1906)
- ramose; ectosomal skeleton finely reticulate, porous, arenaceous; choanosomal skeleton composed of
an open reticulate network of heavy spongin fibres, with long ascending primary fibres and shorter
transverse secondary fibres both cored with sand grains and foreign spicule fragments; megascleres are
absent; microscleres are birotulates. References: Whitelegge (1906:482); de Laubenfels (1936:127)
[possibly a synonym of Iotrochota].
Iotrochota Ridley, 1884 (type species: Halichondria birotulata Higgin, 1877) (syn.
Hiattrochota de Laubenfels, 1950; Iotrochostyla de Laubenfels, 1954) - encrusting, massive, digitate or
bushy growth forms, often black colouration with a purple staining mucus; ectosomal skeleton thick,
difficult to detach, composed of single spicules or poorly defined short tracts of spicules; choanosomal
skeleton a regular reticulation of multispicular tracts; collagen is heavy and usually darkly pigmented;
megascleres are smooth choanosomal styles, or oxeas, or only ectosomal strongyles inside fibres (all
about the same size, without definite localization of strongyles to the ectosomal skeleton); microscleres
are birotulates. Reference: van Soest (1984:38); Wiedenmayer (1977:138); Higgin (1877:297); de
Laubenfels (1936:95); Ridley (1884:433).
Isodictya Bowerbank, 1864 (type species: Spongia palmata Lamarck, 1814) - flabellate,
digitate growth forms; choanosomal skeleton is reticulate or plumoreticulate; megascleres are diactinal,
usually oxeas; microscleres are palmate isochelae. References: Bowerbank (1864:133); Bergquist &
Fromont (1988:39).
Melonanchora Carter, 1874 (type species: Melonanchora elliptica Carter, 1874) - fistulose
growth form, with paper-like thin ectosome composed of smooth ectosomal tylotes, smooth
choanosomal styles, microscleres are melonchelas or spherancres (modified anchorate isochelae);
Desqueyroux-Faundez & Van Soest (1995).
Myxilla Schmidt, 1862 (type species: Halichondria rosacea Lieberkuhn, 1859) (syn.
Dendoryx Gray, 1867; Ectyomyxilla Lundbeck, 1909; Emplocus Gray, 1867; Tereus Gray, 1867;
Stegxella Bowerbank, 1874; Hastatus Vosmaer, 1880; Stelodoryx Topsent, 1904; Crellomyxilla
Dendy, 1924; Burtonanchora de Laubenfels, 1936; Pseudomyxilla Koltun, 1955; Styloptilon Cabioch,
1968) - ectosomal skeleton composed of tylotes with variable terminations - mucronate, strongylote or
tornote, frequently with 1 or few prominent spines on apices - forming both a tangential skeleton and
vertical spicule brushes; choanosomal skeleton stylote, usually styles with or without spined bases,
forming a tight meshed isotropic reticulation or looser isodictyal skeleton; echinating acanthostyles
usually present but sometimes lost; microscleres are anchorate, unguiferous or spatulate isochelae often
of several sorts, and sigmas (the latter sometimes lost). References: Dendy (1924:364); Bergquist &
Fromont (1988:33,86); Bakus (1966:496); Desqueyroux-Faundez & Van Soest (1995); Lévi
(1963:35).
Five subgenera:
Myxilla (Myxilla) Schmidt, 1862 (type species: Halichondria rosacea Lieberkuhn, 1859)
Myxilla (Burtonanchora) de Laubenfels, 1936 (type species: Myxilla crucifera Wilson, 1925)
- ectosomal tylotes with smooth bases, choanosomal styles smooth never spined.
Myxilla (Stelodoryx) Topsent, 1904 (type species: Stelodoryx procera Topsent, 1904) -
microscleres are unguiferous anchorate isochelae instead or, or in addition to spatulate
anchorate chelae.
56 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
Onychomyxilla Topsent, 1927 (type species: Dendoryx pectinata Topsent, 1892) - smooth
ectosomal tylotes, choanosomal acanthostyles forming isotropic reticulation echinated by smaller
acanthostyles, microscleres are unguiferous isochelae verging on birotulates, no sigmas; Desqueyroux-
Faundez & Van Soest (1996).
Paresperia Burton, 1930 (type species: ) - subectosomal and ectosomal skeletons have loose
paratangential and tangential reticulations of acanthose or basally spined auxiliary subtylostyles,
choanosomal skeleton with reticulate tracts of the same spicules, echinating acanthostyles absent,
microscleres are palmate isochelae.
Plocamiancora Topsent, 1927 (type species: ) - smooth ectosomal tornotes with polytylote
tyles, choanosomal acanthostrongyles and long acanthostyles forming isotropic reticulation, echinating
acanthostyles present, microscleres are anchorate isochelae.
Plocamissa Burton, 1935 (type species: Plocamia igzo de Laubenfels, 1932) - encrusting;
ectosomal skeleton composed of smooth asymmetrical tornotes (one end pointed, the other inflated);
choanosomal skeleton "plocamiid" with an isotropic basal reticulation of single acanthostrongyles or
acanthotornotes echinated by long microspined styles in ascending plumose axial columns, with
abundant short acanthostyles echinating skeletal tracts; tylotes also occur interstitially; microscleres are
spatulate anchorate isochelae. References: de Laubenfels (1932:102; 1936:78); Burton (1935:401);
Desqueyroux-Faundez & Van Soest (1995). Note: possibly synonymous with Plocamionida.
Plumocolumetta de Laubenfels, 1936 (type species: Gellius bidens Topsent, 1901) -
subcylindrical; ectosomal skeleton detachable, hispid, but no specialised spicules; choanosomal skeleton
reticulate, slightly confused, with tracts cored by robust oxeas; microscleres are sigmoid anchorate
isochelae (vestigial, small teeth). References: de Laubenfels (1936:54); Topsent (1901:14).
Rotuloplocamia Lévi, 1952 (type species: ) - ectosomal tylotes or subtylote tornotes,
choanosomal acanthostrongyles forming isodictyal reticulation, microscleres birotulates.
Sigmarotula Bergquist & Fromont, 1988 (type species: Sigmarotula lamellata Bergquist &
Fromont, 1988) - lamellate; ectosomal skeleton with a tangential layer of tylotes (often with
asymmetrical bases), and also in weak brushes supporting the ectosomal membrane; choanosomal
skeleton consists of an isodictyal reticulation with triangular meshes, composed of smooth styles, with
thinner styles dispersed within the mesohyl; microscleres are birotulates and sigmas. References:
Bergquist & Fromont (1988:).
Stelotrochota Bakus, 1966 (type species: Stelotrochota hartmani Bakus, 1966) - encrusting;
ectosomal skeleton composed of an irregular tangential layer of tylotes with microspined swollen bases;
choanosomal skeleton composed of an isotropic or irregular isodictyal reticulation of large
choanosomal acanthostrongyles, with vaguely triangular meshes; microscleres are unguiferous
(anchorate) isochelae. References: Bakus (1966:501).
Tedandoryx de Laubenfels, 1954 (type species: ) - smooth ectosomal tylotes, smooth
choanosomal subtylostyles forming halichondroid confused architecture, microscleres are very thin
acanthostyles or roughened microstyles (resembling onychaetes), and arcuate isochelae [possibly a
Tedania contaminated with arcuate chelae (Desqueyroux-Faundez & Van Soest, 1995)].
network of fibres incorporating sand and foreign spicules, and with plumose tracts of strongyles;
microscleres are unguiferous isochelae and sigmas. References: Burton (1934:555); Hentschel
(1911:321).
Batzella Topsent, 1894 (type species: Halichondria inops Topsent, 1891) (syn.
Collosclerophora Dendy, 1917) - encrusting; ectosomal skeleton membraneous, without specialised
spiculation; choanosomal skeleton reduced to a hymedesmoid loosely plumose skeleton of strongyles
(tornotes); no microscleres. References: van Soest (1984:47); Wiedenmayer (1989:69).
Chondropsis Carter, 1886 (type species: Chondropsis arenifera Carter, 1886) (syn.
Sigmatella Lendenfeld, 1888) - skeleton incorporating varying amounts of sand and foreign material and
the skeleton organisation overall is plumose or plumo-reticulate; the spicule skeleton is always reduced
quantitatively to the foreign elements; megascleres are diactinal (strongyles, strongyloxeas); microscleres
are unquiferous isochelae and sometimes sigmas. References: Bergquist & Fromont (1988:42);
Wiedenmayer (1989:72).
Hemimycale Burton, 1934 (type species: ) - styles and anisostrongyles in both ectosomal and
choanosomal skeletons forming plumose tracts.
Kaneohea de Laubenfels, 1950 (type species: ) - single category of ectosomal and
choanosomal spicule (styles or strongyles), sand incorporated into fibres, microscleres are raphides.
Phoriospongia Marshall, 1880 (type species: Phoriospongia solida Marshall, 1880) (syn.
Psammotoxa de Laubenfels) - arenaceous ectosomal skeleton with heavy sand cortex, with or without
megascleres forming brushes protruding through the surface; choanosomal skeleton formed mainly by
sand grains but also with strongyles or styles, forming radial, plumose or dendritic tracts within the
choanosome and ascending to the surface; megascleres frail strongyles or styles, with frequent
intermediates, sometimes rudimentary; microscleres absent or may include sigmas, palmate isochelae
and toxas. References: Wiedenmayer (1989:71); Bergquist & Fromont (1988:42); Topsent (1933:19);
de Laubenfels (1936:99); Wiedenmayer (1989:67).
Psammascus Marshall, 1880 (type species: Psammascus decipiens Marshall, 1880) - tubular
growth form; choanosomal skeleton reticulate, with regular ascending primary fibres, and spongin fibres
cored by sand grains; no megascleres or microscleres. References: Wiedenmayer (1989:67); Marshall
(1880:92).
Psammoclemma Marshall, 1880 (type species: Psammoclema ramosum Marshall, 1880) (syn.
Psammopemma Marshall, 1880; Sarcocornea Carter, 1885) - ramose and massive growth forms;
ectosomal skeleton arenaceous, detachable; choanosomal skeleton dendritic tracts composed only of
sand grains united by spongin, or amorphously packed with sand; no megascleres or microscleres.
References: Wiedenmayer (1989:75); Marshall (1880:109). [This genus allegedly differes from
Psammascus by its skeletal structure (dendritic/amorphous versus distincly reticulate choanosomal
structure) (Wiedenmayer, 1989:75)].
Psammodoryx Burton, 1934 (type species: Dysidea chaliniformis Carter, 1885) - branching
growth forms; ectosomal skeleton arenaceous; choanosomal skeleton composed of a reticulation of
heavy spongin fibres cored with sand grains and foreign spicules; no proper megascleres; microscleres
are unguiferous isochelae. References: Burton (1934:554); Carter (1885:217).
Strongylacidon Lendenfeld, 1897 (type species: Strongylacidon sansibarensis Lendenfeld,
1897) (syn. Xytosiphum de Laubenfels, 1950) - encrusting to massive; ectosomal skeleton with
protruding brushes of stongyles from the ascending choansomal fibres; choanosomal skeleton plumose
in encrusting specimens becoming plumoreticulate in more massive specimens; fibres cored by
strongyles and some detritus; microscleres are unguiferous-arcuate isochelae and sigmas. References:
Wiedenmayer (1989:68); van Soest (1984:42); Bergquist & Fromont (1988:38).
Xytopsues de Laubenfels, 1936 (type species: ) - strongyles in ectosomal and choanosomal
skeletons, together with sand, microscleres are unguiferous-arcuate isochelae and sigmas.
GENERA:
Hemitedania Hallmann, 1914 (type species: Amorphina anonyma Carter, 1886) - massive;
ectosomal skeleton reticulate but without any specialised spicules; choanosomal skeleton composed of
reticulate or plumo-reticulate fibres cored by smooth choanosomal oxeas or tornotes, with an
arenaceous skeleton; megascleres are sharply pointed oxeas of a single category; microscleres are
onychaetes with an asymmetrical tyle. References: Hallmann (1914:431); Carter (1886:49).
Strongylamma Hallmann, 1917 (type species: Chondropsis carteri Dendy, 1895) - massive;
ectosomal skeleton arenaceous, without special spicules, but with a soft dermal cortex containing sand
grains and whispy loose tracts of strongyles supporting the ectosomal membrane; choanosomal skeleton
plumoreticulate, composed of arenaceous skeletal tracts, predominantly plumose ascending with fewer
transverse connecting fibres, cored by strongyles and large sand grains enclosed in moderately heavy
spongin fibres; microscleres are onychaetes of two morphologies, the larger asymmetrical. References:
Hallmann (1916:643); de Laubenfels (1936:95).
Tedania Gray, 1867 (type species: Reniera digitata Schmidt, 1862, jun. syn. of Halichondria
anhelans Lieberkühn, 1859) (syn. Trachytedania Ridley, 1881; Tedaniopsis Dendy, 1924;
Paratedania Burton, 1929; Oxytedania Sarà, 1978) - massive; ectosomal skeleton composed of tylotes
or tornotes with microspined bases forming tangential or paratangential surface tracts; choanosomal
skeleton composed of styles with smooth or microspined bases, producing reticulate, plumo-reticulate,
plumose or even dendritic architecture; microscleres are onychaetes. References: Dendy (1924:367); van
Soest (1984:50); Schmidt (1862:75); Wilson (1902:395); Hentschel (1912:340); Ridley (1884:417);
Koltun (1959:133); Bergquist & Fromont (1988:57); Wiedenmayer (1977:133; 1989:87);
Desqueyroux-Faundez & Van Soest (1995); Burton (1929:441); Kirkpatrick (1907:289).
Three subgenera:
Tedania (Tedania) Gray, 1867 (type species: Reniera digitata Schmidt, 1862) - smooth,
relatively small styles, occasionally strongylote styles as structural megascleres and
microspined tylotes as ectosomal megascleres.
Tedania (Tedaniopsis) Dendy, 1924 (type species: Tedaniopsis turbinata Dendy, 1924) -
relatively long thick smooth styles occasionally modified to anisostrongyles or anisoxeas as
structural megascleres, ectosomal megascleres are mucronate or tylostrongylote tornotes
occasionally with one or more vestigial spines.
Tedania (Trachytedania) Ridley, 1881 (type species: Trachytedania spinata Ridley, 1881) -
structural spicules are relatively small styles at least some of which show a few spines,
ectosomal megascleres are mucronate or oxeote tornotes.
Tedanione Wilson, 1894 (type species: Tedanione foetida Wilson, 1894) - encrusting, cryptic,
smooth strongyles the only megascleres forming a loosely plumose skeleton, microscleres onychaetes in
single or 2 size classes, with distinct tyles; Desqueyroux-Faundez & Van Soest (1995).
Tedaniopsamma Burton, 1934 (type species: Hircinia flabellopalmata Carter, 1885) -
flabellate digitate growth form; ectosomal skeleton arenaceous; choanosomal skeleton composed of a
reticulate skeleton with well developed fibres cored by and axial column of sand and smooth styles;
microscleres are raphides. References: Burton (1934:599); Carter (1885:313); Lendenfeld (1889:616)
[possible synonym of Hemitedania].
Asbestopluma Norman, 1882 (type species: Cladorhiza pennatula Schmidt, 1875) (syn.
Cotylina Lundbeck, 1905; Lycopodina Lundbeck, 1905) - erect stalked growth forms, or long stalked
cup; upper part penniform or with side-branches arising from all sides of an axis; skeleton consists of a
spicule axis divided into parallel fibres; in axis are spongin fibres, one running through each side branch,
or a coat of densely interwoven spicules; megascleres are styles or subtylostyles in the axial and extra-
axial skeletons, and minutely spined tylostyles or tylostrongyles found in the coat on the stalk;
microscleres are large palmate anisochelae, asymmetrical, with teeth on the smaller end, small fusiform
palmate isochelae and sigmas (and forceps in Lycopodina). References: de Laubenfels (1936:122);
Lundbeck (1905:44-72).
Chondrocladia Thomson, 1873 (type species: Chondrocladia virgata Thomson, 1873) (syn.
Neocladia Koltun, 1970) - symmetrical sponges with a basal stem and apical projections arising from a
rounded head; the skeleton consists of a spicule axis which branches into plumose tracts in the head of
the sponge; megascleres are styles; microscleres are unguiferous isochelae and sigmas. References:
Bergquist & Fromont (1988:28); Bergquist (1972:125); Boury-Esnault & van Beveren (1982:56).
Cladorhiza Sars, 1872 (type species: possibly Cladorhiza abyssicola Sars, 1872) (syn.
Axoniderma Ridley & Dendy, 1886; Exaxinata de Laubenfels, 1936; Raoa de Laubenfels, 1936;
Trochoderma Ridley & Dendy, 1886) - erect, branched or unbranched growth forms, often consisting of
a middle axis sending off branches more-or-less regularly (or hollow cup shaped in Raoa); base usually
with richly branched root; skeleton with axis of thick spongin and multispicular fibres with smaller
lateral fibres inserted supporting the lateral branches; megascleres styles, subtylostyle or tylostyles,
sometimes oxeas only (in Exaxinata); microscleres are anchorate anisochelae and sigmas; some species
also with straight anchorate isochelae with long shaft and small teeth "pseudo-amphiasters (in
Axoniderma). References: Ridley & Dendy (1886:342, 1887:97); Lévi (1964:73); Lundbeck (1905:78);
de Laubenfels (1936:343).
Crinorhiza Schmidt, 1880 (type species: ).
Meliiderma Ridley & Dendy, 1887 (type species: Meliiderma stipitata Ridley & Dendy, 1887)
(syn. Helophloeina Topsent, 1928) - erect, stipitate; ectosomal skeleton with erect tylostyles protruding
from choanosomal skeleton; choanosomal skeleton a dense band of fibres cored by tylostyles, forming
most of the stem, becoming plumose near the surface; megascleres styles or tylostyles, and may include
desmas; microscleres may include anchorate isochelae, anisochelae, sigmas and microstrongyles.
References: Topsent (1928); Hooper & Lévi (1989:440); Ridley & Dendy (1887:102).
of these spicules, occur within the mesohyl; microscleres sharp-toothed diancistras or cyrtancistras,
sometimes with toxas, individual or bundles of raphides (trichodragmata), or sigmas.
