Guide To Sponge Identification

SPONGUIDE.
GUIDE TO SPONGE COLLECTION AND IDENTIFICATION

(Version August 2000)
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 In any case, this guide will be superceded eventually by a more comprehensive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.
LIST OF SPONGE HIGHER TAXA
Family Clionidae Gray, 1867.
Family Hemiasterellidae Lendenfeld, 1889.
Family Latrunculiidae Topsent, 1922.
Class Archaeocyatha (extinct)
Family Placospongiidae Gray, 1867.
Family Polymastiidae Gray, 1867.
Class Demospongiae.
Family Spirastrellidae Ridley & Dendy, 1886.
Subclass Homoscleromorpha.
Family Stylocordylidae Topsent, 1928.
Order Homosclerophorida.
Family Suberitidae Schmidt, 1870.
Family Plakinidae Schulze, 1880.
Family Tethyidae Gray, 1867.
Subclass Tetractinomorpha.
Family Timeidae Topsent, 1928.
Order Spirophorida.
Family Trachycladidae Hallmann, 1917.
Family Tetillidae Sollas, 1886.
Order Lithistida".
Family Scleritodermidae Sollas, 1888.
Suborder Triaenosina.
Order Astrophorida.
Family Corallistidae Sollas, 1888.
Family Ancorinidae Schmidt, 1870.
Family Pleromidae Sollas, 1888.
Family Calthropellidae Lendenfeld, 1906.
Family Theonellidae Lendenfeld, 1903.
Family Coppatiidae Topsent, 1898.
Suborder Rhabdosina.
Family Geodiidae Gray, 1867.
Family Cladopeltidae Sollas, 1888.
Family Pachastrellidae Carter, 1875.
Family Neopeltidae Sollas, 1888.
Family Theneidae Sollas, 1886.
Suborder Anoplina.
Family Thrombidae Sollas, 1888.
Family Azoricidae Sollas, 1888.
Order Hadromerida.
Family Desmanthidae Topsent, 1893.
Family Chondrillidae Gray, 1872 (Schmidt,
Family Vetulinidae Lendenfeld, 1903.
1862).
Subclass Ceractinomorpha.
Phylum Porifera
Subphylum Cellularia
Sponguide - Version August 2000. John N.A. Hooper. Qld. Museum, Australia
Order Verticillitida.
Family Cryptocoeliidae Steinmann, 1982.
Order Agelasida.
Family Agelasidae Verril, 1907.
Family Astroscleridae Lister, 1900.
Order Poecilosclerida.
Suborder Microcionina .
Family Iophonidae Burton, 1929.
Family Microcionidae Carter, 1875.
Family Raspailiidae Hentschel, 1923.
Family Rhabderemiidae Topsent, 1928.
Suborder Myxillina.
Family Anchinoidae Topsent, 1928.
Family Coelosphaeridae Hentschel, 1923.
Family Crambidae Lvi, 1963.
Family Crellidae Hentschel, 1923.
Family Hymedesmiidae Topsent, 1928.
Family Myxillidae Topsent, 1928.
Family Phoriospongiidae Lendenfeld, 1888.
Family Tedaniidae Ridley & Dendy, 1886.
Suborder Mycalina.
Family Cladorhizidae de Laubenfels, 1936.
Family Guitarridae Burton, 1929.
Family Desmacellidae Ridley & Dendy, 1886.
Family Hamacanthidae Gray, 1872.
Family Mycalidae Lundbeck, 1905.
Order Halichondrida.
Family Axinellidae Carter, 1875.
Family Desmoxyidae Hallmann, 1917.
Family Dictyonellidae van Soest,Diaz &
Pomponi, 1990.
Family Halichondriidae Vosmaer, 1887.
Order Haplosclerida.
[Marine haplosclerida families]:
Family Callyspongiidae de Laubenfels, 1936.
Family Chalinidae Gray, 1867.
Family Niphatidae Van Soest, 1980.
Family Phloeodictyidae Carter, 1882.
Family Petrosiidae van Soest 1980.
[Freshwater haplosclerid families].
Family Spongillidae Gray, 1867.
Family Metaniidae Volkmer-Ribeiro, 1986.
Family Potamolepidae Brien, 1967.
Family Lubomirskiidae Brien, 1969.
Order Dictyoceratida.
Family Ircinidae Gray, 1867.
Family Thorectidae Bergquist, 1978
Family Spongiidae Gray, 1867.
Order Dendroceratida.
Family Dysideidae Gray, 1867.
Family Darwinellidae Merejkowsky, 1879.
Family Dictyodendrillidae Bergquist, 1980.
Family Halisarcidae Vosmaer, 1885.
Order Verongida.
Family Aplysinidae Carter, 1875.
Family Druinellidae Lendenfeld, 1889.
Family Ianthellidae Hyatt, 1875.
Class Calcarea.
Subclass Calcinea.
Order Clathrinida.
Family Clathrinidae Minchin, 1900.
Family Soleniscidae Borojevic et al., 1990.
Family Levinellidae Borejevic et al., 1986.
Family Leucaltidae Dendy & Row, 1913.
Family Leucascidae Dendy, 1893.
Family Leucettidae de Laubenfels, 1936.
Order Murrayonida.
Family Murrayonidae Kirkpatrick, 1910.
Family Paramurrayonidae Vacelet, 1967.
Family Lelapiellidae Borojevic, Boury-Esnault
& Vacelet, 1990 .
Subclass Calcaronea.
Order Leucosoleniida.
Family Leucosoleniidae Minchin, 1898.
Family Amphoriscidae Dendy, 1892.
Family Grantiidae Dendy, 1892.
Family Heteropiidae Dendy, 1893.
Family Lepidoleuconidae Vacelet, 1967.
Family Sycettidae Dendy, 1892.
Family Staurorrhaphidae Jenkin, 1908.
Order Lithonida.
Family Lelapiidae Dendy & Row, 1913.
Family Minchinellidae Dendy & Row, 1913.
Family Petrobionidae Borojevic, 1979.
Subphylum Symplasia
Class Hexactinellida.
Subclass Amphidiscophora.
Order Amphidiscosida.
Family Hyalonematidae Gray, 1857.
Family Monorhaphididae Ijima, 1927.
Family Pheronematidae Gray, 1870.
Subclass Hexasterophora.
Order Hexactinosida.
Family Aphrocallistidae Gray, 1867.
Family Aulocalycidae Ijima, 1927.
Family Craticulariidae Rauff, 1893.
Family Euretidae Zittel, 1877.
Family Farreidae Gray, 1872.
Family Tretodictyidae Schulze, 1886.
Order Lychniscosida.
Family Aulocystidae Schulze, 1886.
Family Dactylocalycidae Gray, 1867.
Order Lyssacinosida.
Family Caulophacidae Schulze, 1886.
Family Euplectellidae Gray, 1867.
Family Leucopsacasidae Ijima, 1903.
Family Rossellidae Gray, 1872.
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.
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.
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 ?
qHOW MANY SPONGES ARE THERE ?
l Frankly, we don't really know the magnitude of sponge biodiversity, but we have some theories ...
l In some habitats sponges are the dominant macrobenthic life form; in others they are very scarse.
l Some places in the world have a relatively well-known sponge faunas (e.g. Mediterranean, Caribbean,
British Isles), basically reflecting the relatively long period of time scientists have worked on these
faunas. Each of these regions contains about 500-800 species, with no more than about 100 species
shared between the regions (i.e. so-called cosmopolitan species).
l Some areas have moderately well known faunas (e.g. Madagascar, New Caledonia, New Zealand, Sri
Lanka, Micronesia, Japan) mainly as a consequence of recent scientific exploration during the past 20
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 havent 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).
qWHY ARE WE INTERESTED IN SPONGES ?
l Sessile marine invertebrates are largely an unknown biological resource. Sponges, hard corals, soft
corals, gorgonians and ascidians in particular are megadiverse in the Indo-west Pacific region. We need
to know this biological diversity as a matter of priority, for reasons of potential genetic and chemical
resources as well as matters of ecology and conservation management.
l Many of these marine species are rapidly disappearing because of habitat destruction and overfishing
(e.g. indiscriminate trawling, dynamite fishing etc.).
l There is now a REALISED POTENTIAL that sessile marine invertebrates are economically important,
apart from providing "benthos" or shelter/ habitats for other commercial species such as prawns and
fishes, they have the most active biochemicals potentially useful for mankind in the pharmaceutical
industry.
l "Bioprospecting", seeking out new species for their genetic and chemical resources, is also the most
accurate and comprehensive way to discover as many new and unusual species as possible because new
species and genera often equate to new types of chemicals and chemical activities, some already found
useful against human pathogens and other human ailments.
l Of all life forms marine animals contain the highest number of completely new molecules discovered
over the past 20 years, and many of these are "biologically active" against human pathogens and other
ailments.
l Of all marine species the phyla of sessile marine invertebrates show greatest "biological activity"
agains pathogens and other human ailments.
qEXAMPLES OF SPONGES DISCOVERED TO HAVE MEDICAL PROPERTIES
l Inorganic basis for toxicity (Tedania charcoti from Antarctica with extraordinarily high levels of
cadmium and zinc accumulated from the sea water - extremely effective antibiotic, but also toxic
towards mammalian tissues).
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.
qFROM SEABED TO SICKBED": PROCESS FROM DISCOVERY TO MEDICAL
APPLICATION
l It is a long and complex process from discovery of a new sponge and new molecules to the production
of a safe, effective commercial compound with medical application.
qIMPORTANCE OF ACCURATE, CONTINUALLY MAINTAINED MUSEUM ("VOUCHER
SPECIMEN") COLLECTIONS FOR TAXONOMY AND "BIOPROSPECTING" MEDICAL
RESEARCH
l Why is it important to maintain museum collections of sponges for medical research, and why is
taxonomy (science of identification and evolutionary relationships) important to this process ?
l We MUST know the species of sponge from which the "magic molecule" was discovered, exactly
where it was collected, what season it was collected in, and whether there are morphologically similar
species living nearby that may be confused with the target species. Museum collections, specimen
database, and species "profile" databases are essential for this process.
l A fragment or duplicate "voucher specimen" of a target species MUST be kept (preserved) in a
museum (i.e. separate from the samples analysed chemically or genetically). In sponge taxonomy it is
usually always essential to re-check this "voucher specimen" against new samples to be sure whether it is
the same or different. Because there are so many morphologically similar species that are often
taxonomically unrelated, and certainly chemically very different, using a published species description is
often not adequate by itself to identify a sponge, particularly when the description is pre-1900 (which is
largely the situation for the Indo-west Pacific sponge fauna).
l Collections and collection databases (containing accurate locality data, accurate descriptions of
species' characteristics) are mainly the provinces of museums, providing a unique resource that many
other scientists often do not have (e.g. no time to both build the collections and databases, and carry out
the target research on these collections) .
l The taxonomist's role is to publish these species descriptions and distributions, with the (maybe
impossible) aim of FULLY document regional biological resources in a systematic, scientific way.
l Good taxonomy underpins every other branch of biological and biochemical science ! Comprehensive,
accurate descriptions, with recognisable illustrations and accurate locality data are essential for
determining the relationships between species and regions (biogeography), how many species are shared
between closer or more distant regions (species' dispersal potential, reproductive mechanisms etc.),
habitat preferences (ecology), population variability (genetics) and chemical relationships
(chemotaxonomy, marine natural products chemistry).
2. METHODS OF DEALING WITH SPONGES IN THE LABORATORY AND PREPARATION
FOR THEIR IDENTIFICATION.
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.
q 2.2. Fixation and preservation.
Sponges should be frozen immediately upon collection, which to a certain extent fixes the colour, or live
material may be placed directly in 80-90% ethanol solution. A 5% concentration of buffered
formaldehyde is a less preferable alternative for fixation, and should be used for only brief periods (e.g.
24 hours), after which specimens should be transferred immediately to ethanol. Calcareous sponges
should not be fixed or preserved in formalin.
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.
q 2.3. Histological preparation for light microscopy.
Usually sponge identifications require two forms of histological preparation: one, a spicule preparation
(for those species with a mineral skeleton), to determine the diversity and geometry of spicules in the
skeleton; and a second, a perpendicular section though the sponge 'tissue', to determine the structure of
the skeleton, the water-canal system, and other aspects of histology.
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.).
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 50-100m 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 (>50m), 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 60C, 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.
q 2.4. Histological preparation for scanning electron microscopy.
Much of the preparation outlined below involves hazardous chemicals and noxious fumes, and certain
safety precautions should be taken (laboratory coat, gloves, safety glasses, eye-wash bottle handy, fume
extraction system). Similarly, it essential that instruments used to preparate SEM preparations are
thoroughly cleaned to prevent contaminants from being introduced.
2.4.1. Section preparation and viewing.
1. Cut a 1-1.5mm thick section of sponge ensuring that both the ectosome and choanosome are
represented. Cut another similar section for later use in spicule preparation.
2. Place section in cavity block and cover with several drops of sodium hypochlorite (houshold bleach)
to etch the collagen from the skeleton. Monitor the etching process through a dissecting
microscope in order to prevent the skeleton falling apart. Delicate structures (plumose,
halichondroid, hymedesmoid skeletons) may only require a few seconds treatment with bleach;
robust skeletons (reticulate, fibrous, articulated skeletons) may several minutes; generally 30
seconds is adequate time in bleach.
3. Pipette off bleach at the appropriate time and immediately add 70% ethanol. Leave stand for several
minutes to ensure bleach is completely neutralised.
4. Omit this step if section is very delicate. Without removing section from cavity block, repeat steps 2-3
substituting concentrated hydrogen peroxide in place of sodium hypochlorite (bleach). Finally
rinse in ethanol.
5. Place section on clean microscope slide and let dry.
6. Mount section on SEM stub with double-sided tape, copper dag, or Superglue. An alternative method
of fixing sample to stub, and one that produces a perfectly smooth background, is to cover
stubs with "Aquadhere" wood glue, let dry completely (usually several days), then prior to use
expose dried glue to vigorous steam (e.g. boiling kettle), which softens the set glue, and simply
place the section on top of the stub (it sinks in a short way but is bonded reasonably well to
stub).
7. Sputter-coat the specimen well to ensure that all fibres are well coated. In some cases uncoated
sections can be viewed successfully under low accelerator voltage, but better results are
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.
2.4.2. Spicule preparation and viewing.
1. Take a thinly cut section (including both ectosome and choanosomal regions) and place in a durham
tube (micro-test tube). Using a clothes peg or similar device to hold the tube add a drop of
concentrated nitric acid to the tube (with opening pointed away from the face).
2. Wait for the vigorous reaction to finish and add another drop of acid. Repeat this step several more
times, using drop-by-drop addition so as to control the reaction and production of oxide byproducts.
3. After the acid digestion process appears to be complete, add enough nitric acid to nearly fill the tube.
Directing the tube away from the face gently heat it over low heat (e.g. an alcohol flame),
ensuring that only small bubbles form but not boiling rapidly. Maintain low boiling for 1-2
minutes and let stand until cool.
4. Centrifuge (4000rpm for 30 seconds is adequate).
5. Pipette off nitric acid leaving the spicule mass at the bottom of the tube undisturbed.
6. Refill the tube with fresh nitric acid and resuspend the spicules using clean, fine, glass rod.
7. Repeat steps 3-5.
8. Fill the tube with firstly demineralised water, 70% ethanol, then two series of 100% ethanol
solutions, resuspending spicules, centrifuging and decanting the supernatant between each
change of solution, finally ending with suspended spicules in a solution of absolute ethanol.
9. Adhere a micro-cover glass onto an SEM stub using double-sided tap or copper dag; place stub in stub
holder.
10. Pipette a couple of drops of spicule solution onto the cover glass, ignite and spread out spicule
solution with a glass rod or forceps until all ethanol is vaporised. Spicules bond to glass
relatively firmly, but excess spicules can be blown off glass using compressed air, or spread out
over the glass by adding further ethanol. Monitor the distribution of spicules on the cover glass
under compound or dissecting microscope (magnification depending on spicule size). Add
more drops of spicule solution and repeat this step if too few spicules are present, but do not
overcrowd field of view for photographic purposes.
11. The alternative method of coating the stub with "Aquadhere" glue can also be used, although ethanol
should not be burnt but evaporated, however, single spicules may sink into glue too far if it is
too soft (left in steam too long).
11. Sputter coat stub, and view at 25kV, minimum working distance and smallest apperture for best
resolution.
3. OUTLINE OF CHARACTERS USED FOR DEMOSPONGIAE IDENTIFICATION.
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.
q 3.1. Brief summary of sponge morphology.
A simple analogy of a sponge is a flexible balloon containing a gelatinous ground substance, a roving
cell population, water canals and water pumping stations, and organic and/or inorganic structures
producing a definite body form. The 'simple sponge' is in fact a very complex histological unit, which
even today is not well understood. For the present purposes of sponge identification we provide here a
brief outline of the main structural features of two of the three classes of Porifera (the Hexactinellida, a
deep water group known as glass sponges, is substantially different and not dealt with here). More
thorough, detailed sources of reference are provided at the end of this book.
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
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
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 socalled 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.
13
3.2.11.2. Spicules. These are classified according to five major criteria:

3.2.11.2.1. Chemical composition. These may be silicate or calcitic, indicating division between the
classes Demospongiae and Calcarea.
3.2.11.2.2. Spicule size. Larger spicules, called megascleres, contribute to the skeletal framework
within the sponge, whereas smaller ones, microscleres, are packed between tracts of megascleres,
supporting the soft parts. Spicule sizes are essential criteria in defining species, in some examples
providing the only easy clues to distinguishing related species, whereas absolute spicule dimensions are
less important at higher taxonomic levels.
3.2.11.2.3. Spicule fusion. Most spicules are free within the mesohyl or bound together by the organic
skeleton, whereas some are characteristically fused together, producing an interlocking or articulated
skeleton. In the Calcarea order Murrayonida these spicules consist of rigid monospicular skeletons
composed of modified triaxons; in the Demospongiae order Lithistida these are special spicules called
desmas.
3.2.11.2.4. Spicule distribution. Localisation of spicules to particular regions is a relatively common
phenomenon. These include ectosomal spicules (or cortical spicules, found on the surface of the
sponge), principal spicules (forming the major structural tracts, or found exclusively inside spongin
fibres), auxiliary spicules (found outside the fibres, scattered within the mesohyl and/or just below the
surface), and echinating spicules (or accessory spicules, poking through the fibres, perpendicular to
them). Only few groups of sponges have all four categories of spicules.
3.2.11.2.5. Spicule shape or geometry. There is an extremely diverse range of shapes known for the
phylum, and this is probably the single most important character in the current system of sponge
taxonomy. Even in a single species there may be many sorts of spicules.
To assist in defining spicule shapes a specialised terminology has evolved.
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
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.
4. MAJOR CHARACTERS USED TO DESCRIBE DEMOSPONGIAE, BASED ON THE
'DELTA' (CSIRO COMPUTERISED DESCRIPTIONS) FORMAT.
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
developmental stage and may be frustrating to use
*SHOW Sponge character list. Revised JNAH 28NOV-1993
*CHARACTER LIST
#1.
<EXTERNAL
MORPHOLOGY>
<COMMUNITY> sponge/
1. macrobenthic, fixed to the substrate/
2. macrobenthic, fixed to rolling over substrate/
3. endolithic, burrowing into soft substrate/
4. excavating, bioeroding, boring into substrate/
5. encrusting, photophilic, exposed/
6. encrusting, sciaphilic, sheltered in overhangs or
caves/
7. zoophytic, overgrowing live organic substrates/
#2. <EXTERNAL MORPHOLOGY> <GROWTH
FORM>/
1. thinly encrusting/
2. thickly encrusting/
3. insinuating, boring calcitic substrates/
4. enlarged basal portion below substrate, fistules
protruding through substrate/
5. lobate, spherical-bulbous/
6. lobate, massive/
7. lobate, stoloniferous, spreading over substrate/
8. simple whip-like (flagelliform), unbranched/
9. arborescent, simple branching, cylindrical
digitate
branches, few bifurcatations/
10. arborescent, cylindrical digitate branches,
complex
branching, repeatedly bifurcate/
11. arborescent, flattened digitate branches,
complex
branching, repeatedly bifurcate/
complex
reticulate branching in one plane/
complex
reticulate branching in more than one plane/
14. arborescent, bushy, irregular branches, thickly
branching in more than one plane/
15. arborescent, bushy, flattened branches, thickly
branching in more than one plane/
16. tubulo-digitate, solid construction/
17. hollow, single tubular digit/
18. hollow, bifurcate tubular digits/
19. hollow, several tubular digits attached to

common
base/
20. lamellate, plate-like/
21. fan-shaped (flabelliform), with lamellae in one
plane/
22. fan-shaped (flabelliform), with lamellae in more
than one plane/
23. capitate/
24. club-shaped/
25. cup-shaped/
26. vase-shaped/
27. spherical (golf ball)/
#3. <EXTERNAL MORPHOLOGY> <POINT OF
ATTACHMENT> attached/
1. directly to substrate/
2. directly to substrate, with enlarged basal
holdfast/
3. to substrate with basal portion burrowed into
sediment/
4. to substrate, insinuating into cavities/
#4. <EXTERNAL MORPHOLOGY> <HUE>/
1. /
1. pale/
2. light/
3. bright/
4. dark/
5. olive/
6. drab/
7. mottled/
8. deep/
9. speckled/
10. reddish/
11. greyish/
12. brownish/
#5. <EXTERNAL
COLOURATION>/
1. unknown/
2. white/
3. beige/
4. yellow/
5. blue/
6. turquiose/
7. green/
8. blue-green/
9. orange/
10. pink/
11. red/
12. red-orange/
13. red-brown/
MORPHOLOGY>
<LIVE
14. maroon/
15. brown/
16. grey-brown/
17. grey/
18. black/
#6.
<EXTERNAL
MORPHOLOGY>
<PRESERVED COLOUR> colour in
ethanol/
1. white/
2. beige/
3. pale beige/
4. light yellow/
5. yellow/
6. brown/
7. dark brown/
8. grey-brown/
9. grey/
10. black/
#7. <EXTERNAL MORPHOLOGY> <OSCULE
SHAPE>/
1. not visible/
2. conspicuous, discrete, with a slightly raised
membraneous lip/
3. conspicuous, discrete with a slightly raised
membraneous lip, and subectosomal drainage canals
radiating away from oscules forming stellate
grooves on
surface/
4. large, terminal, raised, on apex of small surface
papillae/
5. large, terminal, raised, on the apex of fistules/
6. small, evenly scattered over the surface,
producing
porous-reticulate appearance/
#8. <EXTERNAL MORPHOLOGY> <OSCULE
DISTRIBUTION>/
1. /
2. on apex of sponge, confined to distinct pore
areas
(sieve-plates)/
areas
(porocalyces)/
area
(capitum)/
5. mainly on lateral sides of branches/
6. mainly on external surface of lamellae/
7. mainly on apex of digits/
8. scattered evenly over surface/
#9. <EXTERNAL MORPHOLOGY> <OSTIA>
inhalant pores/
1. not visible/
2. minute, dispersed evenly over entire surface/
3. minute, dispersed over external surfaces/
5. minute, dispersed over internal surfaces/
6. minute, dispersed over lateral surfaces/
#10.
<EXTERNAL
MORPHOLOGY>
<TEXTURE>/
1. unknown/
2. soft, slimy/
3. soft, mucusy/
4. firm, mucusy/
5. insubstantial, collapses out of water/
15
6. soft, spongy, compressible/

7. firm, rubbery/
8. tough, compressible, difficult to tear/
9. brittle, easily crumbled/
10. firm, barely compressible/
11. firm, incompressible/
12. stony/
#11.
<EXTERNAL
MORPHOLOGY>
<SURFACE>/
1. translucent, membraneous/
2. translucent, membraneous, optically smooth/
3. translucent, membraneous, hispid/
4. opaque/
5. opaque, membraneous, optically smooth/
6. opaque, membraneous, hispid/
7. opaque, slightly collagenous, lightly pigmented/
8. arenaceous, with crust of detritus/
9. fleshy, collagenous, heavily pigmented/
10. dense, spiculose, heavily pigmented/
#12.
<EXTERNAL
MORPHOLOGY>
<SURFACE SCULPTURING>/
1. even, unornamented/
2. even, choanosomal fibres clearly visible below
surface/
3. even, spicules perpendicular to surface/
4. even, spicules tangential to surface forming
cobweblike network/
5. even, with detachable crust of spicules,
paratangential or tangential to surface/
6. uneven, subectosomal drainage canals, grooves
or
ridges clearly visible below surface/
7. uneven, conulose, prominently sculptured, with
tangential spicule skeleton forming cobweb-like
network
running between surface conules/
8. uneven, with regularly dispersed microconules/
9. uneven, clathrous, with bifurcated surface
processes/
10. uneven, shaggy, with protruding fibres scattered
over surface/
11. uneven, digitiform, with large widely spaced
tapering surface processes/
12. uneven, papillose, with close-set, long, tapering
surface processes/
13. uneven, with paratangential reticulated
ectosomal
fibres/
#13.
<SKELETON>
<ECTOSOMAL
SKELETON>/
1. membraneous, without spicule skeleton/
2. membraneous, heavily collagenous/
3. membraneous, arenaceous, with thick crust of
sand
grains and foreign spicule fragments lying on
surface/
4. membraneous, arenaceous, with sand grains and
foreign spicule fragments incorporated into
peripheral
fibres/
5. membraneous, with special ectosomal fibres
tangential to surface/
6. membraneous, with peripheral choanosomal
fibres
lying close to, and tangential to, surface/
7. membraneous, hispid, with erect spicules from

choanosomal skeleton protruding through surface/
8. unispicular, isotropic, single spicules lying
9. unispicular, isodictyal, with single spicules lying
10. uni- or paucispicular, isodictyal tracts of
spicules lying tangential to surface/
11. uni- or paucispicular, radial tracts of spicules
perpendicular to surface/
12. paucispicular, with sparse bundles of spicules
paratangential or perpendicular to surface/
13. pauci- or multispicular, with discrete bundles of
spicules standing perpendicular spicules/
14. multispicular, with a continuous palisade of
spicules perpendicular to surface/
15. multispicular, with a thick crust with spicules
tangential or paratangential to surface/
16. membraneous, but with a felt of microscleres/
#14.
<SKELETON>
<ECTOSOMAL
SPECIALISATION>/
1. /
2. composed of undifferentiated choanosomal
spicules/
3. composed of undifferentiated subectosomal
spicules/
4. composed of special category of ectosomal
spicules
scattered individually on surface/
spicules
forming erect bundles on surface/
spicules
forming continuous palisade on surface/
7. composed of dense crust of subectosomal
spicules,
tangential or paratangential to surface/
8. composed of acanthostyles in plumose brushes
around
protruding subectosomal spicules/
9. raspailiid, with bundles of ectosomal auxiliary
spicules
surrounding
larger
protruding
subectosomal
spicules/
10. raspailiid, with plumose bundles of
choanosomal
spicules surrounding larger protruding spicules/
11. composed of bundles of raphides dispersed
over
ectosome and surrounding larger protruding
spicules/
#15.
<SKELETON>
<SUBECTOSOMAL
SKELETON>/
1. /
2. undifferentiated from choanosomal skeleton/
3. collagenous, lacking any region skeleton/
4. compressed, with fibres forming a more closemeshed
reticulation in periphery than in axial skeleton/
5. cavernous, with peripheral fibres less
compressed
than in axial skeleton/
6. vestigial, with spicules sparsely dispersed
throughout peripheral skeleton/
7. compressed, with more-or-less disorganized
tangential tracts of spicules lying below ectosome/
8. compressed, with more-or-less disorganized