SCOPE: Eight nominal genera are included here, although only two are considered to be valid.
REVIEWS: Hentschel (1923), Lévi (1973), van Soest (1984)
GENERA:
Hamacantha Gray, 1867 (type species: Hamacantha johnsoni (Bowerbank, 1862)) (syn.
Athnacama de Laubenfels, 1936; ? Crellancistra de Laubenfels, 1936; Evomerula de Laubenfels, 1936;
Hypsispongia de Laubenfels, 1935 [in part]; Vomerula Schmidt, 1880; Zygherpe de Laubenfels, 1932)
- ectosomal reticulation, choanosomal subtylostyles, fibres echinated by diancistra microscleres which
also form rosettes dispersed between bundles of megascleres. References: van Soest (1984:143).
Pozziella Topsent, 1896 (type species: Pozziella clavisaepta Topsent, 1896) - encrusting;
ectosomal skeleton a detachable opaque membrane with tangential tylostyles; choanosomal skeleton
cavernous, with long multispicular fibres cored by long styles; microscleres are cyrtancistras and sigmas.
References: Topsent (1896; 1904:223); Lévi (1973:611).
spicule tracts completely enveloped in a spongin coat; interior is cavernous with little collagen;
megascleres are subtylostyles; microscleres are anisochelae and sigmas.
Mycale (Aegogropila) (type species: Halichondria aegogropila Johnston, 1842) - ectosomal skeleton
is a neat tangential reticulation of subtylostyles and supported by subectosomal brushes from
the terminal fibres of the choanosome; choanosomal skeleton of plumose tracts of
subtylostyles, with loose reticulation of single spicules dispersed between tracts; microscleres
are anisochelae, sigmas and raphides.
Mycale (Grapelia) (type species: ) -
Oxymycale Hentschel, 1929 (type species: Esperia intermedia Schmidt, 1875) - megascleres
are oxeas; microscleres are anisochelae in rosettes or alone. References: Hentschel (1929:932); Thiele
(1903:381; 1914).
Paresperella Dendy, 1905 (type species: Esperia serratohamata Carter, 1880) - encrusting or
massive; ectosomal skeleton if present consists of subtylostyles; choanosomal skeleton consists of
plumose or plumoreticulate tracts of subtylostyles; megascleres are subtylostyles of a single category;
microscleres are palmate anisochelae and serrated sigmas, and occasionally toxas. References: Bergquist
& Fromont (1988:25); Dendy (1905:362).
Sceptrella Schmidt, 1870 (type species: )
Semisuberites Carter, 1877 (type species: ) - reticulate skeleton.
Stylinos Topsent, 1891 (type species: Isodictya uniformis Bowerbank 1864) (syn. Ulosa de
Laubenfels, 1936; Stylaxinella Vacelet, 1960) - branching growth form; choanosomal skeleton consists
of an anisotropic rectangular reticulation of spongin fibres cored by styles and modified strongyles, less
frequently by oxeas, and differentiated into primary and secondary transverse lines, with tighter
anastomoses near the axis than in the periphery; extra-axial skeleton forms dendritic tracts near the
periphery; ectosomal skeleton is absent; microscleres are absent.
Stylotrichophora Dendy, 1895 (type species: Stylotrichophora rubra Dendy, 1895) -
compressed lobate growth form, smooth surface; ectosomal skeleton composed of plumose tufts of
choanosomal styles protruding from terminal fibres, piercing a regular, close-meshed reticulation of
foreign spicules held together by spongin, with circular meshes; choanosomal skeleton consists of a
reticulation of well developed spongin fibres cored by broken foreign spicules and sand grains, divided
into primary ascending and secondary transverse fibres, with styles dispersed within the mesohyl between
tracts, becoming more plumose, whispy near the surface; microscleres are raphides. References: Dendy
(1895:259).
Zygomycale Topsent, 1930 (type species: Raphiodesma parishi Bowerbank, 1875) - lobate;
tangential reticulate skeleton cored by styles, supported by a cavernous subectosomal skeleton and
plumose tracts of spicules; choansomal skeleton composed of multispicular tracts forming a reticulate
skeleton; megascleres are subtylostyles of a single category; microscleres are anisochelae, isochelae,
toxas, raphides and sigmas. References: Lévi (1956:16); Burton & Rao (1932:228); Wiedenmayer
(1989:84).
q ORDER HALICHONDRIDA
DEFINITION: Choanosomal skeleton composed of styles, oxeas, strongyles or intermediate spicules;
spicules not usually functionally localised to any particular region of the skeleton; skeletal structures
range from disorganised plumoreticulate, criss-crossed "halichondroid skeleton" to distinctly
compressed axis (or basal) region and a differentiated extra-axial (radial, plumose or plumoreticulate)
region; spongin fibres usually poorly developed or absent; ectosomal skeleton sometimes organised into
a tangential layer of spicules or erect spicule bundles, with minimal collagenous spongin, typically with
large subectosomal cavities; microscleres sparse including only raphides, microxeas, or spined
microxeas with a central bend.
Four families are included.
differentiation may be reduced (vestigial), but rudiments of these structures are always present;
microscleres usually absent, although a few genera have raphides or microraphides, sometimes forming
bundles (trichodragmata); reproduction oviparous.
SCOPE: Fifty one nominal genera are currently assigned to the Axinellidae, of which 42 are possibly
valid, although some of these may be moved elsewhere in the future based on a reappraisal of
synapomorphies for the genus.
REVIEWS: Hallmann (1916-1917); Hechtel (1969); Wiedenmayer (1977); Bergquist & Hartman
(1979); Hooper (1984b, 1986a); Hooper et al. (1992); Hooper & Bergquist (1992); Hooper & Lévi
(1993); Alvarez (in press).
GENERA:
Acanthella Schmidt, 1862 (type species: Acanthella acuta Schmidt, 1862) - encrusting, fan-
shaped, branching, bushy, or reticulate growth forms, with cartilaginous consistency; surface is fleshy,
porous, with conules raised into bushy folds; choanosomal skeleton is not obviously differentiated into
axial and extra-axial components, but there are vestiges of this differentiation in the form of with a
central skeleton of compressed branching and rejoining spongin fibres, cored by intermingled styles,
vermiform strongyles and/or oxeas, in uni- or paucispicular columns, forming a compressed
anastomosing network or completely compacted without any obvious separation of fibre and mineral
components; encrusting forms have a basal layer of strongyles lying adjacent to the substrate, with erect
styles embedded in the basal layer; ectosome is fleshy, without specialized megascleres, typically with
choanosomal spicules projecting through the surface.
Auletta Schmidt, 1870 (type species: Auletta sycinularia Schmidt, 1870) - specialized hollow
tubular, branching or cylindrical growth forms, with terminal oscules; choanosomal skeleton has a
basally condensed layer of sinuous strongyles and styles lining the endopinacoderm, and radial plumo-
reticulate extra-axial tracts of long styles/rhabdostyles, of two sizes, embedded perpendicular to the axial
skeleton; these extra-axial tracts ascending towards the surface in longitudinal bands, united by abundant
fibre and collagenous spongin, interconnected by occasional uni- or aspicular fibres; ectosome lacks a
specialized skeleton, but extra-axial spicules may pierce the surface singly or in brushes.
Axinectya Hallmann, 1917 (type species: ) - with short axial styles and long extra-axial styles.
Axinella Schmidt, 1862 (type species: Axinella polypoides Schmidt, 1862) (syn. Chalinissa
Lendenfeld, 1887; Astrospongia Gray, 1867) - flabellate and digitate growth forms; surface typically
hispid, conulose; choanosomal skeleton always with some axial compression of spongin fibres, with or
without differentiated primary and secondary fibre elements; fibres usually cored by styles, sometimes
oxeas, occasionally strongyles, or sometimes all three in various combinations; extra-axial skeleton
plumose or plumo-reticulate; ectosome without special megascleres but often with single spicules or
bundles of extra-axial spicules protruding through the surface; microscleres may include raphides or
microraphides, but these are not widely distributed amongst all species.
Axinosia Hallmann, 1914 (type species: Axinella symbiotica Whitelegge, 1907) - flabellate,
ramose or lamellar growth forms; surface bears small conulose processes; choanosome has
anastomosing axial spongin fibres cored by relatively small styles, together with fewer stongyles and/or
oxeas, which are all of approximately the same size; extra-axial skeleton is ascending, multispicular,
with transverse uni- or paucispicular fibres forming a regular plumo-reticulate to subrenieroid
reticulation, together producing a more-or-less regular branching pattern. Ectosome with dense plumose
brushes of extra-axial styles piercing the surface.
Bubaris Gray, 1867 (type species: Hymeraphia vermiculata Bowerbank, 1866) (syn.
Ommatosa de Laubenfels, 1936) - choanosomal skeleton with a condensed reticulation of smooth
flexuous or vermiform strongyles, sometimes only or straight oxeas, with projecting bundles or
individual styles ascending to the surface.
Bubaropsis Lévi & Vacelet, 1958 (type species: Bubaropsis curvisclera Lévi and Vacelet,
1958) - encrusting growth form; choanosome is a condensed basal layer of spongin cored by scattered
acanthostrongyles with tylote bases, lying adjacent to the substrate; extra-axial skeleton is disordered,
without an obvious radial structure, containing scattered tracts of curved oxeas, sharply angular or
toxiform oxeas, strongyles and raphides in bundles.
Cerbaris Topsent, 1898 (type species: Cerbaris torquatus Topsent, 1898) - encrusting growth
form; choanosomal skeleton consists of a basal layer of spongin fibres cored by twisted, U-shaped,
diactinal megascleres which are evenly acanthose, rarely smooth, often with strongylote points, and form
a discrete but overlapping basal reticulation; extra-axial skeleton consists of smooth styles-subtylostyles,
with subterminal swellings, standing perpendicular to the substrate, protruding through the surface.
Cymbastela Hooper & Bergquist, 1992 (type species: Pseudaxinyssa stipitata Bergquist and
Tizard, 1967) - lamellate, lobate or cup-shaped growth forms; choanosomal skeleton with compressed
reticulate axial spongin fibres cored by oxeas, sometimes with telescoped ends, in which major tracts run
longitudinally through lamellae; axial fibres generally decrease in spongin content towards the periphery
64 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
of lamellae; extra-axial skeletal tracts generally ascending, diverging, radial, plumose or plumoreticulate,
becoming plumose below the surface; specialised ectosomal spicules (smaller oxeas) present only in one
species, otherwise choanosomal spicules usually protrude through ectosome individually or in sparse
bundles. Predominantly autotrophic nutrition, most species containing symbiotic cyanobacteria.
Oviparous reproduction.
Dragmacidon Hallmann, 1917 (type species: Thrinacophora agariciformis Dendy, 1905) -
massive, thick-walled, stalked, cup-shaped growth form; surface is minutely conulose and grooved;
choanosomal skeleton without axial condensation, consisting of irregular, close-set plumose
multispicular spongin fibres and spicule tracts running perpendicular to the surface, and containing
intermingled oxeas and styles (or only oxeas), and with transverse, connecting pauci- or unispicular
fibres cored by the same megascleres; extra-fibre skeleton consists of larger oxeas or styles, which may
or may not pierce the surface; microscleres are raphides which occur singly or in bundles
(trichodragmata).
Dragmatella Hallmann, 1916 (type species: Desmacella aberrans Topsent, 1892a) - massive
growth form; surface with digitiform tapering processes; choanosome is cavernous, without axial
condensation, consisting of well-developed, non-plumose, multispicular spongin fibres cored by styles,
arranged in more-or-less dendritic structure, diverging and ascending to the surface; ectosome with
choanosomal styles lying tangential to the surface, criss-crossing in every direction; microscleres are
raphides with occur singly or in bundles (trichodragmata).
Dragmatyle Topsent, 1904 (type species: Dragmatyle lictor Topsent, 1904a) - encrusting
growth form; choanosome consists of a basal layer of spongin without coring megascleres; extra-axial
skeleton is composed of long subtylostyles embedded in the basal layer, erect on the substrate, and
protruding a long way through the ectosome; ectosomal skeleton consists of a special category of
sinuous, diactinal tornotes or oxeas forming a tangential layer on the surface; microscleres are raphides
occurring singly or in bundles (trichodragmata).
Dragmaxia Hallmann, 1916 (type species: Spongosorites variabilis Whitelegge, 1907) -
lamellate, fan or cup-shaped growth forms; surface has longitudinal radiate and grooved ridges;
choanosomal axial skeleton is condensed into a dense spicule axis, cored by smaller styles, occupying the
central portion of lamina; extra-axial skeleton consists of whispy plumose tracts, composed of larger
styles and poorly developed fibres, which radiate from the axis, sparsely interconnected by transverse
uni- or paucispicular tracts; ectosomal skeleton is fleshy, without specialized spiculation, but containing
numerous microscleres are raphides, occurring singly or in bundles (trichodragmata).
Homaxinella Topsent, 1916 (type species: Axinella supratumescens Topsent, 1907) -
branching, frondose, digitate or whip-like growth forms; choanosomal skeleton consists of a dense axial
reticulation of smaller styles-subtylostyles organized into rigid or loosely interlocking fibres; extra-axial
skeleton is radial, plumose, and may be confined to peripheral fibres only, comprised of larger styles;
ectosomal spicule layer is perpendicular or paratangential to the extra-axial spicule tracts, consisting of
larger styles.
Hymerhabdia Topsent, 1892 (type species: Hymerhabdia typica Topsent, 1892a) - encrusting
growth form; choanosomal skeleton consists of a basal layer of spongin fibres on the substrate, without
coring megascleres, but with extra-axial styles-tylostyles, or sometimes oxeas, and rhabdostyles erect on
basal layer and protruding through the surface.
Ketosus de Laubenfels, 1936 (type species: Dictyocylindrus setosus Bowerbank, 1873b) - fan-
shaped, branching growth form; surface raised into long papillae; choanosomal axial skeleton contains
sinuous strongyles coring compacted, anastomosing fibres; extra-axial skeleton consists of ascending
plumose tracts of sinuous oxeas; ectosomal skeleton has plumose brushes of subectosomal oxeas
projecting through the surface.
Lithobubaris Vacelet, 1969 (type species: Lithobubaris tenens Vacelet, 1969) - encrusting
growth form; choanosome consists of a solid condensed basal crust of monocrepidial desmas with
mammilliform and tuberculate lateral projections, which forms a rigid interlocking skeleton; extra-axial
skeleton consists of long smooth styles-subtylostyles with slightly rhabdose bases, perpendicular to and
entirely embedded in the basal skeleton, enveloped by the interlocking desmas, and only rarely protruding
beyond the surface.
Microstylifer Vacelet, 1969 (type species: Microstylifer rugosus Vacelet, 1969) - encrusting;
growth form; choanosomal skeleton is composed of irregular bundles of large basally spined styles,
forming a vague basal reticulation on the substrate; ectosomal skeleton is not detachable but contains
obliquely orientated auxiliary styles which are lightly basally spined; microstyles are dispersed
throughout the choanosome, but these probably represent immature auxiliary spicules.
Monocrepidium Topsent, 1898 (type species: Monocrepidium vermiculatum Topsent, 1898) -
encrusting growth form; choanosomal skeleton consists of a basal layer of globular or granular,
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 65
vermiform, diactinal monocrepidial desmas; with extra-axial styles or rhabdostyles embedded in basal
spongin, perpendicular to substrate, and protruding through the surface; mesohyl contains more slender
diactinal, vermiform megascleres than those of the axis.
Pachaxinella Burton, 1930 (type species: Halichondria subdola Bowerbank, 1866) -
branching growth form; choanosomal skeleton without axial condensation, consisting of an irregular
reticulation of fibres, cored by ascending paucispicular primary tracts of long styles and connected by
transverse uni- or paucispicular tracts of the same spicules; extra-axial skeleton is not markedly
differentiated from the deeper choanosomal region, although peripheral ascending tracts, cored by
choanosomal styles, protrude through the surface; ectosome has specialized spiculation consisting of
smaller styles scattered over the surface in dense paratangential or erect brushes.
Pararhaphoxya Burton, 1934 (type species: Pararhaphoxya tenuiramosa Burton, 1934a) -
stalked, branching growth forms; choanosomal skeleton is an axial core of sinuous strongyles,
intermingled with curved oxeas and styles in longitudinal tracts running along branches; extra-axial
skeleton consists of perpendicular radial tracts of curved oxeas and styles, more slender than
choanosomal spicules and with irregular terminations; ectosome is membraneous, with subectosomal
oxeas or styles poking through the surface.
Perissinella Topsent, 1928 (type species: Stylotella (Perissinella) madeirensis Topsent,
1928a) - massive, lobate-digitate growth forms; surface is microconulose and pitted; choanosome with a
more-or-less compressed axial skeleton of heavy spongin fibres, forming broad tracts which typically
anastomose, cored by thicker styles-subtylostyles, some with subterminal basal swellings and rhabdose
bases, in multispicular and irregularly arranged tracts; extra-axial skeleton consists of multispicular
longitudinal and ascending tracts, branching towards the periphery, with unispicular connecting spongin
fibres, cored by thinner styles-subtylostyles; ectosomal skeleton lacks specialized spiculation, but
diverging brushes or individual extra-axial styles protrude through the surface.
Phacanthina Vosmaer, 1912 (type species: Acanthella obtusa Schmidt, 1862) - lobate,
flabellate-digitate growth form; choanosome with an axially condensed core, without marked axial and
extra-axial differentiation, and axial fibres consist of dendritic or plumose, non-anastomosing, whispy
tracts, with very little spongin uniting spicules, cored by smaller styles; extra-axial skeleton is absent, but
long styles near the periphery of the skeleton project through the surface; ectosome without specialized
spiculation, although longer styles extend a long way through the surface.
Phakellia Bowerbank, 1863 (type species: Spongia ventilabra Linnaeus, 1767) (syn.
Querciclona de Laubenfels, 1936) - compressed flabellate or cup-like growth forms predominant;
surface is smooth or microconulose; oscules often surrounded by stellate subectosomal drainage canals;
choanosomal axial skeleton is a dense mass of interwoven spicules, typically composed of only
interwoven styles, or may include intermingled sinuous strongyles and styles, or occasionally only
strongyles, organized into multispicular-ascending and paucispicular-transverse tracts, together forming
a compressed reticulation at the axis; spongin fibres and collagenous spongin are sparse; extra-axial
skeleton consists of sparse plumose bundles or individual styles or oxeas standing perpendicular to the
axis, with or without transverse connecting megascleres; ectosomal skeleton membraneous without
specialized spiculation, usually fleshy, often with spicules protruding through the surface; microscleres
absent.
Phycopsis Carter, 1883 (type species: Phycopsis hirsuta Carter, 1883b) - stalked, branching or
massive, cup-shaped growth forms; surface is shaggy with filamentous processes; choanosomal skeleton
is axially condensed, cored by oxeas, with diverging ascending multispicular tracts forming the extra-
axial skeleton, interconnected by irregular uni- or paucispicular transverse fibres forming an overall
plumo-reticulate architecture; axial and extra-axial skeletons are not clearly differentiated near the core,
but tracts become increasingly plumose towards the periphery, forming a characteristically radial extra-
axial and ectosomal skeleton which extends into the surface processes.
Plicatellopsis Burton, 1932 (type species: Plicatellopsis arborescens Burton, 1932) - erect,
branching or fan-shaped growth forms; choanosome is an irregular or confused reticulation of long
styles-tylostyles forming definite spicule tracts, without axial condensation; extra-axial skeleton consists
of ascending and diverging plumose tracts of longer tylostyles; ectosomal skeleton consists of brushes of
small styles or tylostyles.
Pseudaxinella Schmidt, 1875 (type species: Pseudaxinella sulcata Schmidt, 1875) -
encrusting, massive, subspherical, club-shaped or unbranched lobate growth forms; surface often finely
conulose, tuberculate or corrugated; choanosomal skeleton plumo-reticulate, without a compresed axial
region, or differentiated axial and extra-axial regions; skeletal tracts crowded, plumose, more-or-less
parallel and anastomosing, composed of smaller, thicker oxeas (or anisoxeas) and styles in equal
proportion; ectosome fleshy, with a unispicular core of thinner, slightly longer spicules than in axial
66 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
region; megascleres typically include only (anis-)oxeas and styles in equal proportions, but some species
have flexuous styles or strongyles confined to the surface; microscleres absent.
Pseudotrachya Hallmann, 1914 (type species: Trachya hystrix Topsent, 1892a) - cylindrical
digitate, or subglobular semi-encrusting growth forms; choanosome contains an axial core of dispersed,
interwoven small oxeas; extra-axial skeleton consists of longitudinal, multispicular ascending tracts of
large styles embedded in the axis, protruding through the surface; ectosome with specialized skeleton
consisting of a dense palisade of small oxeas erect on the surface, in bundles, between which protrude
the longer extra-axial styles.
Ptilocaulis Carter, 1883 (type species: Ptilocaulis gracilis Carter, 1883b) (syn. Plicatella
Schmidt, 1864) - erect, cylindrical, clavate, bushy, and lamellate growth forms; surface prominently
conulose, with elongate, overlapping papilliform projections, often bifurcate at their points;
choanosomal skeleton plumo-reticulate, with clearly differentiated axial and extra-axial components;
axis compressed, composed of irregularly anastomosing close-set spongin fibres cored by styles,
subtylostyles, anisoxeas or strongyles (usually asymmetrical), sometimes including sinuous forms;
extra-axial skeleton plumo-reticulate or plumose, with heavy fibres cored by ascending multispicular
tracts of the same spicules, interconnected by paucispicular transverse tracts forming subisodictyal
reticulation (s.s.), or without transverse spicule skeleton and simply with meandering, plumose extra-
axial spicule tracts; ectosomal skeleton is fleshy, without specialized spiculation, but surface may be
pierced by plumose brushes of choanosomal styles; microscleres absent.
Reniochalina Lendenfeld, 1888 (type species: Reniochalina stalagmitis Lendenfeld, 1888)
(syn. Axiamon Hallmann, 1917) - branching, fan-shaped or lobate growth forms; surface is highly
conulose, tubercular, ridged; choanosomal skeleton without marked axial compression, and without
marked differentiation between axial and extra-axial regions, but with a lattice-like plumo-reticulation
of spongin fibres, cored by interchangeable oxeas, anisoxeas or styles, many with microspined points, in
approximately equal proportions and not restricted to any particular region of the skeleton; fibres divide
and rejoin to form rhomboidal meshes, without obvious differentiation between primary and secondary
elements; ectosome with individual spicules or loose bundles of anisoxeas piercing the surface;
microscleres are absent or may include raphides.
Rhabdoploca Topsent, 1904 (type species: Microciona curvispiculifera Carter, 1880) -
laminate encrusting growth form; choanosomal skeleton consists of a condensed basal layer of spongin,
with acanthose curved strongyles, with aspinose bases, standing erect; extra-axial skeleton consists of
long smooth rhabdostyles embedded in and perpendicular to the basal layer, projecting through the
surface, interdispersed with echinating smooth rhabdostyles; mesohyl contains acanthose strongyles
distributed without apparent order.
Rhaphoxya Hallmann, 1917 (type species: Rhaphoxya typica Hallmann, 1916c) (syn.
Acanthellina Carter, 1885) - massive growth form; surface with papilliform conules; choanosomal
skeleton without axial compression and without differentiated axial and extra-axial regions, but
consisting of a loose, irregularly reticulate, often meandering. slender, non-plumose, longitudinal and
transverse spongin fibres, poorly invested with spongin, cored by slender, sinuous, curved or flexuous
styles, oxeas and/or strongyles of one size category, differing only in their extremities; extra-axial spicule
tracts and fibres ascend to the surface, which may (s.s.) or may not protrude through the surface;
ectosome fleshy, lacking specialised spiculation; microscleres are raphides occurring singly or in bundles
(trichodragmata).
Siphonocalypta Burton, 1931 (type species: Auletta elegans Vosmaer, 1885) - hollow tubular
growth form; choanosomal skeleton is cavernous constructed around a central siphon ("gastral cavity"),
lined by a basally condensed, halichondroid, tangential layer of small, slightly curved or sinuous styles;
extra-axial skeleton is cavernous, consisting of ascending radial spongin fibres, non-branching, not
rejoining, cored by multispicular tracts of larger styles; ectosome with a tangential layer of smaller
styles.
Skeizia Cabioch, 1968 (type species: Skeizia minuta Cabioch, 1968) - encrusting growth form;
ectosome with a dense palisade of small auxiliary styles-subtylostyles standing perpendicular to the
surface, with long choanosomal (extra-axial) styles-subtylostyles also protruding a long way through the
surface. Choanosomal skeleton is leptoclathriid/hymedesmoid, with a layer of basal spongin lying on the
substrate, into which are embedded the bases of larger choanosomal megascleres, and surrounding which
are irregularly dispersed vermiform flexuous strongyles lying more-or-less tangential to the substrate,
which are ornamented by rounded tubercles.
Stylissa Hallmann, 1914 (type specices: Stylotella flabelliformis Hentschel, 1912) - fan, cup-
shaped and foliose growth forms; surface shaggy, often with small papillae or grooved ridges;
choanosomal skeleton disorganised plumo-reticulate, with slightly condensed axis and slight
differentiation between axial and extra-axial skeletons; well developed spongin fibres cored by parallel
tracts of styles of 1 or 2 sizes, more-or-less ascending and diverging towards the periphery; peripheral
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 67
styles often slightly larger than those in the axis; ectosome is fleshy, without specialized spiculation, but
with bundles of styles diverging and protruding through the surface; microscleres present or absent,
consisting of individual raphides or bundles (trichodragmata).
Stylotella Lendenfeld, 1888 (type species: Stylotella digitata Lendenfeld, 1888) - encrusting,
massive and branching growth forms; ectosomal skeleton membraneous with only protruding
choanosomal spicules; choanosomal skeleton with loosely plumose tracts and scattered strongyles found
throughout the skeleton; microscleres absent. References: Topsent (1891:); Wiedenmayer (1989:67);
Burton (1934:555); van Soest (1984:47); Lendenfeld (1888:185).
Teichaxinella de Laubenfels, 1936 (type species: Teichaxinella shoemakeri de Laubenfels,
1936a) (syn. Phakettia; de Laubenfels, 1936) - thin fan-shaped, lamellate or foliaceous growth forms;
choanosome is bread-like, halichondroid, without definite fibre organization, little collagenous spongin,
and without axial condensation, composed of short thick styles, occasionally modified to strongyles, and
thin oxeas; extra-axial specialization may be developed in the periphery only, consisting of scattered,
erect, long thin styles.
Thieleia Burton, 1932 (type species: Hymeniacidon rubiginosa Thiele, 1905) - encrusting
growth form; choanosomal skeleton is a basal layer of styles on the substrate; extra-axial skeleton
consists of plumose, ascending, delicate tracts of styles of a single category running to the surface, with
irregular transverse connecting fibres cored by unispicular tracts of styles, which together form an
irregular reticulation; ectosomal skeleton is pierced by brushes of styles diverging from the ascending
extra-axial tracts, forming a more-or-less continuous, dense dermal palisade.
Tragosia Gray, 1867 (type species: Spongia infundibuliformis Linnaeus, 1791) (syn.
Axidragma Hallmann, 1916) - funnel-shaped or thin fans, stipitate or branching growth forms;
choanosomal and extra-axial skeletons are regularly reticulate, without marked axial condensation,
consisting of plumose ascending primary spongin fibres and tracts of styles, with transverse secondary
(connecting) tracts of oxeas; ectosomal skeleton without megasclere specialization, but with
subectosomal styles poking through the surface; microscleres are raphides which occur singly or in
bundles (trichodragmata).
Uplexoa de Laubenfels, 1936 (type species: Bubaris oxeata Dendy, 1924a) - encrusting
growth form; choanosome consists of a basally condensed reticulation of small oxeas lying on the
substrate, with an extra-axial skeleton of long thick hastate styles perpendicular to the substrate, with
their bases embedded in the basal (oxeote) skeleton and projecting through the surface.
? Viles de Laubenfels, 1934 (type species: Viles ophiraphidites de Laubenfels, 1934) -
massively encrusting growth form; choanosomal skeleton is irregular (halichondroid), composed of
larger auxiliary oxeas intermingled with smaller oxeas and sinuos strongyloxeas. Megascleres are
apparently not localized to any particular region within the choanosome.
Waltherarndtia de Laubenfels, 1936 (type species: Hymeniacidon caliculatum Kirkpatrick,
1903) - stalked, vase-shaped growth form; choanosomal skeleton without axial condensation, consisting
of anastomosing spongin fibres cored by styles, forming more-or-less regular elongate meshes;
ectosomal skeleton with tufts of choanosomal megascleres protruding through the surface.
ordered and compact near the periphery; ectosome skeleton contains only spongin and scattered, curved
acanthoxea microscleres.
spicules and diverging spicule tracts ascending to the surface, protruding slightly beyond the ectosome,
and producing a finely conulose surface pattern, although ectosomal skeleton absent (from van Soest et
al. 1990).
Epipolasis de Laubenfels, 1936 (type species: Spongosorites suluensis Wilson, 1925) -
Massive-amorphous to flabellate, without a collagenous choanosome and without any spongin, resulting
in an utterly confused arrangement of spicules, but with a parchment-like feltwork of tangential oxeas
and sinuous trichodragmata (raphides) (from van Soest et al. 1990).
Halichondria Fleming, 1828 (type species: Spongia panicea Pallas, 1766) (syn. Amorphina
Schmidt, 1870; Densa de Laubenfels, 1934; ? Ciocalapata de Laubenfels, 1936; Cioxeamastia de
Laubenfels, 1942; Eumastia Schmidt, 1870; Halichondriella Burton, 1931; Menanetia Topsent, 1896;
? Pyloderma Kirkpatrick, 1908; Raspaigella Schmidt, 1868; Seriatula Gray, 1867; Trachyopsilla
Burton, 1931; Spuma Miklucho Maclay, 1870) - Massive-amorphous; ectosomal skeleton with thin,
tangential peel, clearly detachable, composed of single or vague bundles of smaller oxeote spicules,
supported by choanosomal columns of oxeas traversing subdermal spaces; choanosome has the
appearance of a disordered loose reticulation ("halichondroid" structure), and spicule tracts are poorly
defined, directionless; oxeote spicules of intermediate to relatively small size, sometimes modified to
quasistylote or even true styles; ectosomal structure may be secondarily lost; microscleres absent (s.s.) or
may include raphides occurring singly or in bundles (trichodragmata) in "Raspaigella"; larvae are
incubated and tufted-ciliate (modified from van Soest et al. 1990, Diaz et al., 1993).
Hymeniacidon Bowerbank, 1861 (type species: Hymeniacidon caruncula Bowerbank, 1859,
jun. syn. of Spongia perlevis Montagu, 1818) (syn. Amorphilla Thiele, 1898; Thielia Burton, 1932;
Rhaphidostyla Burton, 1935; Nailondra de Laubenfels, 1954; Rhaphoxiella Burton, 1934;
Stylohalina Kirk, 1909; Thieleia Burton, 1932; Uritaia Burton, 1932) - Encrusting to fistulose;
skeletal plan similar to that of Halichondria whereby ectosomal skeleton is thin, membraneous,
detachable, containing only tangential ectosomal styles or stylotes; ectosomal spicules are sometimes
aligned (e.g. in tracts like in Mycale), but generally ectosomal skeleton is poorly developed;
choanosomal skeleton varies from true "halichondroid" to disorganised with whispy, slightly plumose
ascending spicule tracts of styles or stylotes, and most species have a characteristic fleshy consistency;
oxeas have been lost completely and megascleres consist only of styles or stylotes (modified from van
Soest et al. 1990, Diaz et al., 1993).
Petromica Topsent, 1898 (type species: Petromica grimaldi Topsent, 1898) (syn. Monanthus
Kirkpatrick, 1903) - Massive encrusting, often fistulose; tangential ectosomal skeleton; choanosomal
skeleton confused, with little collagen and no spongin fibres, but showing some radial orientation near
the surface; with monocrepidial desmas producing a sublithistid basal skeleton, without monaxons
(from van Soest et al., 1990; Diaz et al., 1993).
Spongosorites Topsent, 1896 (type species: Spongosorites placenta Topsent, 1896) (syn.
Anisoxya, sensu Topsent, 1898 [not Anisoxia Mulsant, 1856]; Aponastra Topsent, 1927; Trachyopsis
Dendy, 1905) - Massive-amorphous to subspherical; ectosomal skeleton with smooth, thick, flaky crust
of paratangentially arranged, relatively thin spicules; choanosomal skeleton with spicules strewn in
confusion, occasionally grouped in spongin-enforced tracts, directed mainly parallel or oblique to the
surface; choanosome with poor collagen; many species with aerophobic pigments (modified from van
Soest et al. 1990, Diaz et al., 1993). Megascleres are oxeas of various sizes, sometimes with their points
modified to strongyles or styloids, but typically centrangulate; extra-axial region is cavernous without
any obvious skeletal tracts.
Topsentia Berg, 1899 (type species: Anisoxya glabra Topsent, 1898) (syn. Trachyopsis
Dendy, 1905; Alloscleria Topsent, 1927; Coelocalypta Topsent, 1928; Oxeostilon Ferrer-Hernandez,
1923; Laminospongia Pulitzer-Finali, 1983) - Massive-anorphous to lobate, brittle and rough texture;
bright colours typically absent; ectosomal skeleton composed of crust of compact, smaller ectosomal
oxeas lying paratangentially producing a microhispid surface (but optically smooth); choanosomal
skeleton with very poor collagen, no spongin fibres, resulting in an utterly confused, directionless
arrangement of spicules packed around canals and subdermal spaces; megascleres are oxeas of a wide
size range, usually in 2-3 size classes; twisted, bent or doubly-bent spicules sometimes present; no
raphide microscleres (modified from van Soest et al., 1990, Diaz et al., 1993).
Trachostyla Topsent, 1928 (type species: Trachostyla semota Topsent, 1928) - encrusting,
halichondroide, spongin absent, megascleres styles and subtylostyles completely microspined;
microscleres absent. Reviews: de Laubenfels (1936: 79).
q ORDER HAPLOSCLERIDA.
DEFINITION: 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,
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 71
rectangular or polygonal meshes; megascleres are exclusively oxeote or strongylote, bonded together
with collagenous spongin or enclosed within spongin fibres; microscleres, if present, may include sigmas
(frequently centrangulate), smooth toxas or microxeas.
Nine families of sponges are included, seven of which are viviparous, with parenchymella
bearing various patterns of ciliation, one oviparous group (Petrosiidae), and one uncertain
(Lubomirskiidae).
choanosomal skeleton unispicular isodictyal, with poorly developed spongin fibres; no microscleres (van
Soest, 1980).
Rhizoniera Griessenger, 1971 (type species: ) - ectosomal skeleton tangential, unispicular,
slightly irregular; choanosomal skeleton of primary uni- or paucispicular tracts and interconnecting
single spicules is further strengthened by longitudinal spicule tracts (van Soest, 1980).
Sigmadocia de Laubenfels, 1936 (type species: Gellius textapatina de Laubenfels, 1926) -
ectosomal tangential unilayered reticulation of single spicules bound at their nodes by spongin;
microscleres sigmas (van Soest, 1980).
Toxadocia de Laubenfels, 1936 (type species: Gellius abbreviatus Topsent, 1918) (syn.
Neoadocia de Laubenfels, 1950) - ectosomal tangential unilayered reticulation of single spicules bound
at their nodes by spongin; microscleres toxas (van Soest, 1980).
Toxiclona de Laubenfels, 1954 (type species: Siphonochalina gaussiana Hentschel, 1914).
reticulation of spicules and/or fibres lying tangential to the surface; ectosomal skeleton a two
dimensional tangential reticulation of close-set primary, secondary and sometimes tertiary spongin
fibres, sparsely cored with small or vestigial oxeas or strongyles; choanosomal skeleton more widely
spaced, composed of a reticulation of primary ascending (bi- or multispicular) and secondary connecting
spongin fibres (uni- or aspicular), composed of well developed fibres, cored by oxeas or strongyles;
spongin characteristically abundant; megascleres sometimes vestigial, with blackened axial canals, absent
entirely or replaced by sand grains; microscleres, if present, include only toxas.