paratangential tracts of spicules lying below
ectosome/
9. scattered tangential tracts of spicules lying below
ectosome/
10. single or paucispicular isodictyal tracts/
11. cavernous, with wide-meshed reticulate tracts
of
spicules lying below ectosome/
12. plumo-reticulate, with diverging and
anastomosing
tracts of spicules supporting ectosome/
13. plumose, with diverging brushes of spicules
supporting ectosome/
14. radial, with perpendicular bundles of spicules
supporting ectosome/
15. radial, with single spicules perpendicular to axis
protruding through ectosome/
#16.
<SKELETON>
<SUBECTOSOMAL
SPECIALISATION>/
1. /
2. composed of undifferentiated choanosomal
spicules/
3. composed of undifferentiated ectosomal
spicules/
4. composed of special category of auxiliary
spicules,
geometrically different from choanosomal spicules/
spicules,
differing from ectosomal spicules only in
dimensions/
spicules,
differing from choanosomal spicules only in
dimensions/
7. composed of echinating spicules concentrated in
peripheral skeleton/
8. supplemented by sand grains and other foreign
particles dispersed throughout peripheral skeleton/
#17.
<SKELETON>
<CHOANOSOMAL
ARCHITECTURE>/
1. collagenous/
2. hymedesmoid/
3. microcionid/
4. plumose/
5. plumo-reticulate/
6. irregularly reticulate/
7. regularly reticulate/
8. renieroid-subisodictyal reticulate/
9. isodictyal reticulate/
10. isotropic reticulate/
11. disorganised halichondroid reticulate/
#18. <SKELETON> <CHOANOSOMAL AXIAL
AND EXTRA-AXIAL
DIFFERENTIATION> with/
1. /
2. undifferentiated axial or extra-axial regions/
3. wide-meshed relatively homogeneous fibres/
4. wide-meshed skeletal tracts/
4. compressed basal fibres, and radial extra-axial
skeleton/
5. compressed basal fibres and plumose extra-axial
skeleton/
6. compressed basal skeleton composed of short
bulbous
fibre nodes, cored by plumose tufts of radiating

extraaxial spicules/
7. differentiated, compressed, reticulate axis, and
plumo-reticulate extra-axis/
plumose extra-axis/
radial extra-axis/
10. differentiated plumose axis, and radial extraaxis/
11. mostly non-anastomosing fibres and spicule
tracts/
12. differentiated primary (plumose-dendritic) and
secondary (renieroid-subrenieroid) skeletons/
13. differentiated primary multispicular skeleton and
secondary renieroid skeleton/
14. radially dispersed single spicules or
paucispicular
tracts of spicules throughout/
15. a criss-cross of spicules, vaguely recognisable
in
structure only at the periphery/
16. -out apparent skeletal organization/
#19.
<SKELETON>
<CHOANOSOMAL
SKELETAL TRACTS>/
1. /
2. basal fibres aspiculose, although bases of
spicules
embedded in spongin fibres, standing perpendicular
to
substrate/
3. homogeneous unispicular skeletal tracts cored by
choanosomal spicules/
4. homogeneous uni- or paucispicular skeletal tracts
cored by choanosomal spicules/
5. homogeneous multispicular skeletal tracts cored
by
choanosomal spicules, occupying only proportion
of fibre
diameter/
by
choanosomal spicules, occupying entire fibre
diameter/
by
auxiliary spicules identical to those in peripheral
skeleton/
by
choanosomal spicules with echinating spicules
secondarily incorporated into fibres/
9. larger primary (ascending) and smaller secondary
(transverse) skeletal tracts cored by principal
spicules/
10. larger primary (ascending) skeletal tracts cored
by
principal spicules, smaller secondary (connecting)
tracts cored by auxiliary spicules/
by
principal spicules, smaller secondary (connecting)
tracts aspiculose/
by
auxiliary spicules, smaller secondary (connecting)
tracts aspiculose/
17
13. only primary (ascending) skeletal tracts present,

cored by principal spicules, without connecting
tracts/
14. skeletal tracts in axial skeleton cored by
auxiliary spicules, whereas tracts in peripheral region
wholly arenaceous/
15. primary spongin fibres interconnected by
smaller
secondary fibres/
16. without spongin fibres or spicule tracts,
although
collagen fibrils present/
#20.
<SKELETON>
<CHOANOSOMAL
SPICULES> choanosomal
spicules/
1. /
2. absent/
3. completely enclosed in spongin fibres/
4. core spongin fibres as well as echinate fibre
endings, protruding through fibres in "spicate"
arrangement/
5. echinates spongin fibres as well as form plumose
ascending brushes in peripheral skeleton/
6. echinate fibre nodes/
7. strewn in loosely aggregated tracts within
mesohyl/
8. strewn in halichondroid tracts within mesohyl/
9. form secondary renieroid skeleton, without a
fibre
component, bound at nodes by collagen/
10. form a rigid, interlocking skeleton/
#21.
<SKELETON>
<CHOANOSOMAL
ECHINATING SPICULES>
echinating spicules/
1. /
2. absent (presumed lost)/
3. absent, although second category of larger,
acanthose choanosomal spicule present/
4. absent, although cladotylotes echinate skeletal
tracts/
5. clumped on basal spongin/
6. concentrated in tufts at fibre nodes/
7. concentrated on exterior edges of skeletal tracts/
8. concentrated on primary skeletal tracts/
9. confined to peripheral skeleton, forming plumose
brushes on ectosome, surrounding protruding
subectosomal
spicules/
10. clumped at junction of axial and extra-axial
skeletons, around bases of subectosomal spicules/
11. secondarily incorporated into fibres/
12. form a dense, rigid, interlocking secondary
skeleton/
13. forming plumose or subrenieroid tracts only/
14. sparse, evenly distributed over skeletal tracts/
15. heavy, evenly distributed over skeletal tracts/
16. supplemented by cladotylotes also echinating
skeletal tracts/
#22.
<SKELETON>
<CHOANOSOMAL
SPONGIN FIBRES> spongin
fibres/
1. /
2. absent/
3. absent, spicule tracts bound together at their
nodes by collagen/
4. poorly developed/
5. poorly developed, lightly invested with spongin,

homogeneous in cross-section/
6. poorly developed, lightly invested with spongin,
spicule tracts mostly bound together by granular
collagen/
7. poorly developed, replaced mostly by algal
filaments/
8. well developed, with optically diffuse central
pith/
9. well developed, supplemented by spongin
spicules/
10. well developed, supplemented by collagen
filaments/
11. well developed, supplemented by a mesentarylike
tertiary "fibre" network/
12. well developed, fibres strongly and
concentrically
lamellated and stratified, with both bark and pith
elements/
13. reduced to a layer of spongin lying on
substrate,
without coring spicules/
#23.
<SKELETON>
<INCORPORATED
DETRITUS>
1. /
2. with sand grains incorporated into fibres/
3. with sand grains and foreign spicule fragments
incorporated into fibres/
4. fully cored by detritus, obscuring most spicule
tracts/
#24.
<SKELETON>
<CHOANOSOMAL
MESOHYL> collagen/
1. seen mainly around spicule tracts and spicule
nodes/
2. abundant, lightly pigmented, dispersed evenly
throughout mesohyl/
3. abundant, darkly pigmented, dispersed evenly
throughout mesohyl/
4. abundant, granular, with pigment bodies clumped
within mesohyl, darkly pigmented/
5. abundant, with abundant collagenous filaments/
#25.
<SKELETON>
<CHOANOCYTE
CHAMBERS> choanocyte
chambers/
1. /
2. small, oval <40-200 micrometres diameter>/
3. small, oval-eliptical <20-200 micrometres
diameter>/
4. large, oval-elongate <200-300 micrometres
diameter>/
5. minute, ovoid <5-40 micrometres diameter>/
6. paired <50-350 micrometres diameter>/
#26.
<SPICULES>
<MEGASCLERES,
PRINCIPALS, NO.CATEGORIES>/
1. /
2. principal spicules absent/
3. single category of principal spicule/
4. two categories of principal spicules/
5. three categories of principal spicules/
#27.
<SPICULES>
<MEGASCLERES,
MONACTINAL, PRINCIPALS>
choanosomal principal spicules are/
1. /
2. styles/
3. subtylostyles/
4. tylostyles/
5. acanthostyles/
6. rhabdostyles/
7. quasi-diactinal spicules (asymmetrical)/
8. strongyles/
9. acanthostrongyles/
10. sinuous strongyles/
11. tylotes <amphitylotes>/
12. tornotes/
13. oxeas/
14. anisoxeas/
15. strongyloxeas/
16. tuberculate vermiform strongyles/
17. cladotylotes/
#28.
<SPICULES>
<MEGASCLERES,
MONACTINAL, PRINCIPALS>/
1. /
2. entirely smooth/
3. with lightly acanthose base and smooth shaft/
4. with heavily acanthose base and smooth shaft/
5. with acanthose base, partly acanthose neck, but
remainder of shaft smooth/
6. acanthose at both base and apex, with smooth
shaft/
7. acanthose only at extremities/
8. entirely acanthose/
9. with spines in verticillate rows/
#29.
<SPICULES>
<MEGASCLERES,
MONACTINAL, PRINCIPALS>
with/
1. /
2. evenly rounded bases, fusiform points/
3. evenly rounded bases, hastate points/
4. subtylote bases, fusiform points/
5. subtylote bases, hastate points/
6. tylote bases, fusiform points/
7. tylote bases, hastate points/
8. tapering fusiform bases, fusiform points/
9. tapering fusiform bases, hastate points/
10. tapering pointed bases, fusiform points/
11. tapering pointed bases, hastate points/
12. rounded and mucronate bases, fusiform points/
13. rounded and mucronate bases, hastate points/
14. tuberculate bases, fusiform points/
15. tuberculate bases, hastate points/
16. slightly rhabdose bases, fusiform points/
17. slightly rhabdose bases, hastate points/
18. rhabdose bases, fusiform points/
19. rhabdose bases, hastate points/
20. rhabdose bases with a spiral twist, fusiform
points/
21. rhabdose bases with a spiral twist, hastate
points/
22. fusiform points/
23. hastate points/
24. tapering rounded points/
25. evenly rounded points/
26. hastate, telescoped points/
27. sinuous, flexuous points/
#30.
<SPICULES>
<MEGASCLERES,
TETRACTINAL, PRICIPALS>/
1. /
2. short-shaft mesotriaenes/
3. plagiotriaenes/
4. anatriaenes/
5. protriaenes/
6. promonaenes/
7. orthotriaenes/
8. dichotriaenes/
9. trichotriaenes/
10. trichodal or heterocladal protriaenes/
11. discotriaenes/
12. phyllotriaenes/
13. oxytylotes (diacts)/
14. absent (presumed secondary loss)/
15. discrete spicules absent, skeleton entirely fused
calcitic or aragonitic basal skeleton (hypercalcified)/
#31.
<SPICULES>
<MEGASCLERES,
TETRACTINAL, DESMAS FORMING
SECONDARY CHOANOSOMAL SKELETON>/
1. /
2. ophirhabds (monocrepidial)/
3. dendroclones (monocrepidial)/
4. tricranoclones (tricrepidial)/
5. rhizoclones (monocrepidial)/
6. fused heloclones (monocrepidial)/
7. megaclones (monocrepidial)/
8. tetraclones (tetracrepidial)/
9. rhabocrepids (monocrepidial)/
10. sphaeroclones (tetracrepidial)/
#32.
<SPICULES>
<MEGASCLERES,
AUXILIARY, NO.CATEGORIES>/
1. /
2. auxiliary spicules absent/
3. single category of auxiliary spicule/
4. two categories of auxiliary spicules in ectosomal
and subectosomal regions, respectively/
5. three categories of auxiliary spicules, with
different
distributions
in
ectosomal
and
subectosomal
regions/
#33.
<SPICULES>
<MEGASCLERES,
AUXILIARY
> auxiliary
spicules are/
1. /
2. styles/
3. subtylostyles/
4. tylostyles/
5. acanthostyles/
6. rhabdostyles/
7. quasi-diactinal spicules (asymmetrical)/
8. strongyles/
10. sinuous strongyles/
11. tylotes <amphitylotes>/
12. tornotes/
13. oxeas/
14. anisoxeas/
15. strongyloxeas/
16. tuberculate vermiform strongyles/
17. cladotylotes/
18. polyactinal (asterose)/
#34.
<SPICULES>
<MEGASCLERES,
SUBECTOSOMAL AUXILIARY>
with/
1. /
2. entirely smooth base and shaft/
3. lightly microspined bases/
19
4. heavily microspined bases/