SCOPE: There are 21 nominal genera of Callyspongiidae, including many names created by Lendenfeld,
but only five of these may be valid.
REVIEWS: Bergquist & Warne (1980), van Soest (1980).
GENERA:
Arenosclera Pulitzer-Finali, 1982 (type species: Arenosclera heroni Pulitzer-Finali, 1982) -.
Callyspongia Duchassaing & Michelotti, 1864 (type species: Callyspongia fallax
Duchassaing & Michelotti, 1864) (syn. Ceraochalina Lendenfeld, 1887; Chalinella Lendenfeld, 1887;
Chalinopora Lendenfeld, 1887; Cladochalina Schmidt, 1870; Euchalina Lendenfeld, 1887;
Patuloscula Carter, 1882; Platychalina Ehlers, 1870; Siphonella, sensu Lendenfeld, 1887;
Tubulodigitatus Carter, 1881) - with a regular ectosomal tangential reticulation of primary and
secondary and sometimes tertiary spongin fibres; choanosome with a well developed spongin fibre
skeleton; all primary fibres are regularly reticulate producing rectangular meshes; skeletal tracts usually
dominated by spongin, and spicules generally small or rudimentary (van Soest, 1980).
Dactylia Carter, 1885 (type species: ) (syn. Chalinopsis Lendenfeld, 1886; Chalinopsilla
Lendenfeld, 1888) - regularly hexagonal ectosomal tangential skeleton of primary, secondary and
sometimes tertiary spongin fibres, without coring spicules although some detritus may be included into
the primary fibres; choanosomal spongin fibres similarly uncored or with sparse detritus; both
megascleres and microscleres absent (Bergquist & Warne, 1980; van Soest, 1980).
Euplacella Lendenfeld, 1885 (type species: ) (syn. Placochalina Lendenfeld, 1887) - regular
ectosomal tangential reticulation of primary and secondary spongin fibres cored sparsely, in addition to
which there are erect spicule brushes on the primary and secondary fibres; choanosomal spongin fibres
cored sparsely; microsclere absent; megascleres usually very small or vestigial (minute with blackened
axial canals (modified from van Soest, 1980).
Siphonochalina Schmidt, 1868 (type species: Siphonochalina coriacea Schmidt, 1868) -
tubular growth form; irregular ectosomal tangential reticulation in which the sparsely cored primary and
secondary spongin fibres are indistinct; spongin is more poorly developed than in Callyspongia;
choanosomal reticulation irregular; fibres cored sparsely; microscleres absent (van Soest, 1980).
Spinosella Vosmaer, 1885 (type species: ) (syn. Tuba Duchassaing & Michelotti, 1864) - like
Callyspongia but all primary fibres are fasciculate (van Soest, 1980).
Toxochalina Ridley, 1884 (type species: ) - regular ectosomal tangential reticulation of
primary and secondary spongin fibres cored sparingly; choanosomal reticulation regular with rectangular
meshes and fibres sparsely cored; microscleres are toxas - (van Soest, 1980).
Foliolina Schmidt, 1870 (type species: Foliolina peltata Schmidt, 1870) - hollow pagoda-like
habit (van Soest, 1980).
Oceanapia Norman, 1869 (type species: Desmacidon jeffreysi Bowerbank, 1866) (syn.
Rhizochalina Schmidt, 1870; Phloeodictyon Carter, 1882; Biminia Wiedenmayer, 1977) - long
aquiferous fistules on upper surface; spongin-enforced tangential ectosomal crust composed of single
spicules or paucispicular tracts in isodictyal or irregular (isotropic) arrangement; fistule walls in
particular are supported by an irregular isotropically meshed reticulation of spicule tracts bound by
spongin; choanosomal skeleton often pulpy, with multispicular tracts of spicules, with our without
single spicules dispersed between major spicule tracts; microscleres may include sigmas and/or toxas
(modified from Bergquist & Warne, 1980; van Soest, 1980).
Pachypellina Burton, 1934 (type species: Pachypellina podatypa (de Laubenfels, 1934)) - no
fistules; thick ectosomal crust of tangential spicules and a densely spiculous choanosome which is ridled
with holes; no microscleres (van Soest, 1980).
Pellina Schmidt, 1870 (type species: Pellina semitubulosa (Schmidt, 1862)) - simple
complement of small (100-250µm) oxeote megascleres; microscleres absent; fistules walls thin,
consisting only of an ectosomal crust and longitudinal supporting spicule tracts (van Soest, 1980).
Dosilia Gray, 1867 (type species: ) (syn. Asteromeyenia Annandale, 1912; Astromeyenia
Schröder, 1927).
Ephydatia Lamouroux, 1816 (type species: Spongia fluviatilis Linneaus, 1758)
(Trachyspongilla Dybowsky, 1878; Pleiomeyenia Mills, 1884; Meyenia Carter, 1881; Tupha Oken,
1814 [note: the latter has seniority over Ephydatia]).
Eunapius Gray, 1867 (type species: Spongilla carteri Bowerbank, 1863).
Heteromeyenia Potts, 1881 (type species: Spongilla baileyi Bowerbank, 1863) (Carterella
Potts, 1881; Carterius Petr, 1886; Oxyheteromeyenia Schröder, 1927).
Heterorotula Penney & Racek, 1968 (type species: Spongilla capewelli Bowerbank, 1863).
Nudospongilla Annandale, 1909 (type species: ).
Pachyrotula Volkmer-Ribeiro & Ruetzler, 1997 (type species Spongilla (Stratospongilla)
raceki Rutzler, 1968) -
Pectispongilla Annandale, 1909 (type species: Pectospongilla aurea Annandale, 1909).
Radiospongilla Penney & Racek, 1968 (type species: Spongilla sceptroides Haswell, 1882).
Spongilla Lamarck, 1816 (type species: Spongia lacustris Linneaus, 1758) (syn. Euspongilla
Vejdovsky, 1883). – 2 subgenera:
Spongilla (Spongilla) Lamarck
Spongilla (Stratospongilla) Annandale, 1909 (type species: Spongilla bombayensis Carter,
1882).
Trochospongilla Vejdovsky, 1883 (type species: Trochospongilla horrida Weltner, 1893).
Umborotula Penney & Racek, 1968 (type species: Ephydatia bogorensis Weber, 1890).
tracts; spicules bound together with abundant spongin fibres; microscleres absent; larvae apparently
produced parthenogenetically.
SCOPE: This family presently contains seven nominal genera, three of which are well established.
REVIEWS: Nil.
GENERA:
Baikalospongia Annandale, 1914 (type species: ).
Baicalopis Makuschok (type species: ).
Cortispongilla Annandale, 1909 (type species: ).
Lubomirskia Dybowsky, 1880 (type species: ).
Ochridaspongia Arndt (type species: ).
Pachydictyum Weltner (type species: ).
Swartschewskia Makuschok (type species: ).
q ORDER DICTYOCERATIDA.
DEFINITION: "Keratose sponges" lacking mineral spicules, although detritus and contaminating
spicules may be acquired; sponges usually tough, difficult to tear, and frequently with differences in
pigmentation between the surface and subectosomal regions; main skeleton a reticulation of spongin
fibres, often organised into primary, secondary and sometimes tertiary networks; fibres usually
homogenous 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 parenchymella, 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.
REMARKS. Four families are included in the dictyoceratids, differentiated by their fibre characteristics,
although there is currently some debate about whether Dysideidae should be included here or with the
Dendroceratida (based on affinities inferred by comparative choanosomal ultrastructures and supporting
chemical evidence for affinities to dendroceratids).
due to density and regularity of spongin fibres; surface not armoured, but producing irregular truncate
conules with dermal membrane stretched between conules and over subdermal canals (Bergquist, 1980).
Hippospongia Schulze, 1879 (type species: Spongia communis Lamarck, 1814) (syn.
Aphrodite Lendenfeld, 1885; Ceratodendron Marshall, 1878; Hippospongia de Laubenfels, 1936) -
Surface finely conulose, darkly pigmented, but not armoured; fibre network highly developed and
characterised by the almost complete absence of primary cored fibres, which occur only near the surface;
tangled secondary fibre network forms most of the skeleton, which is supple and elastic; subsurface
region prominently cavernous (Bergquist, 1980).
Hyattella Lendenfeld, 1888 (type species: Hircina clathrata Carter, 1881) (syn.
Luffariospongia Burton, 1937; Trypespongia de Laubenfels, 1936) - Sponge highly lacunar, with large
subdermal cavities and excavations through the body; growth form tubular, erect and repent, sometimes
partially burrowing with apical tubes; pale yellow to orange brown colours are typical. Primary fibres
are a prominent feature of skeleton, regularly spaced and cored with detritus; secondary fibres dense,
regularly spaced, with no detritus; texture is hard, just compressible; surface unarmoured and conulose,
transparent only where collagenous dermal membrane extends between and over the large surface
cavities produced by repeated folds on the sponge body (Bergquist, 1980).
Leiosella Lendenfeld, 1888 (type species: Leisella elegans Lendenfeld, 1889) - Cup-shaped,
lobed or flabellate growth forms; thin but ordered sand armour; texture firm; primary fibres lightly cored
with detritus, tend to become fasciculate where they arise out of the dense secondary network or just
below surface; dense secondary fibre network and secondary fibres also often contain some spicule
detritus, usually as a single discontinuous line of fragments (Bergquist, 1980).
Lendenfeldia Bergquist, 1980 (type species: Phyllospongia dendyi var. frondosa Lendenfeld,
1889) - Complex interlocking lamellate or lamello-digitate growth forms; texture fleshy and soft;
primary fibres cored with detritus and orientated without rigid relationship to the surface, and often
fasciculate. Secondary and tertiary fibre networks form an irregular but not dense mesh; no sand cortex;
surface covered with very fine conules but not grooved as in Carteriospongia (Bergquist, 1980).
Phyllospongia Ehlers, 1870 (type species: Spongia papyracea Esper, 1806) (syn. Mauricea
Carter, 1877) - lamellate, vasiform, digitate or foliose sponges usually of very thin-walled construction,
up to 4mm thick except in digitate forms; surfaec is smooth macroscopically but irregularly corrugated
and regularly conulose microscopically; oscules small, flush with the surface or elevated on low mounds
emphasised by sand and collagen deposition around each rim; skeleton rectangular reticulation
constituted of primary elements disposed at right angles to the surface and secondary connecting
elements aligned parallel to the surface; primary elements contain coring material but this is contained
will within the investing spongin and never causes the fibre to become irregular in outline; secondary
elements are clear and variable in quantity, their relative dominance proportional to the thickness of the
body construction; the pattern of the primary and secondary skeleton is extremely regular and rectangular
in very thin species; in those with slightly thicker habit it becomes less regular as the secondary network
expands between the primary columns; tertiary fibrous elements are also present, sometimes dispersed
but predominantly disposed as an axial skeleton; these vermiform elements are invariably present in basal
and stalk regions; in forms with digitate morphology they form marked axial fascicles disposed at right
angles to the primary fibres and extending throughout all but the most marginal regions of the body; an
organised sand cortex is usually present on one or both surfaces, but it never becomes a pronounced
crust as in Carteriospongia (Bergquist et al., 1988).
Rhopaloeides Thompson, Murphy, Bergquist & Evans, 1987 - (type species: Rhopaloeides
odorabile Thompson et al., 1987) - texture and fibre quality of a Spongia in which the cored primary
fibres form simple fascicles; this condition is evident throughout the sponge but is most pronounced
near and at the surface where the fasciculate fibre whorls dominate as skeletal elements and support
prominent surface conules; the secondary skeleton is comparable to that of Spongia but is more
compact; the sponge is massive, erect in the form of thick broad-based lamellae or multiple fused thick
clubs; surface conules are thick, tuberculate rather than pointed and simple as in species of Spongia,
each one supported by several fibre tracts; surface is clear of detritus and not armoured by development
of a sand cortex (Thompson et al., 1987).
Spongia Linneaus, 1759 (type species: Spongia officinalis Linneaus, 1759) (syn. Ditela
Schmidt, 1862; Euspongia Bronn, 1859) - Massive, spherical, lamellate or cup-shaped sponges. Texture
is springy, very compressible, supple, elastic; surface never armoured, but covered with low even
conules and frequently pigmented black, brown or grey (offset from white or beige interior). Primary
fibres are reduced in number; secondary fibres are a highly developed network of fine, intertwined fibres
making up bulk of skeleton; primary fibres contain central axis of foreign material and most are found
near surface of skeleton; secondary fibres with no detritus (Bergquist, 1980).
78 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
SCOPE: Twenty five nominal genera are included in the family, 13 of which are probably valid.
REVIEWS: de Laubenfels (1948), Bergquist (1980, 1995), Boury-Esnault et al. (1990: 237), Bergquist
(1995).
GENERA:
Aplysinopsis Lendenfeld, 1888 (type species: Aplysinopsis elegans Lendfeld, 1888) - Digitate
to thick-walled tubular construction; texture is fleshy and compressible; surface membrane supports a
thin but regular sand armour and is covered with rounded conules; primary fibres simple and cored;
secondary fibres clear and form an irregularly branching network in which some branches may be quite
fine (Bergquist, 1980)
Cacospongia Schmidt, 1862 (type species: Cacospongia mollior Schmidt, 1862) - surface
finely and evenly conulose superficially resembling Spongia; surface never armoured; texture
compressible and easily torn; fibres stratified, harsh, brittle; primary fibres are simple and cored with
detritus, to some extent masking the granular pith; secondary fibres clear, regularly spaced, forming an
almost rectangular reticulation, with or without pith (Bergquist, 1980).
Collospongia Bergquist, Cambie & Kernan, 1990 (type species: Collospongia auris
Bergquist, Cambie & Kernan, 1990) - Thin compressed lamellate and simple to multiple cup-shaped
habit with skeletal structure typical of the family but augmented by a surface skeleton in the form of a
tangential network of fine regular meshed fibres on both oscular and poral faces of the flattened body.
Primary skeleton fasciculate and lightly cored, compressed into a central position and oriented parallel to
the substrate. From this fibre core fascicles curve to intersect both surfaces. The secondary skeleton is
compressed and is arranged in similar fashion but the fibres are clear of debris. All fibres are strongly
laminated and pithed (Bergquist, Cambie & Kernan, 1990).
Fascaplysinopsis Bergquist, 1980 (type species: Aplysinopsis reticulata Hentschel, 1912) -
surface marked by prominent conules and delicate tracery of sand, but no complete sand cortex; surface
pigmentation (blackish) markedly different from choanosome (yellowish); texture firm, harsh in ethanol
due to coarse nature of the fibres; primary fibres fasciculated and cored with detritus; secondary fibres
not cored and branch in irregular fashion; fibre meshes together do not form a regular-meshed skeleton;
adjacent components of the primary fascicles sometimes joined in ladder-like fashion producing a simple
arrangement reminiscent of Thorecta; all fibres thick with visible pith; choanosomal region cavernous,
typically gelatinous (Bergquist, 1980).
Fasciospongia Burton, 1934 (type species: Stelospongia australis var. fovea Lendfendfeld,
1888) (syn. Stelospongos Schmidt, 1870; Stelospongia, in part, Schulze, 1879; Stelospongus, in part,
Ridley, 1884) - Globular, tubular, cup or fan shapes with marked exhalant canals running either centrally
or superficially; surface has a shiny skin-like appearance produced by considerable collagenous
deposition in the ectosomal region; conules multiple because of the fibre fascicles, and often truncate
rather than pointed; only slight differential pigmentation at the surface, more frequently colour is
uniform throughout the sponge; primary fibres cored with detritus and arranged in stout fascicles which
curve in plumose fashion towards sponge surface; secondary skeleton clear of detritus, and mesh is
branching but regular (Bergquist, 1980).
Fenestraspongia Bergquist, 1980 (type species: Hircinia intertexta Carter, 1885) - External
surface characterised by the regular fenestrated surface; surface extensions intersect to produce a most
characteristic pattern of ridges; cored primary columns form stout fascicles which curve in plumose
fashion towards the surface and extend in rows into plate-like extensions of the collagenous surface
layer; secondary fibre reticulation is dense, branching and clear of detritus; in the interstices of the
secondary network a fine tertiary fibre reticulation also present (Bergquist, 1980).
Hyrtios Duchassaing & Michelotti, 1864 (type species: Hyrtios proteus Duchassaing &
Michelotti, 1864) (syn. Duriella Row, 1911; Dysideopsis Lendenfeld, 1888; Heteronema, in part,
Keller, 1889; Inodes de Laubenfels, 1957; Oligoceras Schulze, 1879; Thorectopsamma Burton, 1934)
- Texture ranges from compressible to quite firm or even brittle, reflecting the degree of development of
the skeleton and debris in the skeleton; surface distinctly conulose, with primary fibres terminating in the
conules and sandy inclusions within fibres appearing whitish against the dark sponge surface;
collagenous mesohyl containing extraneous detritus; both primary and secondary fibres fully charged
with detritus to some extent obscuring stratified nature of spongin fibres; primary skeleton shows some
fasciculation near the surface (Bergquist, 1980).
Luffariella Thiele, 1899 (type species: Luffaria variabilis Poléjaeff, 1884) (syn. Luffaria, in
part, Poléjaeff, 1884) - similar to Cacospongia having a fine conulose unarmored surface, and cored
non-fasciculated primary fibres, but primary and secondary fibres supplemented by a fine tertiary
network; secondary and tertiary skeletal mesh has a complex branching pattern (Bergquist, 1980).
Petrosaspongia Bergquist, 1995 (type species: Petrosaspongia nigra Bergquist, 1995) -
Massive, speading, lobate; surface finely conulose; ectosomal skeleton clearly set off from choanosome,
heavily collegenated, with large subectosomal lacunae; choanosomal skeleton extremely dense made up
80 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
of predominantly tightly interlocking, strongly laminated uncored secondary fibres forming irregular
polygonal meshes; primary fibres short, cored and confined mainly to surface; detritus common in
choanosome; choanocyte chamber small, spherical; like Hyrtios but with irregular skeletal reticulation,
uncored secondary fibres and distinct cored primary fibres (Bergquist, 1995).
Smenospongia Wiedenmayer, 1977 (type species: Aplysina aurea Hyatt, 1875) - surface
pigmentation variable, brown to purple-brown, choanosomal pigmentation yellowish, but sponge dries
to a dark colour which is not homologous to the Verongida aerophobic colour changes; primary
ascending fibres are much reduced and, when present, are represented by slightly fasciculated columns at
and near the sponge surface; these columns are connected by a dense irregular reticulation of secondary
fibres; fibres contain no foreign coring material but do retain pith elements although in small and
variable quantity; fasciculate primary fibres in superficial region are arranged in an irregular honeycomb
appearance, but this is lost in the interior (Bergquist, 1980).
Taonura Carter, 1882 (type species: Taonura flabelliformis Carter, 1882) - Stipitate, cup or
lamellate shape typical; texture soft, compressible; sponge surface covered with fine small conules,
never armoured, but often has undulating pitted appearance over whole or part of surface; primary fibres
cored and secondary fibres clear; mesh is regular and approximately rectangular (Bergquist, 1980).
Thorecta Lendenfeld, 1888 (type species: Thorecta cribrocusta Carter, 1886) (syn.
Antheroplax Lendenfeld, 1889; Geelongia, in part, Carter, 1885) - , Stalked, globular, fan-shapes
typical; surface always arenaceous, not conulose; primary fibres cored with a regular axial column of
debris and secondary fibres clear; skeleton forms a regular almost perfectly rectangular mesh in which
the spaces between the fibres may be quite large (Bergquist, 1980).
Thorectandra Lendenfeld, 1889 (type species: Halispongia choanoides Bowerbank, 1872)
(syn. Halispongia, sensu Bowerbank, 1866) - Stalked, globular, fan-shaped or tubular, with apical or
fringing oscules; excessive mucus production typical; surface heavily armoured and always thrown into
series of ridges and hollows; primary fibres cored with an axial column of detritus and secondary fibres
clear; skeleton almost perfectly rectangular with huge mesh spaces, up to 4mm, and thick fibres;
elements of fibrous skeleton are reduced in relation to the soft tissue, so the whole sponge inside the
sand cortex is soft, easily crumbled and collapsible (Bergquist, 1980).
q ORDER DENDROCERATIDA.
DEFINITION: "Keratose sponges", without mineral spicules, with dendritic or reticulate skeleton, 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
parenchymella, evenly ciliated, with or without a posterior tuft of long flagella.
Three families are traditionally included here, distinguished by their respective fibre
development and skeleton arrangement, although Dysideidae was also recently included in this order
following ultrastructural and chemical evidence (Boury-Esnault, de Vos, Donadey & Vacelet, 1990;
Berguist Karuso & Cambie, 1990), although this has not yet been universally accepted.
network, secondary fibres produce branching pattern; primary fibres cored by sand particles, secondary
fibres lack foreign inclusions, but all fibres laminated (Bergquist, 1980).
Spongionella Bowerbank, 1862 (type species: Spongionella pulchella Sowerby, 1806) (syn.
Megalopstas, in part, Dendy, 1905; Velinea Vosmaer, 1883) - skeleton is compact reticulation of
uncored primary and secondary fibres in which concentric laminations and a pith component are always
apparent although variable in extent; skeletal arrangement is regular, almost perfectly rectangular in the
type species; primary fibres are extended into fine tapered projections and surface is finely conulose
(Bergquist, 1980).
lightly reinforced with collagen no detritus; choanocyte chambers large, oval, eurypylous (Bergquist,
1995).
Dictyodendrilla Bergquist, 1980 (type species: Dendrilla cavernosa Lendenfeld, 1883) (syn.
Megalopastas , in part, Vacelet, 1958) - reticulate fibrous skeleton is regularly rectangular and
composed of pithed laminated fibres; fibres without any coring material; tissue construction is delicate
and cavernous, and the soft tissue is frequently pale contrasting with the dark fibres; the sponges are
often lobate, stalked or spreading with digitate projections (Bergquist, 1980).
Igernella Topsent, 1905 (type species: Euryades notabilis Duchassaing & Michelotti, 1864) -
fibrous skeleton is irregular, reticulate and frequently reduced in relation to the bulk of the soft tissue;
fibres may be concentrated superficially; fibres are laminated and have a pith although this can be
obscured by the incorporated detritus; fibre skeleton is augmented by diactinal and triactinal horny
spicules comparable in structure to those of Darwinella; reticulate skeleton arises from a basal spongin
plate (Bergquist, 1980).
Pseudobasta Topsent, 1931 (incertae sedis) (type species: Pseudobasta hermanni Topsent,
1931) - fibres with pith but these do not dry back to leave a hollow canal as in the Aplysinidae; possibly
in Dictyodendrillidae (Bergquist, 1980).
q ORDER VERONGIDA.
DEFINITION: "Keratose sponges" lacking spicules, typically fleshy and soft, with pigment that oxidizes
to purple colouration; skeleton with large, widely spaced spongin fibres forming dendritic or reticulate
structures; fibres may be aggregated (fasciculated) into bundles; 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.
surface, or localised oscular region; sponge is cavernous with extremely thick fibres which form an
irregular anastomosing reticulum; soft tissue is sparse in relation to fibrous material; fibres have
substantial pith and a bark component in which numerous spongocytes are arranged in concentric annuli;
no evidence of compression of the skeleton into a single plane as in Ianthella; choanocyte chambers are
basically eurypylous but are elongate-oval and sometimes branched in a manner reminscent of Halisarca
(Bergquist, 1980).
Bajalus Lendenfeld, 1885 (type species: Bajalus laxus Lendenfeld, 1885) - fibrous skeleton
absent and strong collagen deposition marks a cortex distinct from the endosomal region; choanocyte
chambers eurypylous, elongate-oval to tubular and occasionally branched (Bergquist, 1980).
Ianthella Gray, 1869 (type species: Ianthella flabelliformis Pallas, 1776) (syn. Basta Oken,
1815; Haddonella Sollas, 1903) - Planar or multiplanar fan shaped; fibrous skeleton highly developed
and makes up by far the great bulk of the sponge body; skeleton is reticulate structure typically
developed in a single plane to produce a regular fan shape; fibres have both bark and pith elements, and
bark contains many spongocytes arranged in concentric annuli (Bergquist, 1980).
q SUBCLASS CALCINEA.
DEFINITION: Regular triradiate spicules, equiangular and equiradiate or exceptionally parasagittal or
sagittal, and a basal system of quadriradiates; most species have at least some spicules with the rays and
angles between the rays being equal, with or without monactinal or diactinal free spicules; young
sponges may have only triradiate spicules; choanocytes basinucleate, nuclei spherical, and basal body of
flagellum not adjacent to the nucleus; larvae are entirely ciliated hollow blastula (coeloblastula).
With two orders.
q ORDER CLATHRINIDA.
DEFINITION: Skeleton composed exclusively of free spicules, without hypercalcified non-spicular
reinforcements or spicule tracts.
With six families.
Clathrina Gray, 1867 (type species: Grantia clathrus Schmidt, 1864) (syn. Ascetta Haeckel,
1872; Ascaltis Haeckel, 1872; Leucopsis Lendenfeld, 1885) - cormus is composed of anastomosing,
irregular reticulation of tubes, usually encrusting, sessile, low growing; smooth surface; oscules not
apparent; soft texture; choanoderm is flat or rarely raised up into conuli by the apical rays of the
quadriradiates, but never forming true folds, at least when the sponge is in the extended state; spicules
are regular equiangular and equiradiate triradiate and/or quadriradiates alone, but sometimes also with
biradiates or tripod spicules (Burton, 1963; Borojevic et al., 1990).
Guancha Miklucho-Maclay, 1868 (type species: Guancha blanca Miklucho-Maclay, 1868) -
cormus composed of a peduncle and a clathroid body, often composed of small budded colonies of
tubular individuals attached to stipitate base; peduncle may be formed by true tubes with a normal
choanoderm, or may be solid with a special skeleton; skeleton composed of regular equiangular and
equiradiate spicules, to which parasagittal triradiate spicules may be present at least in the peduncle; in
some species only parasagittal spicules are present and their unpaired ray is basipetally oriented (Burton,
1963; Borojevic et al., 1990).
GENERA:
Ascandra Haeckel, 1872 (type species: Ascandra falcata Haeckel, 1872) - solitary or
compound, subpyriform, clathrate, substipitate; surface minutely hispid; oscules apical; soft texture;
choanoderm forms folds inside the choanocoel which isolate radially arranged shallow cavities or true
radial tubes; choanosomal folds are supported by apical rays of external quadriradiates only; spicules
regular triradiates, regular quadriradiates and sickle-shaped oxeas (Burton, 1963; Borojevic et al.,
1990).
Leucaltis Haeckel, 1872 (type species: Leucaltis clathria Haeckel, 1872) (syn. Leucaltusa
Haeckel, 1872; Artynaltis Haeckel, 1872; Heteropegma Poléjaeff, 1884) - body composed of large
ramified and anastomosed clathrous tubes; surface even, smooth; oscules small, scattered; texture
friable; each tube has a distinct cortex, a choanoderm composed of elongated and ramified choanocyte
chambers and central atrium; choanoderm and atrial wall have a secondary skeleton composed of small
triradiates and quadriradiates; ectosomal skeleton of several tangential layers of regular triradiates with
facial rays of subectosomal regular quadriradiates; chamber layer with centipetally-directed apical rays
of subectosomal quadriradiates, regular to sagittal, and small irregularly scattered quadriradiates;
choanosomal skeleton with several tangential layers of quadriradiates (Burton, 1963; Borojevic et al.,
1990).
Leucettusa Haeckel, 1872 (type species: Leucetta corticata Haeckel, 1872) - composed of a
clathrate mass of anastomosing tubes; ectosomal skeleton of several tangential layers of regular
triradiates; skeleton of chamber layer and of choanosomal surfaces of small sagittal triradiates, with
those of inner parts of chamber layer with swollen ends to rays (Burton, 1963); simple tubular body a
large atrium and choanocyte chambers which are either elongated, spherical or both (Borojevic et al.,
1990).
Leuclathrina Borojevic & Boury-Esnault, 1987 (incertae sedis) (type species: Leuclathrina
asconoides Borojevic & Boury-Esnault, 1987) - skeleton restricted to the ectosomal cortical region,
and the choanosome lacks spicules completely (Borojevic et al., 1990); globular growth form, apical
oscules with slightly raised fringe; smooth, porous surface; ectosomal cortex supported by equiangular
and equiradiate triactines; choanosome without mineral skeleton; leuconoid aquiferous system
(Borojevic & Boury-Esnault, 1987).
Leucetta Haeckel, 1872 (type species: Leucetta primigenia Haeckel, 1872) (syn. Sycothamnus
Haeckel, 1869; Leucetta Haeckel, 1872; Teichonella Carter, 1878) - a homogeneous organisation of
the wall and a typical leuconoid aquiferous system; there is no clear distinction between the cortex and
the choanoskeleton, nor the presence of a distinct layer of subcortical inhalant cavities; the atrium is
frequently reduced to a system of exhalant channels that open directly into the osculum (Borojevic et al.,
1990); solitary or compound, tubular, clathrate with tubular oscules, or lobate; surface even, harsh;
oscules apical; firm texture; skeleton of regular to subregular triradiates arranged tangentially in
ectosomal and choanosomal region, irregularly arranged in chamber layer (Burton, 1963).
Pericharax Poléjaeff, 1884 (type species: Pericharax heterorhaphis Poléjaeff, 1884) - with a
large central cavity, probably a pseudogaster, surrounded by a thick wall; the wall is divided into a
choanoderm and a thin subcortical layer of inhalant cavities supported by a peculiar skeleton partially
composed of the centripetal rays of the special cortical triradiates (Borojevic et al., 1990); solitary or
compound, cup-shaped, subspherical or massive lobose; surface even, smooth, harsh often thrown into
irregular folds in places; apical oscule, with or without margin; firm texture; ectosomal skeleton a
tangential layer of subregular triradiates; skeleton of chamber layer of large and small regular triradiates
irregularly scattered; choanosomal skeleton a tangential layer of regular quadriradiates (Burton, 1963).
Leucomalthe Haeckel, 1872 (incertae sedis) (type species: Leucomalthe bomba Haeckel,
1872) - with a solid body and a large central cavity; choanocyte chambers greatly elongated, tubular and
copiously branched; skeleton consists of regular triradiates and microbiradiates in the cortical skeleton,
regular triradiates in the choanoskeleton and sagittal quadriradiates both in the atrial skeleton and in the
exhalant channels; large longitudinal biradiates present throughout the body (Borojevic et al., 1990);
solitary, spherical; surface even; apical oscule with well developed fringe; texture firm; ectosomal
skeleton of tangential layers of microxeas and regular triradiates; skeleton of chamber layer of regular
triradiates and longitudinally placed oxeas, with larger canals lined with subregular quadriradiates;
choanosomal skeleton a tangential layer of sagittal quadriradiates (Burton, 1963).
q ORDER MURRAYONIDA.
DEFINITION: Reinforcement of the skeleton composed of either spicule tracts, calcareous plates or a
rigid aspicular skeleton; canal system leuconoid; diapasons (tuning-fork shaped triradiates) or modified
biradiates present and generally fasciculated.
There are 3 families and only 3 known species..
DEFINITION: Choanosomal skeleton composed of bundles of biradiates without any rigid structure;
cortical skeleton composed chiefly of a tangential layer of tripods (triradiates) and curved oxeotes
(biradiates).
SCOPE: Single genus.
REVIEWS: Vacelet (1977, 1981), Hartman (1982).
GENERA:
Lelapiella Vacelet, 1977 (type species: Lelapiella incrustans Vacelet, 1977) - cortex
reinforced by large equiangular triradiates ornamented on the external convex side with mamelons, and
also with cortical diactinal spicules curved at 120° at the center. The basal layer, in contact with the
substrate, is made up of a dense network of curved oxeas. Tracts of oxeas run from the cortex diagonally
through the choanosome to the basal layer of spicules (Hartman, 1982).
q SUBCLASS CALCARONEA.
DEFINITION: Calcarea with incubated amphiblastula larvae flagellated only on the anterior half; nuclei
of choanocytes apical, and the flagellum arises directly from the nucleus; spicules are triradiate and
sagittal (two rays are paired and the third ray is longer than the others), as well as free monaxonic
(monactinal or diactinal) forms; aquiferous system ranges from asconoid to leuconoid grades of
construction.
Two orders recognized.
q ORDER LEUCOSOLENIIDA.
DEFINITION: Only with free spicules, without calcified non-spicular reinforcements.
With seven families.
Haeckel, 1872; Nardoraltis Haeckel, 1872; Nardorandra Haeckel, 1872; Nardoretta Haeckel, 1872;
Nardorilla Haeckel, 1872; Nardorortis Haeckel, 1872; Nardomandrum Haeckel, 1872; Nardopsandra
Haeckel, 1872; Nardopsetta Haeckel, 1872; Nardopsortis Haeckel, 1872; Tarraltis Haeckel, 1872;
Tarrandra Haeckel, 1872; Tarretta Haeckel, 1872; Tarrilla Haeckel, 1872; Tarromandra Haeckel,
1872; Tarropsandra Haeckel, 1872; Tarropsetta Haeckel, 1872; Auloplegmaltis Haeckel, 1872;
Auloplegmetta Haeckel, 1872; Auloplegmilla Haeckel, 1872; Auloplegmandra Haeckel, 1872;
Auloplegmortis Haeckel, 1872; Ascaltometra Haeckel, 1872; Ascandrometra Haeckel, 1872;
Ascettometra Haeckel, 1872; Homandra Lendenfeld, 1891; Nardoris Delage, 1899; Aulorhiza Dendy
& Row, 1913) - encrusting network of more or less erect asconoid tubes which may branch, or throw
out lateral diverticula, but seldom anastomose except occasionally at base of main tubes; surface
minutely hispid; texture soft; skeleton of sagittal triradiates and sagittal quadriradiates, with apical rays
projecting into cloacal cavity, and oxeas with lanceolate distal ends (Burton, 1963).
Jenkin, 1908; Tenthrenodes Jenkin, 1908) - tubular, spherical to elongate and cylindrical; surfaec
minutely papillate and hispid; apical oscule with ot without fringe; texture soft to firm; tubar skeleton of
basal rays of subgastral sagittal triradiates and several rows of tubar sagittal or subregular triradiates,
and ectosomal surface with distal cones ornamented with oxeas; choanosomal skeleton of paired rays of
subgastral sagittal triradiates and choanosomal sagittal or subregular triradiates and quadriradiates;
skeleton of chamber layer typically articulate and showing no marked differences in size between
radiates of ectosome, chamber layer and choanosome except where tubar skeleton is reduced to sagittal
subgastral radiates (Burton, 1963).
occasionally replaced by, oxeas; ectosomal cortex sometimes with quadriradiates in association with
choanosomal triradiates; skeleton of the chamber layer ranging from regularly articulate to irregularly
scattered, and typically with subgastral sagittal radiates; some subdermal pseudosagittal triradiates may
occur but these are derived from normal choanosomal spicules, and do not form a continuous distict
layer as in the Heteropiidae; subgastral quadriradiates, if present, always associated with chamber-layer
skeleton containing confused triradiates; nuclei of collared cells probably always apical; choanocyte
chambers asconoid, elongate and radially arranged, or small, spherical and irregularly scattered in the
choanosome (leuconoid).
SCOPE: This family contains 80 generic names, of which 21 are recognised here.
REVIEWS: Lendenfeld (1885d, 1885m), Dendy (1893a), Dendy & Row (1913), Tanita (1943),
Borojevic (1966).