5. with microspined bases and points, smooth
shaft/
6. entirely spined/
#35.
<SPICULES>
<MEGASCLERES,
SUBECTOSOMAL AUXILIARY>
and/
1. /
2. evenly rounded bases, fusiform points/
3. evenly rounded bases, hastate points/
4. subtylote bases, fusiform points/
5. subtylote bases, hastate points/
6. tylote bases, fusiform points/
7. tylote bases, hastate points/
8. tapering rounded base, fusiform points/
9. tapering rounded base, hastate points/
10. fusiform points/
11. hastate points/
12. tapering rounded points/
13. evenly rounded points/
14. hastate, telescoped points/
15. sinuous, flexuous points/
#36.
<SPICULES>
<MEGASCLERES,
CALTHROPS AUXILIARY
SPICULES>/
1. /
2. absent (presumed secondary loss)/
3. undifferentiated calthrops/
4. tetrapod calthrops/
5. centrangulate diacts/
6. monolophs (lophotetractines)/
7. candelabra/
8. amphimesodichotriaenes/
9. short-shafted dichotriaenes/
#37.
<SPICULES>
<MEGASCLERES,
ECHINATING, NO.CATEGORIES>/
1. /
2. echinating spicules absent/
3. identical in size and geometry to principal
spicules/
4. single category of echinating spicule/
5. two categories of echinating spicules/
6. three categories of echinating spicules/
#38.
<SPICULES>
<MEGASCLERES,
ECHINATING, CURVATURE>
echinating spicules/
1. /
2. straight/
3. slightly curved at centre/
4. greatly curved at centre/
5. slightly curved in distal third of shaft/
6. slightly curved in basal third of shaft/
#39.
<SPICULES>
<MEGASCLERES,
ECHINATING, SHAPE>/
1. /
2. club-shaped/
3. cylindrical, rod-like/
4. subcylindrical, sausage-shaped/
5. triactinal <three rays>/
6. tetractinal <four rays>/
7. polyactinal <many rays>/
8. di-, tri- and tetractinal/
#40.
<SPICULES>
<MEGASCLERES,
ECHINATING, GEOMETRY>/
1. /
2. acanthostyles, rounded bases, fusiform points/
3. acanthostyles, rounded bases, hastate points/
4. acanthostyles, subtylote bases, fusiform points/
5. acanthostyles, subtylote bases, hastate points/
6. acanthostyles, prominently tylote bases, fusiform
points/
7. acanthostyles, prominently tylote bases, hastate
points/
8. smooth styles, rounded bases, fusiform points/
9. smooth styles, rounded bases, hastate points/
10. smooth styles, subtylote bases, fusiform points/
11. smooth styles, subtylote bases, hastate points/
12. smooth strongyles, evenly rounded bases/
14. smooth rhabdostyles/
15. acanthorhabdostyles/
16. sagittal triacts (acanthoplagiotriaenes), including
tetracts and quasi-diacts/
17. cladotylotes with smooth shaft/
18. cladotylotes with spined shaft/
19. choanosomal rhabdostyles, identical in
geometry to
principal spicules/
20. diactinal acanthorhabds/
21. robust, fusiform oxeas/
22. acanthoxeas/
#41.
<SPICULES>
<MEGASCLERES,
ECHINATING, SPINATION>/
1. /
2. entirely smooth/
3. entirely acanthose, with small granular spines
dispersed evenly/
4. entirely acanthose, well large spines dispersed
evenly/
5. with small spines on base only/
6. with heavy spines on base only/
7. with spines on base and "neck" only/
8. with smooth base and spines on point and shaft
only/
9. with spines on base and shaft but smooth point/
10. with spines evenly dispersed over spicule
except
for aspinose "neck"/
11. with smooth shaft and spines only on
extremities/
12. with large recurved clads on apex, swollen
base,
smooth shaft/
13. with large strongly recurved clads on both
ends,
smooth shaft/
14. with large strongly recurved clads on apex,
small
clads on base, lightly spined shaft/
15. with small clads on both base and apex, and
lightly
spined shaft/
16. with small clads on both base and apex, and
smooth
shaft/
17. with large erect spines covering, and nearly
perpendicular to, shaft/
18. with small scattered spines over shaft, and
clads
(large, strongly recurved hooks) present on both

base
and apex of spicule/
19. with spines on major axis (longest arm) only/
20. with spines on all axes/
21. with spines in regular, verticillate rows, covering
most of spicule/
#42.
<SPICULES>
<MICROSCLERES,
NO.CATEGORIES>/
1. single category microsclere/
2. two categories microscleres/
3. three categories microscleres/
4. four categories microscleres/
5. five categories microscleres/
6. six categories microscleres/
7. seven categories microscleres/
8. eight categoris microscleres/
9. nine categories microscleres/
10. absent/
11. /
#43.
<SPICULES>
<MICROSCLERES,
MENISCOID, GEOMETRY>
consisting of/
1. /
2. unmodified palmate isochelae/
3. palmate isochelae, including contort forms/
4. cleistochelae/
5. arcuate isochelae/
6. anchorate isochelae/
7. bidentate sigmoid isochelae/
8. unguiferous isochelae/
9. Isodictya-like isochelae/
10. crocae (J-shaped) chelae/
11. birotulates/
12. bipocillae/
13. palmate anisochelae/
14. placochelae/
15. sphaerancora/
16. canonochelae/
17. clavidiscs (compound diancistras <Merlia>/
18. diancistras, sigmoid, with central and apical
notches/
19. cyrtancistras, with serrated inner margin
<sigmoid>/
20. sigmancistras, with vestigial central notch/
21. c-sigmas/
22. s-sigmas/
23. spirosigmas, small, contort, rugose/
24. thraustosigmas, large, irregular, crooked,
rugose/
25. centrangulate sigmas/
26. tetrapocillae/
27. spined isancorae/
28. spined isochelae/
#44. <SPICULES> <MICROSCLERES, TOXAS,
GEOMETRY>/
1. /
2. oxhorn toxas, completely, smooth, reflexed
arms,
wide central curvature/
3. oxhorn toxas, reflexed arms with spines on
points,
wide central curvature/
4. accolada toxas, slight central curvature, long
straight arms, straight points, /
5. thin, deeply curved toxas, straight points/
6. v-shaped toxas, sharp angular central curvature,

straight arms, straight points/
7. v-shaped toxas, sharp angular central curvature,
straight arms, slightly reflexed points/
8. asymmetrical, sinuous toxas/
9. oxeote toxas, no or vestigial curvature, straight
points/
10. acanthose cladotoxas/
11. smooth forceps/
12. acanthose forceps/
#45. <SPICULES> <MICROSCLERES, OTHER
MONAXONIC,
GEOMETRY>/
1. /
2. discorhabds <Didiscus>/
3. anthosigmas/
4. sanidastoid discorhabds/
5. anisodiscrohabds/
6. spirasters/
7. spinispiras/
8. spirulas/
9. toxaspires/
10. sigmaspires/
11. selenasters/
12. smooth microxeas/
13. acanthose microxeas/
14. microstrongyles/
15. oval microstrongyles/
16. centrotylote microxeas/
17. smooth microstyles/
18. rugose microstyles/
19. microtylostyles/
20. "microrhabds", degenerate microxeas/
21. smooth commata/
22. rugose commata/
23. spheres/
24. raphides/
25. trichites, bundles of raphides/
26. onychaetes/
27. spined centrotylote rods/
28. spear-shaped microstyles/
29. thraustoxeas/
21
30. sanidasters/
31. "ecailles", monocrepidial disc/
#46.
<SPICULES>
<MICROSCLERES,
ASTROSE >/
1. /
2. plesiaster streptasters/
3. amphiaster streptasters/
4. metaster streptasters/
5. spiraster streptasters/
6. oxyaster euasters/
7. oxyspheraster euasters/
8. pycnaster euasters/
9. strongylaster euasters/
10. tylaster euasters/
11. anthaster euasters/
12. anthospheraster euasters/
13. sterrospheraster euaster/
14. sterraster euasters/
15. aspidaster euasters/
16. ataxasters euasters (acanthosphaera, or
degenerate
euasters)/
#47. <SPICULES> <DIMENSIONS PRINCIPAL
MEGASCLERES>/
#48. <SPICULES> <DIMENSIONS AUXILIARY
MEGASCLERES>/
#49. <SPICULES> <DIMENSIONS CALTHROPS
AUXILIARY
MEGASCLERES>/
#50.
<SPICULES>
<MEGASCLERES,
ECHINATING, DIMENSIONS>/
#51.
<SPICULES>
MICROSCLERES>/
<DIMENSIONS
#52. <REMARKS>/
5. KEY TO THE EXTANT ORDERS OF PORIFERA.