GENERA:
Amphiute Hanitsch, 1894 (type species: Amphiute paulini Hanitsch, 1894) - tubular, solitary
or compound, apical oscule with fringe; ectosomal skeleton of large longitudinally arranged oxeas,
microxeas set at right angles to surface, a tangential layer of triradiates and paired rays of subectosomal
pseudosagittal triradiates; tubar skeleton of basal rays of subectosomal pseudosagittal triradiates and
subgastral sagittal triradiates; choanosomal skeleton of paired rays of subectosomal sagittal triradiates
and a tangential layer of large oxeas and choanosomal quadriradiates (Burton, 1963).
Anamixilla Poléjaeff, 1884 (type species: Anamixilla torresi Poléjaeff, 1884) - colonial
tubular, apical oscule; ectosomal skeleton a tangential layer of triradiates; tubar skeleton of
centrifugally-directed basal rays of subgastral sagittal triradiates, with irregularly arranged tubar
triradiates; choanosomal skeleton of paired rays of subgastral sagittal triradiates and a tangential layer of
triradiates and quadriradiates (Burton, 1963).
Aphroceras Gray, 1858 (type species: Grantia ensata Bowerbank, 1858) (syn. Leucogypsia
Bowerbank, 1864; Cyathiscus Haeckel, 1870; Leucandrena Haeckel, 1872; Artynandrus Haeckel,
1872) - tubular, elongate, apical oscule; skeleton of chamber layer ranging from more or less confused
to articulate, with subgastral or other sagittal radiates; ectosomal skeleton of tangentially placed
triradiates supplemented typically by colossal oxeas placed longitudinally, or less commonly projecting
from the surface (Burton, 1963).
Ebnerella Lendenfeld, 1891 (type species: Ebnerella schulzei Breitfuss, 1898) - with syconoid
organisation; choanosomal skeleton reduced to the unpaired actine of the subatrial spicules; cortex
devoid of longitudinal diactines (Borojevic, Boury-Esnault & Vacelet, in prep.).
Eilhardia Poléjaeff, 1884 (type species: Eilhardia schulzei Poléjaeff, 1884) - solitary cup-
shaped, stipitate, sub-papillate oscules on outer surface of cup only; with large triradiates irregularly
arranged in chamber layer, large triradiates and oxeotes in the ectosomal layer, with a tangential layer of
smaller radiates, and with microxeas or trichodragmata scattered in one or other of principal layers
(Burton, 1963).
Grantia Fleming, 1828 (type species: Spongia compressa Fabricius, 1780) (syn. Sycortis
Haeckel, 1872; Sycortusa Haeckel, 1872; Vosmaeria Lendenfeld, 1885; Dermatreton Jenkin, 1908;
Teichonopsis Dendy & Row, 1913; Paragrantia Hozawa, 1940) - solitary, sac-shaped, laterally
compressed, hispid, apical or marginal oscule; ectosomal skeleton a tangential layer of triradiates, with
tufts of oxeas projecting beyond surface; tubar skeleton of centripetally-directed basal rays of subgastral
sagittal triradiates and numerous rows of tubar triradiates; choanosomal skeleton of paired rays of
subgastral triradiates and a tangential layer of choanosomal triradiates and quadriradiates (Burton,
1963).
Hypograntia Carter, 1886 (type species: Hypograntia infrequens Carter, 1886) (syn.
Grantiopsis Dendy, 1893; Hippograntia Breitfuss, 1898) - tubular, roughened, apical oscule; with
ectosomal cortex as thick as the chamber layer, with many layers of tangential triradiates; tubar skeleton
articulate, composed of the basal rays of subgastral quadriradiates with which are associated sagittal
triradiates practically reduced to the basal ray by suppression of the paired rays; choanosomal skeleton of
the paired and apical rays of the subgastral quadriradiates together with a tangential layer of
choanosomal quadriradiates; without colossal longitudinal oxeotes (Burton, 1963).
Jenkina Brondsted, 1931 (type species: Jenkina cirrata (Jenkin, 1908)) - with a sylleibid or
leuconoid organisation; choanosomal skeleton reduced to the unpaired actine of subatrial spicules;
cortex without longitudinal diactines (Borojevic, Boury-Esnault & Vacelet, in prep.)
Lamontia Kirk, 1894 (type species: Lamontia zona Kirk, 1894) - tubular, hispid, apical oscule
with fringed margin; pores in special area usually subterminal equatorial, but may be irregular
longitudinal; skeleton of chamber layer consisting of small scatterd oxeotes; ectosomal cortex with
triradiates in addition to oxeotes; choanosomal quadriradiates present; sponge consisting of a single
olynthus with a specialised pore-zone below the terminal vent (Burton, 1963).
92 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
Leucandra Haeckel, 1872 (type species: Leucandra egedii (Schmidt, 1870) - with a leuconoid
organisation; internal diactines go through the cortex and keep their free part outside the sponge
(Borojevic, Boury-Esnault & Vacelet, in prep.).
Leuconia Grant, 1832 (type species: Spongia nivea Grant, 1826) (syn. Medon Duchassaing &
Michelotti, 1864; Sycinula Schmidt, 1868; Dyssyconella Haeckel, 1870; Dyssycum Haeckel, 1870;
Coenostomella Haeckel, 1870; Dyssiconella Wright, 1870; Leucortis Haeckel, 1872; Leucaltaga
Haeckel, 1872; Mlea Haeckel, 1872; Leucandraga Haeckel, 1872; Leucandrusa Haeckel, 1872;
Dyssycarium Haeckel, 1872; Coenostomium Haeckel, 1872; Dyssycortus Haeckel, 1872; Dyssycandrus
Haeckel, 1872; Dysscaltella Haeckel, 1872; Dyssycandrella Haeckel, 1872; Dyssycandrium Haeckel,
1872; Lipostomaltis Haeckel, 1872; Lipostomandra Haeckel, 1872; Lipostomortis Haeckel, 1872;
Amphoriscandra Haeckel, 1872; Amphoricortis Haeckel, 1872; Amphoriulandra Haeckel, 1872;
Amphoriidandra Haeckel, 1872; Coenostomandra Haeckel, 1872; Coenostomandium Haeckel, 1872;
Coenostomellium Haeckel, 1872; Coenostomortis Haeckel, 1872; Artynandrium Haeckel, 1872;
Artynellandra Haeckel, 1872; Artynortus Haeckel, 1872; Aphrocerandra Haeckel, 1872; Aphrocerortis
Haeckel, 1872; Leucortmetra Haeckel, 1872; Leucania Lendenfeld, 1885; Dyssicarium Delage, 1899) -
encrusting, smooth surface, small scattered oscules; friable texture; ectosomal skeleton of several
tangential layers of triradiates; skeleton of chamber layer supported by large irregularly scattered radiates
and diacts (microxeas) irregularly arranged; choanosomal skeleton and linings of exhalant canals a
tangential layer of sagittal quadriradiates (Burton, 1963).
Leucopsila Dendy & Row, 1913 (type species: Leuconia stilifera Schmidt, 1870) - tubular,
laterally compressed, single or colonial, apical oscule with thin undulating margin; smooth conulose or
even surface; soft texture; ectosomal skeleton of several layers of tangential triradiates with numerous
microxeas; skeleton of chamber layer a confused mass of microxeas with large quadriradiates scattered
between; choanosomal skeleton a dense layer of microxeas (Burton, 1963).
Leucyssa Haeckel, 1872 (type species: Leucyssa spongilla Haeckel, 1872) - clathrate mass of
anastomosing tubes; even surface, non-hispid; apical oscules with strongly fringed margins; skeleton of
body of oxeas only, with larger oxeas surrounding oscules (Burton, 1963).
Paraleucilla Dendy, 1892 (type species: Leucandra cucumis Haeckel, 1872) - ovate body,
even surface, non-hispid, apical oscules, naked or fringed margins; ectosomal skeleton a tangential layer
of triradiates and oxeas; subectosomal skeleton of two layers of quadriradiates with apical rays
oppositely directed; skeleton of chamber layer of scattered quadriradiates; choanosomal skeleton a
tangential layer of triradiates (Burton, 1963).
Sycophractus Haeckel, 1872 (syn. Sycurandra Haeckel, 1872; Ute, of authors) (type species:
Ute glabra Schmidt, 1864) - [based on S. spenceri (Dendy, 1892:94): spherical or subspherical; uneven
surface, roughened, apical oscule; firm; ectosomal skeleton of longitudinal oxeas, tangential triradiates
and groups of microxeas; skeleton of chamber layer of an outer tubar skeleton of triradiates and an inner
irregular layer of triradiates and quadriradiates of exhalant canals; choanosomal skeleton of several
tangential layers of quadriradiates, and sparsely scattered microxeas (Burton, 1963).
Sycute Dendy & Row, 1913 (type species: Sycon dendyi Kirk, 1895) - tubular, hispid surface
in tufts; apical oscule with fringed margin; ectosomal skeleton of longitudinally placed oxeas, with tufts
of small oxeas; skeleton of chamber layer of triradiates; choanosomal skeleton of quadriradiates
(Burton, 1963).
Sycyssa Haeckel, 1872 (type species: Sycyssa huxleyi Haeckel, 1872) - oval, even surface
strongly hispid; apical oscule with fringed margin; ectosomal skeleton a tangential layer of oxeas, with a
palisade of oxeas set at right angles to the surface, and with large oxeas projecting beyond; skeleton of
chamber layer of proximal parts of large oxeas; choanosomal skeleton a tangential layer of oxeas and a
subgastral layer of large oxeas (Burton, 1963).
Synute Dendy, 1892 (type species: Synute pulchella Dendy, 1892) - with a cormus constituted
by completely fused units with a syconoid organisation, surrounded by a common cortex with a skeleton
of giant longitudinal diactines (Borojevic, Boury-Esnault & Vacelet, in prep.).
Teichenopsis Dendy & Row, 1913 (type species: Teichonella labyrinthica Carter, 1878) -
caliculate growth form when small, becoming infundibular and compressed, to lamellate and much
folded, stipitate in all but early stages; even surface, non-hispid; oscules small and numerous on inner
surface; texture firm and brittle; ectosomal skeleton of several layers of triradiates, with microxeas set at
right angles to surface; tubar skeleton of sugastral sagittal triradiates and numerous rows of tubar
triradiates; choanosomal skeleton of paired rays of subgastral triradiates, a tangential layer of
quadriradiates and microxeas set at right angles to surface (Burton, 1963).
Trichogypsia Carter, 1871 (type species: Trichogypsia villosa Carter, 1871) (syn. Dyssycyssus
Haeckel, 1872; Amphoriscyssa Haeckel, 1872; Aphroceryssa Haeckel, 1872; Sycolepis Lendenfeld,
1885) - encrusting to irregular massive growth forms; surface uneven, irregular, minutely hispid;
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 93
oscules small, scattered; with skeleton composed entirely of distally spined diactines (oxeas) only; canal
system leuconoid (Burton, 1963).
Ute Schmidt, 1862 (type species: Ute glabra Schmidt, 1864) - with leuconoid organisation;
cortex sustained by giant longitudinal diactines; choanosomal skeleton articulated; no radial fascicules
of diactines (Borojevic, Boury-Esnault & Vacelet, in prep.).
Uteopsis Dendy & Row, 1913 (type species: Ute argentea Poléjaeff, 1883) - solitary, tubular,
substipitate; smooth surface, striated; apical oscule; firm texture; ectosomal cortex well developed, with
skeleton composed of triradiates and containing oxeotes and microxeas arranged tangentially; tubar
skeleton of centrifugally-directed basal rays of subgastral sagittal triradiates, tubar quadradiates and
bundles of distally directed oxeas; choanosomal skeleton of paired rays of subgastral sagittal triradiates,
an inner layer of choanosomal triradiates and quadriradiates, and an outer layer of choanosomal
quadriradiates, with microxeas irregularly scattered (Burton, 1963).
apical rays of subgastral sagittal quadriradiates and a tangential layer of choanosomal subregular
quadriradiates; skeleton of root tuft of oxeas and anchoring radiates (Burton, 1963) [incertae sedis].
q ORDER LITHONIDA.
DEFINITION: Generally restricted to shaded habitats such as caves and tunnels; massive reinforced
calcitic (hypercalcified) skeleton, together with tuning fork spicules and saggital tetractines as free
spicules; larvae are amphiblastula; choanocytes are apinucleate.
Three families.
q SUBPHYLUM SYMPLASMA
q CLASS HEXACTINELLIDA (Glass Sponges).
DEFINITION: Skeleton composed of six-rayed siliceous spicules (hexacts), occurring individually or
fused together, usually forming rigid lattice-like skeletons; body wall has a cavernous structure, with
living tissue stretching across a framework around the cavities like a membrane; this tissue is syncytial,
on both the dermal region (pinacoderm) and in the choanosome, in which the multinucleolate
96 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
protoplasm is not divided into cells; uniflagellated choanocytes are absent from this class of sponges,
and the choanocytes are really only collar-flagellum units lining cylindical chambers (hence they are
referred to as "flagellated chambers" rather than "choanocyte chambers" as in the classes Calcarea and
Demospongiae); these unusual choanocytes are embedded in the membraneous protoplasm stretched
between spicules by "plugged bridges"; spicules occur in three different regions, and the localization of
particular spicule types to particular areas is very precise; three zones differentiated: (1) lying on or just
below the dermal membrane (dermal); (2) lying within the trabeculae (parenchymal); (3) lying below the
membrane around the atrial cavity (gastral); diverse geometry of megascleres and microscleres; unlike
other classes of sponges axial canals of spicules are always square in cross-section; larvae are incubated
parenchymella.
Two subclasses and four orders.
REVISIONS: Ijima (1927), Lévi (1964), Reiswig (1992).
q SUBCLASS AMPHIDISCOPHORA.
DEFINITION: With birotulate microscleres but lacking hexaster microscleres; sponges embedded in soft
sediments by single or tufts of basal monactine spicules, not attached directly to substratum; flagellated
chambers are continuous at their openings, not sharply marked off from each other as in other classes of
sponges.
One Recent order containing three families.
q ORDER AMPHIDISCOSIDA
As for subclass.
top of the body around the columella or gastral cone and are sharply set off from the inhalant surface by
the oscular margin; four separate exhalant canals may open around the columella, or the entire exhalant
region may be either inwardly depressed or outwardly bulging to form a "gastral" cavity, sometimes
covered by a lattice-like sieve plate; neither uncinate spicules nor scepters are present; marginal prostals
are pinular rhabdodiactines (i.e. diactinal with the distal end spined); pleural prostals are smooth
diactines; choanosomal supporting spicules are mostly rhabdodiactines, often occurring in association
with macrohexactines or macropentactines.
SCOPE: This family contains 16 nominal genera, but probably only four of these are valid.
REVIEWS: Ijima (1927)
GENERA:
Chalaronema Ijima, 1927 (type species: )
Compsocalyx Schulze, 1904 (type species: )
Hyalonema Gray, 1832 (type species: Hyalanema sieboldi Gray, 1835) (syn. Coscinonema
Ijima, 1927; Cyliconema Ijima, 1927; Euhyalonema Ijima, 1927; Leptonema Lendenfeld, 1915;
Pteronema Ijima, 1927; Paradisconema Ijima, 1927; Thamnonema Ijima, 1927; Phialonema
Lendenfeld, 1915; Prionema Lendenfeld, 1915; Corynonema Ijima, 1927; Onconema Ijima, 1927;
Oonema Lendenfeld, 1915)
Lophophysema Schulze, 1904 (type species: )
q SUBCLASS HEXASTEROPHORA.
DEFINITION: Hexasters microscleres present, birotulate microscleres absent; growth forms are diverse,
usually fixed to substratum by a basal attachment; basal spicules, when present, consist of pentactines or
anisodiactines usually in tufts.
Three extant orders and twelve families are recognized.
q ORDER HEXACTINOSIDA.
DEFINITION: Rigid parenchymal skeleton produced by fusion of hexactines; dermal and gastral spicules
usually pentactines, with the unpaired ray directed inwards, or sometimes stauractines, and these spicules
are usually connected by tissue only.
Six families.
REVIEWS: and revisions: Ijima (1898, 1903, 1927), Reid (1957, 1963); Reiswig (1990, 1991).
q ORDER LYCHNISCOSIDA.
DEFINITION: Sponges firmly attached to substrata; parenchymal megascleres lychniscs, or derivatives,
united in a rigid framework; central part of each spicule surrounded by twelve struts arranged like the
edge of an octahedron.
Two families.
q ORDER LYSSACINOSIDA.
DEFINITION: Parenchymal megascleres vary from hexactines to rhabdodiactines, usually occurring free
in tissues, sometimes secondarily fused to form rigid framework; dermal spicules consist of a single
layer of large pentactines or hexactines, with single, long, proximal ray directed inwards, or with a layer
100 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
of small dermal spicules overlying larger hypodermal pentactines, with the unpaired ray extending
inwards.
Four extant families are recognized.
covering over the sponge; parenchymal spicules hexactines and/or rhabdodiactines; microscleres oxy-
and discohexasters, sometimes discoctasters.
SCOPE: Twenty five genera are included in the family, although not all these may be truly valid. Three
subfamilies are recognised.
REVIEWS: Schulze (1887, 1897, 1899), Ijima (1927).
GENERA:
Subfamily Lanuginellinae Schulze, 1897 - without discoasters, with strobiloplumicomes.
Calycosoma Schulze, 1899 (type species: )
Lanuginella Schmidt, 1869 (type species: )
Lanugonychia Lendenfeld, 1915 (type species: )
Lophocalyx Schulze, 1887 (type species: )
Mellonympha Schulze, 1897 (type species: )
Lithobactrum Kirkpatrick (type species: ) - oxeas, styles and desmas; "Lithistida" of uncertain
placement
Lithospongia Duchassaing & Michelotti - unrecognisable (de Laubenfels, 1936)
Manon Oken - unrecognisable (de Laubenfels, 1936)
Microxistyla Topsent (type speces: ) - microscleres centrotylotes diacts; possibly
Desmacellidae.
Monotria de Laubenfels, 1936 (type species: Coppatias solidissima Wilson, 1902) - simple
spiculation of oxeas and triods which are plausibly calthrops having lost 1 of 4 rays; possibly
Pachastrellidae.
Mycalopsis Topsent (type species: ) - similar to Esperiopsis but with 3 sizes of chelae the
largest of which occur in rosettes; possibly synonym of Esperiopsis.
Nailondria de Laubenfels, 1954 (type species: Nailondria maza de Laubenfels, 1954) -
amorphous, soft, soggy texture; ectosome detachable with tangential layer of mainly styles, and
choanosome cavernous with strongyles and styles in halichondroid arrangement; probably Halichondria.
Neothenea de Laubenfels, 1936 (type species: Neothenea enae de Laubenfels, 1934) -
microscleres metasters and bent microrhabds that may be reduced asters; possibly Ancorinidae.
Nisella Johnson (type species: ) - with tylasters and normal euasters; possibly Ancorinidae.
Oopsacas Topsent (type species: ) - probably in Leucopsacasidae.
Ophiraphidites Carter, 1876 - unrecognisable (de Laubenfels, 1936).
Ophistospongia Gray, 1867 (type species: Ophistospongia australis Gray, 1867) - virtually
unrecognisable (de Laubenfels, 1936).
Orthorachis Gray - unrecognisable (de Laubenfels, 1936).
Oxeosarcodea de Laubenfels, 1954 (type species: Oxeosarcodea oinops de Laubenfels, 1954)
- massive, cheesy gelatinous consistency; surface slightly microconulose without any obvious separable
ectosome; choanosome gelatinous perforated by meandering canals, spiculation only of thin oxeas
scattered and in few vague tracts without apparent associated spongin; megascleres predominantly
oxeote but with occasional strongylote modifications; possibly Axinellidae reminiscent of Axsinyssa.
Parafieldingia Milne-Edwards - unrecognisable (de Laubenfels, 1936)
Pellinula Czerniavsky - virtually unrecognisable (de Laubenfels, 1936)
Pharetronema Sollas - unrecognisable (de Laubenfels, 1936)
Pityrete Topsent (type species: ) - probably belongs to Euretidae (de Laubenfels, 1936).
Pleorhabdus Schulze (type species: ) (syn. [Polyrhabdus] Schulze (preocc.) - possibly
Caulophacidae.
Protoclathria Burton (type species: ) - acanthostrongyles echinating fibres instead of styles; no
microscleres; uncertain validity possibly synonym of Antho.
Protoschmidtia Czerniavsky, 1880 (type species: Protoschmidtia simplex Czerniavsky, 1880).
Psammina Haeckel, 1889 (type species: Psammina nummulina Haeckel, 1889) - virtually
unrecognisable (de Laubenfels, 1936).
Psammophyllium Haeckel, 1889 (type species: Psammophyllum flustraceum Haeckel, 1889) -
virtually unrecognisable (de Laubenfels, 1936)
Pseudochalina Schmidt, 1870 (no type species named).
Quepanetsal de Laubenfels, 1954 (type species: Quepanetsal madidus de Laubenfels, 1954) -
encrusting, soft crumbly consistency, oscules contractile; ectosome with tangential skeleton easily
peeled; choanosome bread-like with spicules dispersed in confusion; 2 categories of megascleres shorter
thick strongyle and longer thin oxeas with microspined points, not restricted to any particular location
within the skeleton; no microscleres; possibly related to Desmoxya or Halichondria.
Quintoxilla (type species: )
Quixilla de Laubenfels, 1936 (type species: Hymeraphis lacazii Topsent, 1891) - ectosomal
spicules with hastate modifications to both ends, few tylostyles and echinating acanthostyles; no
microscleres; possibly Hymedesmiidae.
Rhabdodragma Dendy (type species: ) - with spiny euasters, spiny microrhabds and others that
are smooth; possibly Ancorinidae.
Rhabdostauridium Schmidt, 1880 (type species: Rhabdostauridium retortula Schmidt, 1880)
- unrecognisable (de Laubenfels, 1936).
Rhacodiscula Zittel, 1878 (type species: Rhacodiscula asteroides Zittel, 1878) -
phyllotriaenes oxeas and desmas, microscleres euasters and streptasters; uncertain "Lithistida" affinities.
Rhaphidhistia Carter (type species: ) - principal spicules oxeas, microscleres spirasters;
possibly Spirastrellidae.
Ridleia Dendy (type species: ) - peculiar flask shaped hollow growth form; smaller ectosomal
tylostyles overlaying larger choansomal tylostyles; probably Suberitidae.
Sarcomella Schmidt - virtually unrecognisable (de Laubenfels, 1936)
Sceptrintus Topsent (type species: ) - with a few spiny styles and 2 sizes of rhabds with their
spines arranged nodally; possibly related to Agelasiidae (de Laubenfels, 1936).
Sclerochalina Schmidt, 1868 (type species: Sclerochalina asterigena Schmidt, 1868).
Scleroplegma Schmidt (type species: ) - possibly Euretidae (de Laubenfels, 1936).
Scutastra Ferrer-Hernandez (type species: ) - like Penares but has peculiar euaster with
exceedingly numerous rays, and smooth diactinal microscleres; probably Ancorinidae (de Laubenfels,
1936).
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 105
Scyphia Oken, 1814 (type species: Scyphia scyphiformis Oken, 1814) - unrecognisable (de
Laubenfels, 1936).
Seliscothon Zittel (type species: ) - erect lamellate growth form; only oxeas and desmas;
uncertain affinities within "Lithistida".
Setidium Schmidt (type species: ) - cup-shaped growth form; uncertain "Lithistid" affinities.
Siphonidiella Burton (type species: ) - only dichotriaenes and desmas, but ectosomal desmas
differ from those in the choansomal skeleton; uncertain affinities within "Lithistida".
Siphydora James Clark - unrecognisable (de Laubenfels, 1936).
Solina Gray - virtually unrecognisable (de Laubenfels, 1936)
Somatispongia Bowerbank - unrecognisable (de Laubenfels, 1936).
Spiroxya Topsent (type species: ) - megascleres oxeas, microscleres spinaespirae-like sigmoids
and microstrongyles with spines arranged around them; possibly Spirastrellidae.
Spongocardium Kirkpatrick (type species: ) - peculiarly shaped sigmoid microscleres; possibly
Spirastrellidae.
Stannarium Haeckel, 1889 (type species: Stannarium concretum Haeckel, 1889); virtually
unrecognisable (de Laubenfels, 1936).
Stannoma Haeckel, 1889 (type species: Stannoma dendroides Haeckel, 1889); virtually
unrecognisable (de Laubenfels, 1936).
Stannophyllum Haeckel, 1889 (type species: Stannophyllum zonarium Haeckel, 1889);
virtually unrecognisable (de Laubenfels, 1936).
Stellogeodia Czerniavsky (type species: ) - virtually unrecognisable, possible affinity with
Geodia (de Laubenfels, 1936).
Stylospira de Laubenfels, 1934 (type species: Stylospira mona de Laubenfels, 1934) - peculiar
spirally twisted styles, raphide microscleres; possibly Bubariidae.
Sulcastrella Schmidt (type species: ) - strongyles and tricrepid desmas; uncertain placement
within the "Lithistida".
Symplectella Dendy (type species: ) - possibly Euplectellidae.
Sympyla Sollas (type species: ) - vase-shaped with raised pore areas; tylotes and desmas;
uncertain "Lithistid" affinities.
Technitella Norman - unrecognisable, probably not a sponge (de Laubenfels, 1936)
Tragium Oken, 1814 (type species: Tragium solida Oken, 1814) - unrecognisable (de
Laubenfels, 1936).
Tremaulidium Schmidt (type species: ) - desmas, tylotes slightly spined at each end, tylostyles
with only heads spined resembling plocamiid sponges (de Laubenfels, 1936); uncertain "Lithistid"
affinities.
Tretolophus Sollas (type species: ) - desmas, oxeas, styles; uncertain "Lithistid" affinities.
Tubulodigitus Carter (type species: ) - intermediate between Adocia and Callyspongia;
uncertain affinity (de Laubenfels, 1936).
Uliczka de Laubenfels, 1936 (type species: Cinachyra schistospiculosa Uliczka, 1929) -
peculiar double-ended protriaenes; probably synonym of Cinachyra.
Uncinatera Topsent (type species: ) - referred to same group as Hexactinella; possibly
Tretodictyidae (de Laubenfels, 1936).
Valentis de Laubenfels, 1936 (type species: Desmacidon lentis Vosmaer, 1880) - ectosomal
spicules strongyles, choanosomal spicules tylostyles; microscleres include palmate and tridentate forms;
possibly Myxillidae.
Vazella Gray (type species: Holtenia pourtolesi Schmidt, 1870) - unrecognisable (de
Laubenfels, 1936).
Xylospongia Gray (type species: ) - unrecognisable (de Laubenfels, 1936).
Xytopsoocha de Laubenfels, 1936 (type species: Gellius macrosigma Topsent, 1890) - tylote
megascleres, enormous sigmas, medium sigmas, raphides; uncertain affinity, possibly Anchinoidae.
Ysiphonia de Laubenfels, 1936 (no type species named).
We prefer to receive specimens frozen if at all possible (and posted via courier, packed in dry
ice), given that subsequent genetic studies can then be undertaken on these species. However, outside
Australia this is difficult and expensive, in which case the following techniques should be used.
Ideally sponges should be frozen (to fix soluble pigments), and subsequently transferred to 70-
80% ethanol. Do not use formaldehyde. Specimens should be kept in separate plastic bags, each with a
label stating as a minimum a unique field/registration number.
Live specimens may alternatively be placed directly in 80-90% ethanol (they dilute themselves
to about 70-80%); but beware, soluble pigments from darker species may leach into and discolour
lighter coloured species if placed in the same container.
Specimens should be fixed and preserved for at least 48 hours prior to postage. Excess alcohol
shouls be drained off; delicate specimens should be wrapped in cheese-cloth if damage is possible; and
specimens heat-sealed in several layers of plastic bag.
q 1.4. Postage.
Preserved specimens posted from overseas should contain the words: SCIENTIFIC SAMPLES,
NO COMMERCIAL VALUE, CONTENTS MARINE SPONGES FOR TAXONOMY, or some such
disclaimer.
9. RECOMMENDED READING
Ackers, R.G., Moss, D. & Picton, B. (1992). Sponges of the British Isles ("Sponge V"). A colour
guide and working document. Marine Conservation Society, 9 Glouster Rd., Ross-on-Wye,
Herefordshire, HR9 5BU. 1992 Edition. Pp 1-175. [Provides a good glossary, illustrations of
characters, and examples of descriptions].
Bergquist, P.R. (1978). Sponges. Hutchinson: London. Pp 1-268. [This is still the best general text
available on Porifera, even though it is now out of date for much of the taxonomy and some theories on
sponge biology].
Brien, P., Lévi, C., Sarà, M., Tuzet, O & Vacelet, J. (eds) (1973). Spongiaires. Traité de Zoologie.
Anatomie, Systématique, Biologie. Masson et Cie: Paris. Volume 3. Pp 1-716. [This is a much more
detailed, specialist treatise on sponge biology, but contains good general information on many aspect of
the phylum. Once again, though, much is out of date].
Hartman, W.D. (1982). Porifera. Pp. 640-666 In: Parker, S.P. (ed.) Synopsis and Classification of
Living Organisms. New York: McGraw-Hill. Volume 1. [The first comprehensive description of the
family level classification for the Porifera (although the Calcarea section follows Burton (1963) and is
completely rejected). Major revisions of some orders (Poecilosclerida, Halichondrida) have been
subsequently undertaken and thus this work is now partially out of date].
Hooper, J.N.A. & Wiedenmayer, F. (1994). Porifera. In: Wells, A. (ed.) Zoological Catalogue of
Australia. AGPS: Canberra. Volume 12. Pp 1-621. [This provides family diagnoses and discussions
on sponge taxonomy, in a contemporary framework, as it relates to the Australian fauna only. Provides
biographical and bibliographical information on every named Australian species (although only 30%
have yet been described), but not much use for identifying below the family level].
Hooper, J.N.A. (1993). Phylum Porifera. Sponges. In: Mather, P. & Bennett, I. (eds) A Coral Reef
Handbook. Surrey Beatty & Sons: Sydney. Third Edn. Pp 35-46. [This is an overview of sponges in
coral reef habitats in particular, and should be treated as no more than a general introduction to sponge
taxonomy].
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 107
10. GLOSSARY.
radiate - condition of skeletal architecture whereby spicules are oriented radially from the centre (axis)
of the sponge; sometimes only visible at the surface; also known as radial or choristid structure.
renieroid - condition of skeletal architecture in which there is a regular reticulation of single (or few)
spicules forming square meshes, joined together at their ends (nodes) by an accretion of
collagenous spongin.
reticulate - condition of skeletal architecture whereby spongin fibres and/or spicule columns branch and
rejoin (anastomose) with each other to form 2- or 3-dimensional meshes.
sclerocyte - anucleolate motile secretory cell which produces spicules.
secondary fibre or tract - minor fibre or spicule tract interconnecting the ascending primary fibres or
tracts (cf. primary).
sedentary animals - (or sessile) animals in which the adults do not move by usual forms of locomotion
(cilia, pseudopods, legs, etc.), but usually live attached to the substrate. Adult sponges are
typically sedentary, althoug their larvae swim through the water column using cilia and/or
flagellae.
spicule - discrete element of the skeleton, usually mineralized (silica or calcite), produced by
sclerocytes; divided into two categories based on size (megasclere and microsclere).
spongin - proteinaceous material composed of collagen, forming the organic intercellular matrix
(collagenous filaments or spongin type A), and organic skeleton (spongin fibres or spongin type
B).
spongin fibre - macroscopic collagenous structures made up of many small microfibrils bound together,
producing discrete stands or plaques; fibres may be homogeneous (e.g. Spongiidae), have a light
central pith (e.g. Thorectidae) or a granular medullary portion (e.g. Verongida); fibres
frequently contain the mineralized secreted products of the sponge (spicules) and/or foreign
particles (e.g. arenaceous species); also known as the organic skeleton.
spongocytes - motile nucleolate cells that secrete spongin fibres.
subectosomal spicules - condition where megascleres are localized to a region below the ectosomal
skeleton but not associated with fibres or primary skeletal tracts (cf. ectosomal, choanosomal
spicules).
subisodictyal - condition of skeletal architecture similar to isodictyal reticulation but where meshes
have 2 or more spicules per side.
syconoid construction - body plan produced by folding of both the exterior (pinacoderm) and interior
(choanoderm) walls, such that choanocyte chambers lie within the body wall, and chambers
open directly onto the atrium (Calcarea).
syncytial - where the living "tissue" on the external surface (pinacoderm) or within the internal surface
(choanoderm) consists of an acellular protoplasm with multiple nuclei, usually stretched across
a skeletal framework (Hexactinellida).
tangential - group of fibres or spicules arranged parallel to the surface.
tetract, tetractinal - spicules with 4 rays (found in some Demospongiae, Hexactinellida and Calcarea).
tetraxonid - spicule with 4 rays each containing a central axis; also known as tetraxonic (cf. monaxonid,
triaxonid).
trabeculae - fibre, tract or bundle of spicules with angular cross section (Demospongiae); spicular
framework forming the hexactinellid skeleton, also known as "plugged bridges", across which
syncytial "tissue" is stretched (Hexactinellida).
triact - spicule with 3 rays (common in Calcarea).
triaxonid - spicule with 3 perpendicularly intersecting axes; also known as triaxonic (cf. monaxonic,
tetraxonic).
type locality - original locality from which the original specimen of the species, called the holotype, was
described.
unispicular - single row of spicules in a tract, fibre or reticulation (e.g. isodictyal) (cf. multispicular).
vivipary - method of sexual reproduction whereby the female sponge takes in sperm from another
sponge via the inhalant aquiferous system, eggs are fertilized and the ciliated (parenchymella)
larvae are brooded within the female sponge, and fully developed larvae, usually well-
differentiated cytologically, are subsequently released into the seawater (cf. ovipary).
11. ILLUSTRATIONS
Figures 5-230 refer to: Hooper, J.N.A. & Wiedenmayer, F. (1994). Porifera. In: Wells, A. (ed.)
Zoological Catalogue of Australia. AGPS: Canberra. Volume 12. Pp 1-621. Other figures of growth
forms, surface features, spongin fibres, and skeletal structures are taken recent publications such as
Boury-Esnault & Ruetzler (1997) and Wiedenmayer (1977).