1. Mineral skeleton composed of calcitic spicules ..................................................... [Class Calcarea] (2)
Mineral skeleton composed of six-rayed silica spicules, occurring both individually and fused together
..... .....................................................................................................................[Class Hexactinellida] .... (5)
Mineral skeleton composed of silica spicules and/or spongin fibres ........... [Class Demospongiae] .....
(7)
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].
4. Body plan asconoid
Leucosoleniidae].
....................................................
[Order
Leucosoleniida,
family
Body plan syconoid or leuconoid ............................................... [Order Leucosoleniida, 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)].
9. With siliceous megascleres and/or microscleres, with spongin fibres ..................................................
(10)
Without free spicules, with spongin fibres .............................................................................................
(11)
10. Microscleres include chelae and other diverse forms, megascleres often localized to distinct regions
(e.g. inside fibres), sand/detritus may replace megascleres completely .................. [Order
Poecilosclerida].
Microscleres typically absent, with main skeleton composed of criss-cross (family Halichondriidae) of
monaxonic megascleres (styles, oxeas, strongyles), usually with more organization at surface, or
sometimes condensed into an axial skeleton and a plumose or plumo-reticulate extra-axial skeleton
(family Axinellidae), or plumose-dendritic mineral skeleton (family Dictyonellidae), fibre system poorly
developed or absent ................................................................................................................... [Order
Halichondrida].
Microscleres may be absent or include centrangulate sigmas, toxas or microxeas, megascleres diactinal
usually producing well-formed structure (e.g. isodictyal-reticulate) ........................ [Order
Haplosclerida].
Main skeleton composed of well developed spongin fibres cored and/or echinated by short styles or
oxeas with verticillate spines [Order
Agelasida]
23
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].
7. THE SPONGE CLASSIFICATION (EXTANT TAXA ONLY)
q THE PHYLUM PORIFERA
Porifera, the pore bearers, contains four classes (one completely extinct), 5 subclasses, 28 orders, 232
families and 977 valid genera (although many more nominal genera which have been sunk by various
authors. There are probably at least 15,000 extant species worldwide and many more extinct ones.
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
(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 (100m 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.
q FAMILY PLAKINIDAE SCHULZE, 1880.
SYNONYM: Corticiidae Vosmaer; Oscarellidae Lendenfeld.
DEFINITION: Encrusting or massive growth forms; simple body structure with aquiferous system
varying from simple asconoid construction to more complex folding and elaborate canal systems;
mineral skeleton composed of relatively small calthrops and/or derivatives (diods or triods), often with
branched ends (lophotetractines), generally arranged uniformly within sponge; spicules usually surround
aquiferous system in regular "alveolar" arrangement; siliceous spicules and spongin fibres absent in one
genus (Oscarella), having only collagenous fibrillar spongin in mesohyl; choanocyte chambers with
300-500 choanocytes, usually eurypylous, occasionally aphodal; larvae unique amphiblastula type.
SCOPE: Fourteen nominal genera are known in this family, of which only seven are probably valid.
REVISIONS: Sole-Cava, Boury-Esnault, Vacelet & Thorpe (1992); Maldonado (1992); Diaz & van
Soest (1994).
GENERA:
Astroplakina Dendy & Burton, 1926 (type species: Astroplakina stelligera Dendy & Burton,
1926) (incertae sedis) - thinly encrusting; spiculation diods, triods, calthrops and multirayed forms,
larger usually smooth whereas smaller oxyasters and strongylasters have roughened rays (possession of
euasters probably indicate Astrophoridan affinities (Jaspis or Hemiasterella)) (Dendy & Burton, 1926;
Diaz & van Soest, 1994).
Corticella Sollas (type species: ) - with only calthrops and spiny eusasters.
Corticium Schmidt, 1862 (type species: Corticium candelabrum Schmidt, 1862) (syn. Achinoe
Gray, 1867) - thinly encrusting, contractile surface; spiculation exclusively tetractines of single size and
candelabras, although spicules occasionally absent completely; aphodal choanocyte chambers (Diaz &
van Soest, 1994).
Placinolopha Topsent, 1897 (type species: Placinolopha bedoti Topsent, 1897) (syn.
Acanthoplakina Burton, 1959; Diactinolopha Sar, 1960) - thinly or massively encrusting; spiculation
in two size classes, consisting of smooth diods, triods, calthrops with irregular ramifications and spines
at ends of rays, and large lophotetractines (lophodiacts, lophotriacts and lophocalthrops) (Diaz & van
Soest, 1994).
Plakina Schulze, 1880 (type species: Plakina monolopha Schulze, 1889) (syn. Plakoosa de
Laubenfels, 1936b) - thinly or massively encrusting; development of ectosome variable and
subectosomal cavities may be present; proportion of mesohyl to chambers varies between 0.7 and 1.8:1;
spiculation of of diods, triods and calthrops, and with lophocalthrops having one, two, three or four
lophate rays; eurypylous or aphodal choanocyte chambers usually with a radial arrangement of chambers
around a central excurrent canal; lophocalthrops highly concentrated at sponge surface (Diaz & van
Soest, 1994; Muricy, Boury-Esnault, Bezac & Vacelet, 1997).
PlakinastrellaSchulze, 1880 (type species: Plakinastrella copiosa Schulze, 1880) (syn.
Dercitopsis Dendy, 1905; ? Placinastrella Topsent) - thinly to massively encrusting, sometimes lobate;
skeleton of diods, triods and calthrops in three distinct size categories, the smaller diactines accumulated
on the surface (Diaz & van Soest, 1994).
Plakortis Schulze, 1880 (type species: Plakortis simplexSchulze, 1880) (syn. Roosa de
Laubenfels, 1934) - thinly to massively encrusting; skeleton formed mainly by small diods with triods in
25
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.
q FAMILY TETILLIDAE SOLLAS, 1886.
SYNONYM: Samidae Sollas.
DEFINITION: Sponges with a perfect radial skeleton and consequent near spherical form, often referred
to as `golf ball sponges'; megascleres triaenes and oxeas arranged in radiate pattern; protriaenes
apomorphic for the family, often protruding from the suface; microscleres contorted sigmaspires with
minute spines; sometimes other modified triaene spicules also present (amphiclads, calthrops-like);
inhalant pores grouped in special pore areas in some genera (poriferous pits or porocalices), unique to
the family; reproductive patterns range from extrusion of fertilized eggs (which are fixed to the substrate
and develop directly), to oviparous (with incubation of complete young sponges which are then expelled
by localised breakdown of the pinacoderm); no free larvae yet described.
SCOPE: Seventeen nominal genera are presently included in the family, although not all these are valid,
with a major revision of the order required. Nine genera are currently recognised here.
REVIEWS: Lendenfeld (1903); Ruetzler (1987), Ruetzler & Smith (1992).
GENERA:
Acanthotetilla Burton, 1950 (type species: ) (syn. Acanthocinachyra Lvi, 1964) - without
cortex, with porocalices, with acanthoxea accessory megasclere spicules.
Amphitethya Lendenfeld, 1907 (type species: Amphitethya microsigma Lendenfeld, 1907) without porocalices, with cortex and cortical layer of amphiclads (amphitriaenes and plagiotriaenes).
Cinachyra Sollas, 1886 (type species: Cinachyra barbata Sollas, 1886) (syn. Fangophilina
Schmidt, 1880; Spiretta Lendenfeld, 1888; Tethyopsilla Lendenfeld, 1888; Kaira de Laubenfels, 1936)
- with flask-shaped porocalices, with specialised cortex reinforced by special oxeas, without unusual
spicules.
Cinachyrella Wilson, 1925 (type species: Tetilla hirsuta Dendy, 1889) - with porocalices,
without cortex, without unusual accessory spicules.
Craniella Schmidt, 1870 (type species: ) (syn. Craniellopsis Topsent, 1913) - cortex
reinforced by erect or criss-crossed oxeas; without porocalices or unusual accessory spicules.
Paratetilla Dendy, 1905 (type species: Tethya merguiensis Carter, 1883) - without cortex,
with porocalices, with special dermal layer of calthrop-like triaenes.
Raphidotethya Burton, 1934 (type species: Raphidotethya enigmatica Burton, 1934) - oxeas
only megascleres, triaenes secondarily lost (possible synonym of Tetilla).
Samus Gray, 1867 (type species: Samus anonyma Gray, 1867) (syn. Rachella Sollas, 1886) excavating growth form; with sigmaspires, with small amphitriaenes, cladocalthrops, without
porocalices, without oxeas.
Tetilla Schmidt, 1868 (type species: Tetilla euplocamos Schmidt, 1868) (syn. Chrotella
Sollas, 1886; Psetalia Gray) - without porocalices, without cortex, without unusual accessory
megascleres.
q FAMILY SCLERITODERMIDAE SOLLAS, 1888.
DEFINITION: Bowl, cup-shaped or plate-like growth forms; fused "lithistid" skeleton composed of
monocrepidial desmas (rhizoclones) and free spicules with slender oxeas distributed in tracts within
choanosome, sometimes protruding through surface; triaenes absent, presumed secondarily lost;
specialised ectosomal skeleton composed of contorted sigmaspires and special ectosomal desmas, also
dispersed throughout sponge, sometimes together with minutely spined microstrongyles.
SCOPE: Five nominal and three valid genera are included here.
REVIEWS: nil.
GENERA:
Aciculites Schmidt, 1879 - rhizoclones, oxytyles, ectosomal tangential acanthose microaniostrongyles, loss of sigmaspires.
Amphibleptula Schmidt, 1879 (syn. Microscleroderma Kirkpatrick, 1902, Taprobane Dendy,
1905) - special ectosomal desmas present or absent, sigmaspires, oxeas.
Scleritoderma Schmidt, 1879 - rhizoclones, acanthose microstrongyles, sigmaspires, oxeas.
q ORDER ASTROPHORIDA (syn. Choristida)
DEFINITION: Typically with asterose microscleres (but sometimes lost), microxeas and microrhabds;
with tetractinal megascleres, usually triaenes, calthrops, or short-shafted triaenes, together with oxeas;
with radial skeletal architecture obvious at least at surface; reproduction oviparous although gametes so
far described for very few species; larval stages not yet known.
Seven families are included in this order.
q FAMILY COPPATIIDAE TOPSENT, 1898.
SYNONYMS: [Epipolasidae] Sollas [nomen nudum]; [Streptasteridae] Topsent [nomen nudum];
Sollasellidae Lendenfeld; Jaspinae de Laubenfels; Jaspidae of authors.
DEFINITION: Encrusting to massive growth forms; megascleres only oxeas forming radial skeleton;
triaenes absent; microscleres euasters (never sterrasters), sometimes microxeas and sanidasters; genera
related to stellettids but lack triaenes.
SCOPE: Seven nominal and four valid genera are included here.
REVIEWS: Hajdu & van Soest (1992)
GENERA:
Jaspis Gray, 1867 (type species: Vioa johnstoni Schmidt, 1862) (syn. Coppatias Sollas, 1888;
Dorypleres Sollas, 1888; Astropeplus Sollas, 1888) - with oxeas and euasters.
Paracordyla Hallmann, 1912 (type species: Paracordyla lignea Hallmann, 1912).
Sollasella Lendenfeld, 1887 (type species: Sollasella digitata Lendenfeld, 1888).
Stellettinopsis Carter, 1879 (type species: ) - with oxeas, asters and microxeas.
q FAMILY ANCORINIDAE SCHMIDT, 1870.
SYNONYM: Stellettidae Carter; Euastrosa Sollas.
DEFINITION: Growth forms either encrusting to massive, or more specialised with spherical body and
long inhalant and exhalant tubes at opposite ends (the latter with stellate, spicular, funnel-shaped end);
megascleres long-shafted triaenes (with shaft directed inwards and clads on surface) and oxea
megascleres; microscleres euasters and microrhabds, without sterrasters or amphiasters.
SCOPE: Twenty six nominal genera are included in this family, although only 16 are recognised here.
REVIEWS: Hadju & van Soest, 1992
GENERA: Two groups may be differentiated based (possibly arbitrarily) on growth form.
A. Oscules at apex of long exhalant tube:
Asteropus Sollas, 1888 (type species: Stellettinopsis simplex Carter, 1879) - with oxeas,
oxyasters, true sanidasters and trichodragmata.
Disyringia Sollas, 1888 (type species: Tethyopsis dissimilis Ridley, 1884) - with microrhabds
and asters, complex exhalant tube.
Kapnesolenia de Laubenfels, 1934 - with microrhabds, without asters, simple exhalant tube.
Tethyopsis Stewart, 1870 - without microrhabds.
Tribrachion Weltner, 1882 - with microrhabds and asters, simple exhalant tube.
B. Oscules not raised on long tubes:
27
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. DesqueyrouxFaundez & 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. DesqueyrouxFaundez & Van Soest, 1997).
Stryphnus Sollas, 1886 (type species: Stryphnus niger Sollas, 1886).
q FAMILY GEODIIDAE GRAY, 1867.
SYNONYM: Erylidae Lendenfeld; Sterrastrosa Sollas.
DEFINITION: Thickly encrusting, massive to bowl-shaped growth forms; megascleres long shafted
triaenes and oxeas; sterraster microscleres always present forming superficial ectosomal crust,
sometimes also with euasters, microrhabds and spherules.
SCOPE: The family has four recognised genera, although nine named genera or subgenera have been
included in the group.
REVIEWS: Dendy (1916)
GENERA:
Erylus Gray, 1867 (type species: Stelletta mamillaris Schmidt, 1862) - uniporal afferent and
efferent surfaces oer larger oscules; triaenes short-shafted ortho- or plagiotriaenes; no ana- or
protriaenes; sterrasters usually flattened into surface aspidasters (Ref. Desqueyroux-Faundez & Van
Soest, 1997).
Geodia Lamarck, 1815 (type species: Geodia gibberosa Lamarck, 1815) (syn. Caminella
Lendenfeld; Pyxitis Schmidt, 1870; Geodia (Cydonium) Fleming, 1828; Geodia (Isops) Sollas, 1880;
Isops Sollas, 1880; Synops Vosmaer, 1882; Geodia (Sidonops) Sollas, 1889 - with aquiferous system
independently afferent and efferent, well developed large subectosomal spaces; megascleres triaenes.
microscleres sterrasters and euasters of different types (Ref. Desqueyroux-Faundez & Van Soest, 1997).
Caminus Schmidt, 1870 (type species: Caminus apiarium Schmidt, 1870) - microscleres
sphere-like asters.
Geodinella Lendenfeld (type species: ).
Pachymatisma Bowerbank, 1842 (type species: ) - surface microrhabds.
q FAMILY CALTHROPELLIDAE LENDENFELD, 1906.
DEFINITION: Massive to subsperical growth forms; megascleres calthrops and oxeas, with rays of
calthrops sometimes more than four or reduced to two, and three rays may be bifurcate; one genus
(Chelotropella) with radially oriented dichotriaenes; microscleres usually euasters, usually spherasters,
but sometimes others.
SCOPE: Six nominal genera are included, four of which are probably valid.
REVIEWS: Maldonado (1993).
GENERA:
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: ).
q FAMILY PACHASTRELLIDAE CARTER, 1875.
SYNONYMS: [Halinidae] de Laubenfels [nomen vanum]; Metrastrosa Topsent; Stretastrosa Sollas;
Astrostreptidae Topsent.
DEFINITION. Encrusting, massive and plate-shaped growth forms, with ostia and oscules on opposite
sides; megascleres calthrops, short-shafted triaenes, and oxeas; microscleres streptasters of various types
(metasters, spirasters and amphiasters), but never euasters; desmas common in some genera (`lithistid' or
`sublithistid' grades of construction).
SCOPE: Seventeen nominal genera are included here and 14 genera are currently recognised for this
family.
REVIEWS: Maldonado (1993).
GENERA:
? Acanthotriaena Vacelet, Vasseur & Lvi, 1976 (type species: ).
Brachiaster Wilson, 1925 (type species: ).
Characella Sollas, 1888 (type species: ).
Dercitus Gray, 1867 (type species: Hymeniacidon bucklandi Bowerbank, 1866) (syn.
Calcabrina Sollas, 1888; Stoeba Sollas, 1888) -dichotriaenes and spined microrhabds (sanidasters) and
sometimes toxiform microxeas (lacking in some species, e.g. Stoeba) (Ref. Maldonado, 1993;
Desqueyroux-Faundez & Van Soest, 1997).
? Lamellomorpha Bergquist, 1968 (type species: )
Pachamphilla Lendenfeld, 1907 (type species: Pachamphilla alata Lendenfeld, 1907)
Pachastrella Schmidt, 1868 (type species: Pachastrella monilifera Schmidt, 1868) (syn.
Nethea Sollas, 1888).
Poecillastra Sollas, 1888 (type species: Normania crassa Bowerbank, 1869) (syn. Normania,
sensu Bowerbank, 1869) - plate-like growth form with differentiated upper osculiferous and lower
poriferous surfaces; megascleres oxeas and triaenes, normally orthotriaenes and orthodichotriaenes;
microscleres streptasters and microxeas (Ref. Desqueyroux-Faundez & Van Soest, 1997).
Propachastrella Schrammen, 1910 (type species: ).
Siliquariospongia Hoshino, 1981 (type species: ).
Triptolemus Sollas, 1888 (type species: ).
Vulcanella Sollas, 1886 (type species: Sphinctrella horrida Schmidt, 1870) (replacement
name for Sphinctrella Schmidt, 1870 (preocc. ) (syn. Sphincterella de Laubenfels, 1936) - encrusting or
massive with pores generally dispersed and only few highly specialised oscules opening into a large
cloaca; spicules calthrops of different sizes, oxeas, rugose or microspined microxeas and asters (Ref.
Desqueyroux-Faundez & Van Soest, 1997).
Yodomia Lebwohl, 1914 (type species: ).
q FAMILY THENEIDAE SOLLAS, 1886.
DEFINITION: Mushroom shaped, more-or-less symmetrical sponges, with root-like bases forming
masses of tangled spicules; one or more oscules may be present, and specialized pore areas as well as
scattered pores distributed over entire surface; oscules and pore areas may or may not be fringed by
projecting spicules; megascleres long-shafted trienes (pro-, dicho- and anatriaenes), oxeas and oxytylote
megascleres; microscleres streptaster (plesi-, met-, spir- and amphiasters), but never euasters; asexually
produced buds formed by some species.
SCOPE: Three genera are recognised.
REVIEWS: Lendenfeld (1903), Vosmaer (1932-35)
GENERA:
Cladothenea Koltun, 1964 (type species: Cladothenea andriashevi Koltun, 1964)
Papyrula Schmidt, 1868 (type species: ) - without asters.
Thenea Gray, 1867 (type species: Tethea muricata Bowerbank, 1864) - with asters.
q FAMILY THROMBIDAE SOLLAS, 1888.
SYNONYM: Spirasigmidae Hallmann.
29
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).
q FAMILY CLIONIDAE DORBIGNY, 1852.
DEFINITION: Hadromerida displaying three different stages of growth: excavating (alpha), encrusting
(beta) or massive (gamma). Tylostyles, styles or oxeas as megascleres and streptasters and/or
microrhabds of different types as microscleres. The skeletal arrangement varies depending on the growth
form of the species or specimens. The choanosomal skeleton is confused in specimens in the alpha stage
and confused and arranged in 'ill-defined' tracts in the beta and gamma stages. The ectosomal skeleton
consists of a cortex of megascleres arranged in palisade. In alpha-stage specimens the cortex is only
present in the papillae. Microscleres can appear on the top of papillae, but they never constitute a welldifferentiated layer. Two functional types of papillae, inhalant and exhalant, are present. All the species
included in this family are able to excavate calcareous substrata, where they build galleries following a
species-specific pattern. Some species in the alpha stage are able to spread over the substratum by lateral
growth of the ectosome next to the papillae, reaching a beta or even a gamma stage of growth.
SCOPE: Fifteen nominal genera of clionids have been created, but only four of these may be valid.
REVIEWS: Ruetzler & Stone (1990), Ruetzler (1992), Bavestrello et al. (1997), Rosell & Uriz (1997),
Vicente, Ruetzler & Carballeira, 1991.
GENERA:
Alectona Carter, 1879 (type species: Gummina wallichii Carter, 1874) Boring sponges
characterised by diactinal or polyactinal spicules ornamented by conic or mushroom shaped tubercles;
different kinds of amphiasters occur, often together; oxeas may also be present (Bavestrello et al.
(1997).
Bernatia Rosell & Uriz, 1997 (type species Cliona vermifera Hancock, 1867) Excavating
Clionidae which only develop to the alpha growth stage, with tylostyles in two clearly separated size
classes as megascleres and smooth vermiform microstrongyles as microscleres. The latter are mainly
found in the basal region of the papillae. Skeletal arrangement is uncertain.
Cliona Grant, 1826 (type species: Cliona celata Grant, 1826) (syn. Papillina Schmidt, 1862;
Raphyrus Bowerbank, 1863; Euryphylle Duchassaing & Michelotti, 1864; Myle Gray, 1867; Sapline
Gray, 1867; Idomon Gray, 1867; Pronax, in part, Gray, 1867; Osculina Schmidt, 1868; Archaeocliona
Czerniavsky, 1878; Papillella Vosmaer, 1885; Papillissa Lendenfeld, 1888; Dyscliona Kirkpatrick,
1900; Gapoda de Laubenfels, 1936; Vioa Nardo) - Clionidae displaying 3 different stages of growth:
alpha, beta and gamma. Tylostyles of one size class as megascleres, although those of the ectosome
palisade are usually shorter on average. As microscleres, spirasters with different degress of
development of the actines and with intra-or interspecific variation in the arrangement of actines along
the shaft. The arrangement of microscleres varies depending on the species, sometimes absent (Rosell &
Uriz, 1997).
Clionopsis Bailey (type species: ) - megascleres oxeas with few tylostyles, microscleres long
thin spirasters and short stout spirasters resembling euasters (de Laubenfels, 1936)
Cliothosa Topsent, 1905 (type species: Thoosa hancocki Topsent, 1888) Excavating
Clionidae found only in the alpha stage, with tylostyles in 2 size classes, sometimes with intermediate
sizes. Two types of streptasters as microscleres: 1. Amphiasters with relatively long and thin actines with
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.
q FAMILY CHONDRILLIDAE GRAY, 1872 (SCHMIDT, 1862).
SYNONYM: Gummineae Schmidt, 1862 (emended to Guminida - Carter, 1875, emended to
Gumminidae - Ridley, 1884; Chondrilladae Gray, 1872, emended to Chondrillidae Lendenfeld, 1887;
Chondrosiidae Schulze, 1877.
DEFINITION: Encrusting to massive, liver-like or gelatinous sponges, often mistaken for compound
ascidians; surface often smooth with marked cortex, enriched with fibrillar collagen; megascleres
secondarily lost; euaster microscleres present or absent.
SCOPE: The family includes only five nominal and three valid genera.
REVIEWS: Wilson (1925), Topsent (1928). Under the 1964 edition of the ICZN the '50 year rule'
enabled the senior-most name Gumminidae to become nomen oblita because it had not been used for
over 50 years, and the '5 author/10 paper' provision would have allowed the continued use of
Chondrosiidae because it was considered a 'well established junior name'. However, the 1985 edition of
the ICZN does not recognise these rules, making this situation more complex. It could be argued under
Article 23 of the 1985 ICZN (Rule of Priority), that the synonymy between Chondrosia Nardo, 1847,
and Gummina Schmidt, 1862, has no bearing on the choice of family names even though Gumminidae is
based on a junior synonym. However, Article 40b states that "if a family group name has been replaced
before 1961 because of this synonymy [i.e. synonymy of the type genus], and the replacement name has
won general acceptance, it is to be maintained'. As far as I am aware Topsent (1892) was the last to use
Gumminidae, because his subsequent monographs (1904, 1928) uses Chondrosiidae (or Chondrosidae)
instead. The junior family name (Chondrosiidae Schulze, 1877) came into use over the senior family
name (Gumminidae Schmidt, 1862) around about 1892 so Article 40b can be used to justify
maintaining Chondrosiidae over Gumminidae. Subsequent authors (e.g. Wilson, 1925; Burton, 1934,
1959) use Chondrosiidae for both Chondrosia and Chondrilla, implicitly synonymising the two genera.
So, in fact, it could be argued that the junior name Chondosiidae could be used instead of the senior
available name Chondrillidae because most subsequent authors, especially contemporary ones, merge the
two genera into this family name (e.g. Lvi, 1973; Bergquist, 1978; Hartman, 1982). However, several
contemporary authors (de Laubenfels, 1936; Wiedenmayer, 1977, 1989) keep both family names
31
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) q FAMILY HEMIASTERELLIDAE LENDENFELD, 1889.
SYNONYM: [Astraxinellidae] Dendy [nomen nudum].
DEFINITION: Encrusting, cup-shaped or branching sponges; megascleres styles, oxeas or both enclosed
within compressed axial spongin fibres and plumose to plumo-reticulate extra-axial branches, or
sometimes without a definite axis; microscleres euasters, smooth or partially microspined.
SCOPE: Nine genera have been included in this family, although on five of these are recognised here.
REVIEWS: Hooper (1986), Voultsiadou-Koukoura & van Soest (1991).
GENERA:
Adreus Gray, 1867 (type species: ) - thin, stiff-branched growth form; strongly developed axial
skeleton composed of long styles; poorly developed extra-axial skeleton composed of plumose brushes
of smaller styles; asters with thin strongylote or tylote rays, which may be branched.
Axos Gray, 1867 (type species: Axos cliftoni Gray, 1867) (syn. Echinospongia Gray, 1870) branching or flabelliform growth forms; prominent surface conules; well developed axial skeleton of
styles; extra-axial skeleton of radial spicule tracts of styles at right angles to axis; asters hexaradiate to
cruciform, with thick heavily spined rays.
Hemiasterella Carter, 1879 (type species: Hemiasterella typus Carter, 1879) (syn. Epallax
Sollas, 1888; Kalastrella Kirkpatrick, 1903) - vasiform, plate-like or flattened branching growth forms;
loosely arranged axial and extra-axial skeleton of styles and/or oxeas; ectosomal asters typically with
thick, acanthose, strongylote rays.
Paratimea Hallmann, 1916 (type species: ) - thickly encrusting growth form; extra-axial
skeleton consists of long tylostyles erect on the substrate, surrounded by smaller centrotylote oxeas;
euasters are thick-centred, short-rayed oxyspherasters.
Stelligera Gray, 1867 (type species: ) (syn. Vibulinus Gray, 1867) - branching growth form;
well developed axial skeleton of long styles and oxeas; radial extra-axial skeleton consisting of long,
projecting styles surrounded by brushes of small styles, analogous to the Raspailiidae condition, with an
ectosomal crust of euasters (thick-centred, short rayed oxyspherasters).
q FAMILY LATRUNCULIIDAE TOPSENT, 1922.
SYNONYM: [Podospongiidae] de Laubenfels, 1936 [nomen nudum].
DEFINITION: Encrusting, massive, cylindrical to branched morphology, often with special oscula areas,
oscules elevated on papillae, or pore sieve-plates lying on surface in deep furrows; megascleres styles,
oxeas or strongyles, radial tracts at surface, with axial orientation in stalked forms and more confused
tracts in choanosome of massive forms; microscleres peculiar discorhabds (bearing either two whorls of
spines, two or three disks on a straight or spined axial rod, and with one swollen spined end, both ends
spined, or both ends smooth), often aggregated into dense ectosomal crust ('cortex').
SCOPE: Nine nominal genera and six valid genera are presently included in the family. NOTE. This
family (or most of it) almost certainly belongs in the Poecilosclerida (Kelly, pers. comm.)
REVIEWS: Hallmann (1914), Dendy (1922), Topsent (1928), van Soest (1984), Wiedenmayer (1989).
GENERA:
Alcyospongia de Laubenfels, 1934 (type species: Alcyospongia india de Laubenfels, 1934) megascleres are oxeas, microscleres smooth straight streptasters; externally resembles Podospongia but
its centrum of radiation is basal whereas that of the latter is near the middle of the sphere.
Diacarnus Burton, 1934 (type species: Axos spinipoculum Carter, 1879) - massive, barrelshaped, tubular or lobate-digitate, surface conulose, slippery, rubbery, oscules contractile, ostia stellate
in depressions, tough, elastic, plumo-reticulate skeleton with fibres resembling human cartiliges fully
cored by spicules, radiating towards surface, cortex dense, rubbery, offset from choanosome,
megascleres subtylostrongyles, microscleres spinulate acanthomicrorhabds (spinorhabds) - review KellyBorges & Vacelet (1995).
Latrunculia Bocage, 1869 (type species: Latrunculia cratera Bocage, 1869) (syn. Sceptrella
Schmidt, 1870; Oxylatrunculia Hoshino, 1981) - encrusting to massive; megascleres are styles or
subtylostyles, often polytylote, not differentiated morphologically in parts of the body; microscleres are
sanidasters transitional to discorhabds; whorls of spines more than two but not always distinct,
sometimes asymmetrical. References: Wiedenmayer (1989:40).
Negombata de Laubenfels, 1936 (type species: Latrunculia corticata Carter, 1879) - massive
lobate or digitate, extremely smooth surface, skeleton regular, square-meshed reticulation of fibres
cored by oxeas with strongylote ends, and extra-axial skeleton of thin wavy oxeas that become plumose
in periphery and produce ectosomal brushes, microscleres irregular spinorhabds of 2 sizes predominantly
found on outside of cortex (Kelly-Borges & Bergquist, 1995).
Negombo Dendy, 1905 (type species: )
Podospongia Bocage, 1864 (type species: Podospongia loveni du Bocage, 1864) - radial
skeleton (Topsent, 1928: 219; Wiedenmayer, 1989: 40) (syn. Lovenia du Bocage).
Sanidastra Volkmer-Ribeiro & Watanabe, 1983 (type species: ) - freshwater genus.
Sigmosceptrella Dendy, 1922 (type species: Spirastrella fibrosa Dendy, 1897) - encrusting or
massive; sharply defined fibrous ectosomal cortex containing abundant sigmodiscorhabds microscleres
which typically form a surface crust; choanosomal skeleton composed of dense and well defined fibres,
cored by styles but with light spongin, arising from the base of lamellae and ascending to the surface
nearly parallel with each other, ending at the cortex; megascleres styles or oxeas; microscleres
sigmodiscorhabds with 3-4 whorls of spines derived from sigmaform rhabds.
Strongylodesma Levi, 1969 (type species: Strongylodesma areolata Levi, 1969) [incertae
sedis]
q FAMILY PLACOSPONGIIDAE GRAY, 1867.
DEFINITION: Encrusting to lobate-digitate growth forms; surface often with network of sculptured
grooves or plates, often closable, into which ectosomal pores and oscules open; plate-like grooves on
surface rendered hard by thick cortex of closely packed sterrasters; single layer of upright tylostyles lies
in the floor of each groove; megascleres tylostyles ,also occuring in tracts within choanosomal skeleton;
microscleres sterrasters (spiraster-like) forming both dense surface crust and axial tracts; silica may be
pigmented red.
SCOPE: Only one Recent genus is known.
REVIEWS: Vosmaer & Vernhout (1902).
GENERA:
Placospongia Gray, 1867 (type species: Placospongia melobesioides Gray, 1867) (syn.
Physcaphora Hanitsch).
q FAMILY POLYMASTIIDAE GRAY, 1867.
DEFINITION: Massive, encrusting, globular, discoid, rarely stalked Hadromerida, with oscule and porebearing papillae, fistules, or mammillate projections on the upper surface. Surface slightly velvety or
very hispid. A radial skeleton of megasclere tracts and a complex specialised cortex is developed to a
greater or lesser degree in different genera. The cortex is composed of up to five components or layers in
various combinations, differentiated by megasclere size and disposition. Structural megascleres are tow
or three size categories of tylostyls, subtylostyles, strongyloxeas, or styles. Smooth centrotylote oxeas
are present in two genera, acanthose oxeas are present in one genus, oxea in trichodragmata in another,
and exotyles can be found in many species and several genera. Where known reproduction is oviparous,
with the production of parenchymella larvae following extrusion of eggs and sperm. Several genera are
only found in very deep and or polar waters.
SCOPE: Fourteen nominal genera and 10 valid genera are presently recognised for the family.
REVIEWS: Burton (1930), Boury-Esnault et al. (1992), Kelly-Borges & Bergquist (1997).
GENERA:
Atergia Stephens, 1915 (type species: ).
Polymastia Bowerbank, 1863 (type species: Polymastia mamillaris Johnston, 1842) (syn.
Rinalda Schmidt, 1870) - Massive semispherical, thickly or thinly encrusting Polymastiidae with
smooth to undulating surface; surface usually with conical, cylindrical, or mammillate papillae. The
cortical skeleton consists of up to five components in various combinations: an ectodermal palisade of
erect or fanned brushes of smaller megascleres, a middle cortical layer composed of erect or fanned
intermediate-sized megascleres, a compact basal layer of megascleres that are paratangential in
orientation and loosely arranged. This basal layer can, in some species, be highly developed producing a
33
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: ).
q FAMILY SPIRASTRELLIDAE RIDLEY & DENDY, 1886.
SYNONYMS: [Choanitidae] de Laubenfels [nomen vanum]; Acanthochaetetidae Fischer;
Tabulospongiidae Mori.
DEFINITION: Encrusting Hadromerida with tylostyles (subtylostyles or styles), rarely oxeas as
megascleres. Streptasters (spirasters and amphiasters) as microscleres usually with large actines. Some
streptasters with a short shaft may display the appearance of an euaster. The megascleres are placed in a
somewhat hymedesmoid arrangement in the choanosome, sometimes clustered in more or less welldefined bouquets. Megascleres are absent from the ectosome. The streptasters form a cortical welldefined layer. They can also be widespread in the choanosome and/or concentrated in a basal layer.
SCOPE: Five nominal genera and three valid genera are recognised.
REVIEWS: Hallmann (1914), Reitner & Engeser (1987); Rosell & Uriz (1997).
GENERA:
Acanthochaetetes Hartman & Goreau, 1975 (type species: ) (syn. Tabulospongia Mori, 1976)
- basal calcareous skeleton together with free spicules consisting of tylostyles and spirasters. These
"relict" sponges have a basal skeleton of calcite with a lamellar microstructure. The calcitic skeleton is
provided with calicles featuring vertical rows of spines and walled off at the base of the living tissue by
tabulae that are often irregular in arrangement and secondarily thickened. Increase of calicles occurs by
intramural budding; that is, a depression occurs in a wall between mature calicles, and as the latter grow
up around it, the depression becomes deeper as it developes into a mature, tissue-filled calicle.
Tylostylote siliceous megascleres occur in the tissues lining each calicle, and siliceous spirasteroid
microscleres occur in the ectosomal region. These two kinds of siliceous spicules are not, however,
embedded in a calcareous skeleton. The exhalant canal systems as they converge upon an oscule create
stellate patterns that are impressed into the calcareous skeleton as more or less distinct astrorhizae. A
calcareous epitheca provided with concentric growth lines covers the lower surface of the skeleton.
Anthosigmella Topsent, 1918 (type species: )
Sceptrintus Topsent, 1904 (type species: Sceptrintus richardi Topsent, 1904) Spirastrella Schmidt, 1868 (type species: Spirastrella cunctatrix Schmidt, 1868) (syn.
Chondrillina de Laubenfels, 1936) - with spirasters.
q FAMILY STYLOCORDYLIDAE TOPSENT, 1928.
DEFINITION: Deep water sponges with stalked, asymmetrical, globular or ovoid bodies,
characteristically growing on soft substrates; megascleres include two sizes of oxeote spicules, long
centrotylote spicules and unusual, short, terminally curved spicules, together producing peculiar radial
tracts converging towards stalk, and with spicules disposed along axis within stalk; microscleres absent
or may include microxeas, microstrongyles or asters.
SCOPE: Three nominal and two valid genera are recognised.
REVIEWS: Lvi (1973).
GENERA:
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.
q FAMILY SUBERITIDAE SCHMIDT, 1870.
DEFINITION: Massive, pedunculate, bowl-shaped or encrusting sponges, generally without surface
papillae; skeleton radial at surface, without a distinct cortex, but usually choanosome more disorganised,
occasionally with loose axial organisation and nonradial in arrangement; megascleres typically tylostyles,
subtylostyles, rarely styles or diactinal forms; tylostyles greatly modified in shape and position of head,
being lobate, pear-shaped, drop-shaped or subterminal, or occasionally missing completely;
microscleres, if present, may include spined centrotylote rods; reproduction oviparous, with asexual
buds or stolons also common.
SCOPE: Seventeen nominal genera are included in this family, although only nine of these are presently
recognised.
REVIEWS: Burton (1930), Boury-Esnault (1987), Ruetzler & Smith (1993), Kelly-Borges & Bergquist
(1994).
GENERA:
Aaptos Gray, 1867 (type species: Ancorina aaptos Schmidt, 1864) (syn. Trachya Carter, 1870;
Tuberella Keller, 1880; Tethyophaena Schmidt, 1880) - spherical, subspherical, solitary, smooth or
tuberculate surface, perfectly radial skeleton, sometimes multiple radiating subskeletons in fused
individuals, spicule tracts developed to varying degrees radiating from solid central region, tracts
become plumose below surface, with thickened cortex containing collagen, palisade of two sizes of
small spicules, and intermediate-sized spicules between plumose ectosomal tracts, primary spicules
strongyloxeas, intermediate sized spicules straight or curved styles or subtylostyles, ectosomal spicules
styles, subtylostyles and/or smaller tylostyles, rare oxeas present in some species (Kelly-Borges &
Bergquist (1994).
Caulospongia Kent, 1871 (type species: Caulospongia verticillata Kent, 1871) (syn.
Plectrodendron Lendenfeld, 1888) - distinctive growth form of foliose lamellae arranged in whorls or
plates around a central stalk, smaller tylostyles produce paratangential ectosomal skeleton, choanosomal
skeleton plumo-reticulate, with well developed spongin fibres, cored by larger tylostyles forming
longitudinal primary dendritic tracts through axis of skeleton, and ascending or oblique secondary
reticulate tracts in extra-axial skeleton, microscleres absent.
Cometella Schmidt, 1879 (type species: Cometella gracilor Schmidt, 1870) - peculiar external
stipitate form with small subspherical main body.
Laxosuberites Topsent, 1896 (type species: ) - Encrusting or massive, with ascending or rarely
interconnected strands of tylostyles forming the skeleton, without special ectosomal skeleton.
Poterion Schlegel, 1858 (type species: ) - massive vases with pseudopods for living in soft
sediments.
Prosuberites Topsent, 1893 (type species: Prosuberites longispina Topsent, 1893) Encrusting, with hispid surface, with phalynx of single long tylostyles oriented perpendicularly to (erect
on) the substrate.
Protosuberites Swartschevsky (type species: ) - smaller ectosomal tylostyles overlaying larger
ones in choanosome, the latter lying perpendicular to (erect on) the substrate.
Pseudosuberites Topsent, 1896 (type species: Hymeniacidon hyalinus Ridley and Dendy,
1887) - Massive, structured like Suberites but with a smooth surfaec due to ectosomal skeleton of
tangential (parallel to sponge surface) tylostyles.
Rhizaxinella Keller, 1880 (type species: Rhizaxinella clavigera Keller, 1880) - pedunculated,
with an axial skeleton.
Suberites Nardo, 1833 (type species: Alcyonium domuncula Olivi, 1792) (syn. Ficulina Gray,
1867; Syringella Schmidt, 1868 (sensu stricto); Suberanthus Lendenfeld, 1897; Suberella Thiele,
1905; Laxosuberella Burton, 1930; CarnleiaBurton, 1930; Laxosuberites Burton, 1930; Trachytelia
Topsent) - Massive, compact sponges with interior skeleton of densely packed tylostyles in confusion;
peripheral choanosomal skeleton in closely packed strands and dense ectosomal phalynx of tylostyles
oriented perpendicularly to the sponge surface; ectosomal tylostyles distinctly smaller than choanosomal
ones; choanosomal tylostyles without (s.s.) or with basal spines (Trachytelia).
Terpios Duchassaing & Michelotti, 1864 (type species: Terpios fugax Duchassaing and
Michelotti, 1864) (syn. Terpiosella Burton, 1930) - Thinly encrusting, with tylostyles arranged in
strands traversing the choanosome and protruding brushlike through the ectosome; loose tylostyles in
confusion between strands; spicules of one size class or with large size range, smallest sizes
35
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 FAMILY TETHYIDAE GRAY, 1867.
SYNONYM: Donatiidae Gray.
DEFINITION: Typically spherical, less often encrusting or massive growth forms (subtle differences in
growth forms important to the current generic classification); upper surface often with polygonal plates
and oscula-bearing grooves, and cribriform oscules occur at summit of sponge; basal surface often has
root-like papillae; megascleres styles or strongyloxeas, latter with asymmetrical and/or telescoped ends;
spicules not markedly tylote, frequently occuring in radial tracts; microscleres euasters (including
spherasters and micrasters); asexual reproduction by budding is common.
SCOPE: Thirteen nominal genera are included here, of which only four are presently considered to be
valid.
REVIEWS: Bergquist & Kelly- Borges (1991); Sar & Burlando (1994); Sar (1994).
GENERA:
Burtonitethya Sar, 1994 (type species: Tethya gemmiformis Burton & Rao, 1957) - black
colour; spherical, surface with tubercles, stalked; skeleton radiate, megascleres strongyloxeas;
microscleres include both megasters and micrasters; megasters are oxyasters, large, of heterogeneous
size but consistent shape; micrasters small euasters, never oxyasterose, homogeneous in size and shape.
Columnitis Schmidt, 1870 (type species: Columnitis squamata Schmidt, 1870) - large,
massive, never spherical; skeleton subradiate, megascleres are styles or strongyles; microscleres are both
megasters and micrasters; megasters are spherasters or oxyspherasters, small, homogeneous in size but
with different shapes; micrasters are small homogeneous oxyasters.
Halicometes Topsent, 1898 (type species: Cometella stellata Schmidt, 1870) - small spherical,
with moderately long stalk; skeleton radiate, megascleres strongyloxeas; microscleres include both
megasters and micrasters; megasters are large oxyasters, heterogeneous in size but consistent shape;
micrasters small, homogeneous euasters but never oxyasterote; exotyles present also.
Stellitethya Sar, 1994 (type species: Donatia fissurata var. extensa Hentschel, 1909) massively encrusting; skeleton with parallel bundles of styles or strongyles; microscleres include both
megasters and micrasters; megasters are large spherasters and/or oxyspherasters, heterogeneous in size
but similar in shape; micrasters are small, homogeneous euasters, never oxyasterote.
Tectitethya Sar, 1994 (type species: Cryptotethya crypta de Laubenfels, 1949) - massive;
skeleton subradiate composed of styles or strongyles in plumose bundles; microscleres include both
megasters and micrasters; megasters are small spherasters and/or oxyspherasters, homogeneous in size
but variable in shape; micrasters are small homogeneous oxyasters; sand incorporated into skeleton.
Tethya Lamarck, 1814 (type species: Tethya lyncurium (Linneaus, 1767)) (syn. Donatia
Nardo, 1833; Amniscos Gray, 1867; Alemo Wright, 1881; Lyncuria Nardo; Tethycordylade Laubenfels,
1934; Taboga de Laubenfels, 1936; Tethytimea de Laubenfels, 1936) - medium size spherical sponges,
never stalked but may have basal rootlets; cortex well developed; skeleton radiate composed of
strongyloxeas; microscleres include both megasters and micrasters; megasters are small spherasters
and/or oxyspherasters of homogeneous shape and size; micrasters include both small and large euasters
and oxyasters with different kinds found in ectosome and choanosome.
Tethycometes Sar, 1994 (type species: Tethycometes sibogae Sar, 1994) - small spherical,
with extremely long basal stalk; skeleton radiate composed of strongyloxeas; microscleres include both
megasters and micrasters; megasters are large oxyasters with heterogeneous size but consistent shape;
micrasters are small oxyasters with homogeneous shape and size; microstrongyles also present.
Tethyorrhaphis Lendenfeld, 1888 (type species: Tethyorrhaphis laevis Lendenfeld, 1888) - .
Xenospongia Gray, 1858 (type species: Xenospongia patelliformis Gray, 1858) - arenaceous
surface, large encrusting or disk like with a fringe of projecting spicules surrounding perimeter, adapted
for living on soft sediments; skeleton with parallel bundles of styles or strongyles; microscleres include
both megasters and micrasters; megasters are large oxyasters of heterogeneous size and shape; micrasters
are small homogeneous oxyasters; sand incorporated into both upper surface and basal layer.
q FAMILY TIMEIDAE TOPSENT, 1928.
DEFINITION: Encrusting growth form, rarely massive; surface often sculptured by stellate
subectosomal drainage canals running to oscules, and characteristically with cortex of densely packed
euasters and single layer of erect tylostyles or tracts of tylostyles running to surface; megascleres
exclusively tylostyles; microscleres include euasters (including anthasters and lophasters) or pseudasters
(amphiasters). Diplastrella has in addition a basal layer of large spherasters with branching rays
SCOPE: Two genera are recognised.