DEMOSPONGIAE 8 strongyloxea
FIGS 5 - 65: Geometric forms of megascleres 9 sinuous strongyle
and calthrops 10 anisoxea
Diactinal (monaxonic) Megascleres 11 acanthose oxea
5 oxea 12 sinuous oxea
6 strongyle 13 tuberculate vermiform strongyle
7 tylote 14 acanthotylostrongyle
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 111
HEXACTINELLIDA
FIGS 184 - 230: Geometric forms of
megascleres and microscleres
Microscleres
184 sarule (sceptule)
185 spinoanchorate clavule (sceptule)
186 scopule (sceptule)
187 lonchiole (sceptule)
188 pilate clavule (sceptule)
189 disc-ended clavule (sceptule)
Megascleres
190 normal tylostyle
191 amphiox
192 uncinate
193 terminal end of basal bidentate with anchor
194 diactinal acanthophore
195 diactinal rhabd
196 sinuous rhabd
197 spiny-rayed acanthophore
198 hypodermal pentactine
199 tetractinal acanthophore
200 pentactinal pinule
201 hexactin
202 stauractin
203 triactin
204 superficial pentactine
205 dermal pinule
206 microhexactine
Microscleres
207 strongylhexaster
208 oxydiaster
209 oxyhexaster
210 discohexaster
211 discoctaster
212 codonhexaster
213 tylohexaster
214 graphiocome
215 floricome
216 plumicome
217 discocome
218 hexadisc
219 birotulate amphidisc
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 113
G Hamigera.......................................................48
HAPLOSCLERIDA. ..................................71
Gapoda..........................................................29
Hastatus.........................................................56
Gastrophanella.............................................38
Helophloeina ................................................59
Geelongia ......................................................81
Hemectyon.....................................................46
Gelliodes........................................................73
Hemectyonilla ...............................................45
Gellius............................................................72
Hemiasterella................................................31
Geodia ...........................................................27
HEMIASTERELLIDAE ............................31
Geodia (Cydonium) ......................................27
Hemigellius ...................................................73
Geodia (Isops)...............................................27
Hemihaliclona ..............................................73
Geodia (Sidonops)........................................27
Hemimycale...................................................57
GEODIIDAE................................................27
Hemitedania..................................................58
Geodinella.....................................................27
Herengeria ....................................................37
Gilchristia ...................................................104
Hertwigia.....................................................102
Gomphostegia...............................................62
Heterectya......................................................45
Goreauiella ...................................................40
Heterochone................................................100
Grantessa ......................................................91
Heteroclathria...............................................43
Grantia ..........................................................92
Heterocliona .................................................30
GRANTIIDAE .............................................92 Heterocornulum............................................41
Grantilla........................................................91 Heteromeyenia ..............................................75
Grantiopsis....................................................92 Heteronema...................................................80
Grapelia ........................................................62 Heteropegma.................................................87
Grayax ...........................................................48 Heterophymia..............................................104
Grayella.........................................................52 Heteropia.......................................................91
Grella.............................................................52
HETEROPIIDAE........................................91
Guancha ........................................................86
Heterorete......................................................99
Guitarra.........................................................60
Heterorotula..................................................75
GUITARRIDAE ..........................................60 Heterotella...................................................102
Guminida .......................................................30 Heteroxya ......................................................68
Gummina .......................................................31 Hexactinella ................................................100
Gummineae....................................................30
HEXACTINELLIDA..................................97
Gumminidae ..................................................30
HEXACTINOSIDA.....................................98
Gymnorete .....................................................99
Hexadella ......................................................82
Gymnorossella ............................................102
HEXASTEROPHORA...............................98
H Hezekia ........................................................104
Hiattrochota..................................................55
Haastia ..........................................................81 Higginsia.......................................................69
Habrodictyon ..............................................103 Higginsiinae...................................................68
Haddonella ...................................................85 Hiltonus .........................................................50
HADROMERIDA .......................................29 Hippograntia ................................................92
Halichoclona ................................................72 Hippospongia ...............................................77
Halichondria.................................................70 Hircinella ......................................................79
HALICHONDRIDA....................................63 Hircinia .........................................................79
Halichondriella ......................................70, 72 Hirciniadae.....................................................79
HALICHONDRIIDAE ...............................70 Hircinidae ......................................................79
Haliclona.......................................................72 Hispidopetra .................................................40
Haliclonidae...................................................71 Histioderma...................................................49
Haliclonissa ..................................................73 Histoderma....................................................49
Halicnemia ....................................................68 Histodermella................................................50
Halicometes...................................................35 Histodermion ................................................50
Halina..........................................................104 Histodermopsis .............................................42
Halinastra ...................................................104 Holascella ...................................................101
Halisarca.......................................................83 Holascus ......................................................101
HALISARCIDAE........................................83 Holoplocamia ...............................................43
Halispongia ..................................................81 Holopsamma .................................................44
Hallmannia ...................................................47 Holorodesmia ...................................... 42, 104
Halmopsis......................................................81 Holoxea .........................................................26
Halyphysema...............................................104 Homandra .....................................................90
Hamacantha..................................................61 Homaxinella..................................................65
HAMACANTHIDAE..................................61 Homoderma...................................................91
120 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
Menyllus ........................................................42 N
Merlia ............................................................61
Merriamium ..................................................48 Naauna ..........................................................50
Mesapos.........................................................46 Nailondra ......................................................71
Metania .........................................................76 Nailondria...................................................105
Naniupi..........................................................52
METANIIDAE.............................................76
Nara ...............................................................72
Metrastrosa ....................................................28
Nardoma........................................................89
Meyenia .........................................................75
Nardomandrum ............................................90
Meyerina........................................................97
Nardopsandra...............................................90
MICROCIONIDAE.....................................42
Nardopsetta...................................................90
Microcionina ................................................40
Nardopsis ......................................................89
Microcordyla ................................................34
Nardopsortis .................................................90
Microscleroderma ........................................26
Nardoraltis....................................................90
Microstylifer..................................................65
Nardorandra.................................................90
Microtylostylifer ...........................................61
Nardoretta.....................................................90
Microtylotella ...............................................41
Nardorilla .....................................................90
Microxina ......................................................73
Nardoris ........................................................90
Microxistyla ......................................... 69, 105
Nardorortis ...................................................90
Migas .............................................................70
Nardorus .......................................................89
Milene ............................................................70
Nardosa .........................................................89
Minchinella ...................................................96
Naviculina .....................................................43
MINCHINELLIDAE ..................................95 Negombata ....................................................32
Mlea ...............................................................93 Negombo .......................................................32
Monanchora .................................................51 Neoadocia .....................................................72
Monanthus ....................................................71 Neoaulocystis..............................................101
Monectyon.....................................................46 Neocladia ......................................................59
Monocrepidium ............................................65 Neoesperiopsis..............................................55
Monoplectroninia.........................................96 Neofibularia..................................................61
Monoraphis...................................................98 Neofolitispa...................................................51
MONORHAPHIDIDAE.............................97 Neopelta ........................................................37
Monosyringa.................................................27 NEOPELTIDAE..........................................37
Monotria .....................................................105 Neopetrosia ...................................................75
Murrayona ....................................................88 Neoprosypa ...................................................69
MURRAYONIDA .......................................88 Neosiphonia ..................................................37
MURRAYONIDAE.....................................88 Neothenea....................................................105
Mycale ...........................................................62 Nethea............................................................28
Mycale (Acamasina).....................................62 Niphates.........................................................73
Mycale (Aegogropila) ..................................62 NIPHATIDAE..............................................72
Mycale (Carmia)...........................................62 Nisella..........................................................105
Mycale (Grapelia) ........................................62 Nisibaris ........................................................47
Mycale (Mycale) ...........................................62 Normania ......................................................28
Mycalecarmia ...............................................62 Noronha ........................................................61
MYCALIDAE..............................................61 Nudospongilla ..............................................75
Mycalina .......................................................59
Mycalopsis ..................................................105 O
Myceliospongia ............................................85 Oceanapia .....................................................74
Myle ...............................................................29 Oceanapiidae..................................................74
Myliusia.........................................................99 Ochridaspongia............................................76
Myriastra.......................................................27 Octavella .......................................................24
Myrmekioderma............................................69 Okadaia.........................................................51
Myxichela ......................................................42 Oligoceras.....................................................80
Myxilla...........................................................56 Olynthaltus....................................................89
Myxilla (Burtonanchora).............................56 Olynthandra..................................................89
Myxilla (Ectyomyxilla) .................................56 Olynthandrium..............................................89
Myxilla (Myxilla) ..........................................56 Olynthandrus ................................................89
Myxilla (Stelodoryx).....................................56 Olynthella......................................................89
Myxilla (Styloptilon) ....................................56 Olynthelletta..................................................89
MYXILLIDAE.............................................53 Olynthettus ....................................................89
Myxillina........................................................47 Olynthillus.....................................................89
Myxodoryx.....................................................48
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 123
Olynthium......................................................89 Pararhaphoxya.............................................65
Olynthortus ...................................................89 Parasyringella ..............................................46
Olynthulmus ..................................................89 Paratedania ..................................................58
Olynthus ........................................................89 Paratenaciella ..............................................43
Ommatosa .....................................................64 Paratetilla .....................................................25
Onconema .....................................................98 Paratimea......................................................31
Onychomyxilla ..............................................56 Paresperella..................................................62
Oonema .........................................................98 Paresperia.....................................................56
Oopsacas .....................................................105 Parisociella ...................................................62
Ophiraphidites............................................105 Parmula.........................................................76
Ophistospongia ..........................................105 Patuloscula ...................................................73
Ophlitaspongia.............................................44 Pectispongilla ...............................................75
Ophlitaspongiidae..........................................42 Pellina ...........................................................74
Orina..............................................................72 Pellinula ......................................................105
Orthorachis .................................................105 Penares..........................................................27
Oscarella .......................................................24 Pericharax.....................................................88
Oscarellidae ...................................................24 Periphragella................................................99
Oscaria ..........................................................24 Perissinella....................................................65
Osculina ........................................................29 Petrobiona ....................................................96
Oxeosarcodea .............................................105 PETROBIONIDAE.....................................96
Oxeostilon .....................................................71 Petromica ......................................................71
Oxycarmia .....................................................62 Petromicidae..................................................70
Oxycordyla ....................................................34 Petrosaspongia.............................................80
Oxyheteromeyenia ........................................75 Petrosia .........................................................75
Oxylatrunculia ..............................................32 PETROSIIDAE ...........................................75
Oxymycale .....................................................62 Petrostoma ....................................................96
Oxytedania ....................................................58 Phacanthina............................................66, 69
Phakellia .......................................................66
P
Pharetronema .............................................105
Pachamphilla................................................28 Phelloderma..................................................50
Pachastrella ..................................................28 Pheronema ....................................................97
PACHASTRELLIDAE ..............................27 PHERONEMATIDAE ...............................97
Pachastrissa..................................................27 Phialonema ...................................................98
Pachataxa .....................................................27 PHLOEODICTYIDAE...............................74
Pachaxinella .................................................65 Phloeodictyon ...............................................74
Pachychalina ................................................73 Phlyctaenopora ............................................52
Pachydictyum................................................76 Phorbas .........................................................48
Pachymatisma...............................................27 Phorbasidae....................................................48
Pachypellina .................................................74 Phoriospongia ..............................................57
Pachyrotula...................................................75 PHORIOSPONGIIDAE.............................57
Pandaros .......................................................44 Phycopsis.......................................................66
Papillella.......................................................29 Phyllospongia...............................................78
Papillina........................................................29 Phylosiphonia...............................................72
Papillissa.......................................................29 Phymaraphinidae ...........................................36
Papyrula........................................................28 Phymatellinidae .............................................36
Paracordyla ..................................................26 Physcaphora .................................................32
Paracornulum...............................................42 Pilochrota .....................................................27
Paradisconema .............................................98 Pione..............................................................30
Paradoryx......................................................43 Pitalia ............................................................43
Parafieldingia.............................................105 Pityrete.........................................................105
Paragrantia ..................................................92 Placinastrella................................................24
Paraheteropia...............................................91 Placinolopha ................................................24
Parahigginsia ...............................................69 Placochalina.................................................74
Paralelapia ...................................................97 Placoplegma ...............................................101
Paraleucilla ..................................................93 Placospongia ................................................32
Parameyenia .................................................76 PLACOSPONGIIDAE...............................32
Paramurrayona ............................................88 Plakidium ......................................................38
PARAMURRAYONIDAE.........................88 Plakina ..........................................................24
Paramyxilla...................................................50 Plakinastrella ...............................................24
Pararete.........................................................99 PLAKINIDAE .............................................24
124 ‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia
Plakoosa........................................................24 Protosuberites...............................................34
Plakortis ........................................................24 Psammaplysilla.............................................84
Platychalina............................................55, 73 Psammascus..................................................57
Platylistrum...................................................97 Psammastra...................................................27
Plectispa........................................................44 Psammina....................................................105
Plectrodendron .............................................34 Psammochela ................................................51
Plectronella...................................................47 Psammocinia.................................................79
Plectroninia ..................................................96 Psammoclemma ............................................57
Pleiomeyenia.................................................75 Psammodoryx................................................58
Pleorhabdus................................................105 Psammopemma.............................................57
Pleraplysilla..................................................82 Psammophyllium ........................................105
Pleroma .........................................................37 Psammotoxa..................................................57
PLEROMIDAE ...........................................37 Psetalia..........................................................25
Pleurochorium ..............................................99 Pseudanchinoe..............................................44
Plicatella .......................................................66 Pseudaxinella................................................66
Plicatellopsis.................................................66 Pseudaxinyssa...............................................70
Plocamiancora .............................................56 Pseudobasta..................................................83
Plocamilla .....................................................43 Pseudoceratina .............................................84
Plocamione ...................................................46 Pseudochalina ............................................105
Plocamionida................................................48 Pseudoclathria..............................................52
Plocamiopsis.................................................43 Pseudoesperia...............................................62
Plocamissa ....................................................56 Pseudohalichondria.....................................53
Plumocolumella............................................55 Pseudomyxilla...............................................56
Plumocolumetta............................................56 Pseudosuberites............................................34
Plumohalichondria ......................................48 Pseudotrachya ..............................................66
Pocilloguitarra .............................................60 Psilocalyx ....................................................100
Pocillon .........................................................42 Pteronema .....................................................98
Podospongia.................................................32 Ptilocaulis .....................................................66
Podotuberculum ...........................................48 Ptychodesia ...................................................99
Poecillastra ...................................................28 Pyloderma ...............................................50, 70
Poecilochela..................................................53 Pytheas ..........................................................52
POECILOSCLERIDA ...............................40 Pytheilla ........................................................52
Polejna ..........................................................95 Pyxitis ............................................................27
Poliopogon ...................................................97
Q
Polyfibrospongia..........................................77
Polymastia.....................................................32 Qasimella ......................................................43
POLYMASTIIDAE.....................................32 Quasillina......................................................33
Polytherses ....................................................79 Quepanetsal ................................................105
Poritella.........................................................38 Querciclona...................................................66
Porphyria ......................................................84 Quindesmia ...................................................53
POTAMOLEPIDAE...................................76 Quintoxilla ..................................................106
Potamolepis ..................................................76 Quixilla........................................................106
Potamophloios..............................................76 Quizciona ......................................................43
Poterion.........................................................34
R
Pozziella ........................................................61
Prianos ..........................................................75 Rachella.........................................................25
Prionema .......................................................98 Racodiscula...................................................36
Pronax .....................................................29, 48 Radiella .........................................................33
Pronaxella.....................................................48 Radiospongilla .............................................75
Propachastrella ............................................28 Ramoses.........................................................43
Proraspailia..................................................46 Ramosichela..................................................52
Prostylissa.....................................................70 Raoa...............................................................59
Prosuberites ..................................................34 Raphidotethya...............................................25
Prosycum.......................................................89 Raphioderma ................................................62
Proteleia........................................................33 Raphiodesma ................................................62
Protoclathria ..............................................105 Raphyrus .......................................................29
Protoesperia..................................................62 Raspaciona ...................................................46
Protophlitaspongia ......................................44 Raspaigella ...................................................70
Protoraspailia ..............................................46 Raspailia .......................................................46
Protoschmidtia ...........................................105 Raspailia (Clathriodendron).......................47
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 125
Soleniscilla....................................................89 Sterrastrolepis...............................................76
Soleniscus......................................................86 Sterrastrosa ....................................................27
Soleniscyssa ..................................................89 Stoeba ............................................................28
Solenula.........................................................89 Strepsichordaia ............................................78
Solenulandra ................................................90 Streptoconus .................................................91
Solenuletta ....................................................90 Stretastrosa ....................................................28
Solina...........................................................106 Stromatospongia ..........................................40
Sollasella.......................................................26 Strongylacidon .............................................58
Sollasellidae...................................................26 Strongylamma...............................................58
Somatispongia ............................................106 Strongylodesma ............................................32
Sophax ...........................................................44 Strongylophora.............................................75
Spanioplon....................................................53 Stryphnus.......................................................27
Sphaerotylus .................................................33 Stylaxinella....................................................62
Spheciospongia ............................................30 Stylinos ..........................................................62
Sphenophorina .............................................91 Stylissa...........................................................67
Sphenophorus ...............................................91 Stylocordyla ..................................................34
Sphincterella .................................................28 STYLOCORDYLIDAE..............................33
Sphinctrella...................................................28 Stylohalina ....................................................71
Spinosella......................................................74 Styloptilon .....................................................56
Spinularia .....................................................33 Stylopus .........................................................53
Spirasigmidae ................................................28 Stylorhiza ......................................................34
Spirastrella....................................................33 Stylospira ....................................................106
SPIRASTRELLIDAE.................................33 Stylostichon...................................................48
Spiretta ..........................................................25 Stylotella........................................................67
Spirophora ....................................................36 Stylotellopsis .................................................44
Spirophorella................................................36 Stylotrichophora...........................................62
SPIROPHORIDA .......................................25 Suberanthus ..................................................34
Spirorhabdia.................................................53 Suberea..........................................................84
Spiroxya.......................................................106 Suberella .......................................................34
Spongelia ......................................................81 Suberites........................................................34
Spongia .........................................................78 SUBERITIDAE ...........................................34
SPONGIIDAE..............................................77 Suberotelites..................................................48
Spongilla .......................................................75 Sulcastrella .................................................106
Spongilla (Spongilla)...................................76 Swartschewskia.............................................76
Spongilla (Stratospongilla).........................76 Sycaltaga.......................................................90
SPONGILLIDAE........................................75 Sycandra........................................................90
Spongionella.................................................81 Sycandrarium................................................90
Spongocardium ..........................................106 Sycandrometra..............................................91
Spongosorites ...............................................71 Sycandrophyllum..........................................91
Spongosoritidae.............................................70 Sycantha ........................................................91
Spuma ............................................................70 Sycarium........................................................90
Stannarium..................................................106 Sycetta............................................................90
Stannoma ....................................................106 Sycettaga .......................................................90
Stannophyllum............................................106 SYCETTIDAE .............................................90
Staurocalyptus ............................................102 Sycettopa .......................................................90
STAURORRHAPHIDAE ..........................95 Sycettusa........................................................91
Stegxella ........................................................56 Sycidium ........................................................90
Stelletta..........................................................27 Sycilla ............................................................94
Stellettidae .....................................................26 Sycinula .........................................................93
Stellettinopsis................................................26 Sycinulandra.................................................91
Stelligera .......................................................31 Sycissa ...........................................................90
Stellitethya.....................................................35 Sycocarpus ....................................................90
Stellogeodia ................................................106 Sycocerus.......................................................90
Stelodoryx......................................................56 Sycocubus......................................................90
Stelospongia .................................................80 Sycocystis.......................................................90
Stelospongos .................................................80 Sycocystortis..................................................90
Stelospongus .................................................80 Sycodenandrum ............................................91
Stelotrochota.................................................56 Sycodendrum.................................................90
Stematumenia................................................79 Sycodoranna .................................................90
Stenospongia ................................................79 Sycodorilla ....................................................90
‘Sponguide’ - Version August 2000. © John N.A. Hooper. Qld. Museum, Australia 127