GENERA:
Diplastrella Topsent, 1918 (type species: ).
Timea Gray, 1867 (type species: Hymedesmia stellata Bowerbank, 1866).
q FAMILY TRACHYCLADIDAE HALLMANN, 1917.
SYNONYM: [Spirophorellinae] Lendenfeld [unused name, nomen oblitum].
DEFINITION: Massive or branching growth forms; oscules small (less than 1mm in diameter); ostia
scattered singly or grouped; skeleton condensed in axial region and plumoreticulate in extra-axial region,
with ascending multispicular tracts joined at infrequent intervals by single spicules; skeletal tracts
composed of spongin fibres enclosing intermixed oxeas, strongyles and styles; microscleres smooth
microstrongyles and spined vermiform spiraster-like spirules, rarely of more than two complete turns.
SCOPE: The family is represented by a single valid genus, Trachycladus.
REVIEWS: Hallmann (1917), Bergquist (1970).
GENERA:
Trachycladus Carter, 1879 (type species: Trachycladus laevispirulifer Carter, 1879) (syn.
Spirophora, sensu Lendenfeld, 1889; Spirophorella Lendenfeld, 1888) - diagnosis as for family.
q ORDER LITHISTIDA (syn. Desmophorida Hentschel, Triaenosa Sollas, Tetracladina Zittel)
DEFINITION: A problematic, polyphyletic assemblage of sponges, abundant in the CambrianQuaternary period, with many Recent relatives (all retaining articulated siliceous desma spicules,
producing rigid skeletal structure); desmas classified according to number of secondarily silicified rays
(crepis), from one (monocrepidial) to four (tetracrepidial); many species also with secondary skeletons
composed of free spicules indicating phylogenetic relationships (in this sense most orders of living
sponges have desma-bearing representatives (living relicts ?), and possession of desmas is interpreted as
a primitive feature); "lithistids" lacking free spicules are more difficult to assign to other demosponge
orders, with desma morphology being the only current diagnostic character. Three suborders with nine
families are presently retained in the taxon "Lithistida" awaiting further evidence as to their true
affinities.
Three suborders are recognized with nine families.
REVIEWS: Kelly-Borges & Pomponi (1994).
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.
q FAMILY THEONELLIDAE LENDENFELD, 1903.
SYNONYMS: [Tetracladidae] Sollas [nomen nudum]; Kaliapsidae de Laubenfels; Jereidae de
Laubenfels; Discodermiidae Schrammen; ? Phymatellinidae Schrammen; Phymaraphinidae Schrammen.
DEFINITION: Massive, cup-shaped, vase-shaped or cylindrical sponges with a narrow central cavity;
desma megascleres fused tetraclones (including tri- or tetracrepidial desmas having four arms that do not
have triaenose symmetry); free megascleres include phyllo-, disco- or dichotriaenes; microscleres
microxeas, microrhabds or microstrongyles, and streptoscleres (amphiasters or spirasters).
SCOPE: Ten living genera are presently included in this family, of which only seven are currently
thought to be valid.
REVIEWS: Wilson (1925)
GENERA:
Aulaxinia Zittel, 1878 (type species: ).
Discodermia du Bocage, 1869 (type species: Discodermia polydiscus Bocage, 1869) (syn.
Collinella Schmidt, 1880) - discotriaenes, microrhabds and microxeas.
Kaliapsis Bowerbank, 1868 (type species: ) - tetraclones, phyllotriaenes, amphiasters and
microstrongyles.
Racodiscula Zittel, 1878 (type species: ) (syn. Rimella Schmidt, 1879) - phyllotriaenes and
spirasters.
Neosiphonia Sollas, 1888 (type species: ) ([Jereopsis] Schmidt, 1879 [preocc.]) - dicho- or
trichotriaenes, streptasters, spirasters.
Reidispongia Lvi & Lvi, 1988 (type species: ).
37
Theonella Gray, 1868 (type species: Theonella swinhoei Gray, 1868) - phyllotriaenes and
microrhabds.
q FAMILY CORALLISTIDAE SOLLAS, 1888.
SYNONYM: Coscinospongiidae Lendenfeld.
DEFINITION: Massive, fan-shaped, ridge-like and folded plate-like growth forms; rigid skeleton
composed of tuberculate or arch-shaped monocrepidal desmas (dicranoclones); free megascleres phyllo-,
disco- or dichotriaenes, together with oxeas or strongyles; microscleres streptoscleres (amphiasters,
spirasters) or microxeas.
SCOPE: Six genera are included.
GENERA:
Callipelta Sollas, 1888 (type species: ) - with discotriaenes and amphiasters.
Corallistes Schmidt, 1870 (type species: ) - ectosomal megascleres dichotriaenes, microscleres
spirasters, pores simple.
Herengeria Lvi & Lvi, 1988.
Homophymia Vacelet & Vasseur, 1971 (type species: ) - with phyllotriaenes, strongyles,
amphiasters.
Iouea de Laubenfels, 1955 (type species: ).
Macandrewia Gray, 1859 (type species: ) - with phyllotriaenes, streptasters and oxeas.
q FAMILY PLEROMIDAE SOLLAS, 1888.
SYNONYM: Isoraphiniidae Schrammen; Dorydermidae Moret.
DEFINITION: Cylindrical, vase-shaped or plate-like growth forms; rigid skeleton composed of smooth
arch-shaped or armless monocrepidial desmas (megaclones with terminal cupules); free megascleres in
ectosomal skeleton dicho- or plagiotriaenes, together with oxeas (or strongyles in one genus);
microscleres streptoscleres (amphiasters, spirasters) or microxeas.
SCOPE: Four genera are included.
REVIEWS: Wilson (1925), Lvi & Lvi (1983), Lvi (1991).
GENERA:
Anaderma Lvi & Lvi, 1983 (type species: ) - anatriaenes, dichotriaenes, oxeas and
streptasters.
Costifer Wilson, 1925 (type species: ) - with heloclones, plagiotriaenes, oxeas, streptasters and
spined microxeas or microrhabds.
Lydium Schmidt, 1870 (type species: ).
Pleroma Sollas, 1888 (type species: ) - with dichotriaenes, spirasters, microxeas.
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 FAMILY NEOPELTIDAE SOLLAS, 1888.
DEFINITION: Rounded, massive or papilliform sponges; ectosomal skeleton with monocredial disks;
choanosome with monocrepidial desmas (dicranoclones), with or without free, slender oxeas;
microscleres microxeas and streptoscleres (amphiasters).
SCOPE: Only a single genus.
GENERA:
Neopelta Schmidt, 1880 (type species: ) - with discotriaenes and microxeas.
q FAMILY CLADOPELTIDAE SOLLAS, 1888.
SYNONYM: Siphonidiidae Lendenfeld.
DEFINITION: Solid, lobate sponges with tubular processes, each tube with an apical oscule; rigid
ectosomal skeleton composed of special monocrepidial desmas, branching in one plane, lacking
articulations (i.e. desmas without zygoses), whereas monocrepidial desmas in rigid choanosomal
skeleton branch in all directions, articulating with adjacent desmas (i.e. with zygoses); free ectosomal
megascleres oxytylote or oxystrongylotes, perpendicular to surface, or tangential to surface at ends of
tubular processes; microscleres absent.
SCOPE: A single genus.
REVIEWS: Lendenfeld (1903).

GENERA:
Siphonidium Schmidt, 1879 (type species: ) - with oxystyles or tylostyles.
q Suborder Anoplina
DEFINITION: Both ectosomal megascleres and microscleres absent; affinities uncertain.
Three families are included.
q FAMILY AZORICIDAE SOLLAS, 1888.
SYNONYM: Leiodermatiidae Lendenfeld.
DEFINITION: Cup-shaped, club-shaped, spherical, plate-like or fan-shaped or conical growth forms;
rigid skeleton composed of monocrepid desmas, and free skeleton with tracts of monoaxonid
megascleres (styles or oxeas) in choanosome; special ectosomal skeleton absent; microscleres absent.
SCOPE: Six nominal genera are known, of which three are presently considered to be valid.
REVIEWS: Lendenfeld (1903). Lvi (1991).
GENERA:
Gastrophanella Schmidt, 1879 (type species: ) - with subtylostrongyles and subtylostyles.
Jereicopsis Lvi & Lvi, 1983 (type species: ).
Leiodermatium Schmidt, 1870 (type species: ) (syn. Poritella Schmidt, 1879; Azorica Carter,
1873; Plakidium Lendenfeld, 1906) - with oxeas, without special ectosomal spicules.
q FAMILY DESMANTHIDAE TOPSENT, 1893.
DEFINITION: Encrusting to massive growth forms; rigid skeleton with tetracrepidial desmas
(tetraclones); free spicules consist of tracts of styles and tylostyles dispersed in choanosome; special
ectosomal spicules absent; microscleres absent.
SCOPE: Only one Recent genus.
GENERA:
Desmanthus Topsent, 1893 (type species: ) - (as for family).
Lophacanthus Hentschel, 1914 (type species: ) - only anatriaenes and desmas
q FAMILY VETULINIDAE LENDENFELD, 1903.
DEFINITION: Folded, plate-like growth forms; rigid skeleton composed of acrepid (no rayed) desmas
(sphaeroclones); free spicules strongylote megascleres scattered in choanosome; special ectosomal
spicules absent; microscleres absent.
SCOPE: Only one Recent genus.
REVIEWS:
GENERA:
Vetulina Schmidt, 1879 (type species: ) - with sphaeroclones.
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
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 FAMILY CRYPTOCOELIIDAE STEINMANN, 1982.
SYNONYM: [Neocoeliidae] Hartman (invalid name]; Verticillitidae Senowbari-Daryan.
DEFINITION: Living `sphinctozoans' with solid, cortical aragonitic skeleton consisting of a series of
solitary or colonial chambers one on top of the other, with the lowest (oldest) chambers usually partially
filled with secondary secretions of aragonite and youngest chambers containing living tissue (including a
cuticle and cells such as flattened endopinacocytes and spherule-bearing cells); calcareous chambers
contain reinforcement of radially disposed pillars; lining of atrial cavity uninterrupted (prosiphonate),
with one chamber growing forward into base of next (younger) chamber, and with numerous thin struts
(vesiculae) running from floor to roof of each chamber; struts joined by more or less horizontal
crossbars; walls of chambers and atrial lining have trefoil or multifid perforations, with perforations
corresponding to location of inhalant pores (ostia), whereas larger (oscular) openings are at apex of
chambers with the passage of exhalant water via the atrium; choanocyte chambers aphodal (with a small
canal joining chamber to exhalant canal; larvae parenchymellae that develop from a coeloblastula.
There is some anatomical variability in this group: the lining of the atrial cavity may be
calcified or not; the atrial cavity (spongocoel) may be lacking (=asiphonate, in Cryptocoelia), partly
developed from the apex down, with annular constrictions produced by the bulging inner chamber walls
(=retrosiphonate, in Stylothalamia), or uninterrupted, with a continuous cylindrical wall
[=ambisiphonate in Vaceletia progenitor; or prosiphonate in Vaceletia].
SCOPE: Only a single genus, but "Sphinctozoans" were the main reef constructors in the Middle
Triassic, dating from the Middle Cambrian.
REVIEWS: Vacelet, J. (1979), Pickett (1982), Senowbari-Daryan (1990), Vacelet, Cuif, Gautret,
Massot, Richer de Forges and Zibrowius (1992)
GENERA:
Vaceletia Pickett, 1982 (type species: [Neocoela] crypta Vacelet, 1977 ) (syn. [Neocoela]
Vacelet, 1977 [preocc.]) - with a calcitic basal skeleton composed of an irregular arrangement of
aragonitic crystals; no free spicules.
q ORDER AGELASIDA
DEFINITION: Oviparous sponges, showing (perhaps superficial) resemblance to commercial bath
sponges (Spongiidae) and biochemical similarities to Axinellidae; growth forms branching, tubular, fanshaped 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.
q FAMILY AGELASIDAE VERRIL, 1907.
SYNONYMS: [Ectyonidae] Carter [nomen oblitum].
DEFINITION: Growth form ramose, lamellate, tubular or massive, often "honeycomb" reticulate in
construction; colour frequently orange or red, texture extremely tough but compressible reflecting high
ratio of spongin fibre to spicule; skeletal structure homogeneous, reticulate, with well developed system
of large spongin fibres often containing no primary coring spicules but echinated by unique styles with
verticillate spines (acanthoxeas or acanthostrongyles), with some species having geometrically different
coring and echinating spicules, and others also having styles; sexual reproduction oviparous.
SCOPE: One genus.
REVIEWS: Hartman (1980, 1982)
GENERA:
Agelas Duchassaing & Michelotti, 1864 (type species: Agelas dispar Duchassaing and
Michelotti, 1864) (syn. Ectyon Gray, 1867).
q FAMILY ASTROSCLERIDAE LISTER, 1900.
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.
q Suborder Microcionina Hajdu, van Soest & Hooper, 1993
DEFINITION: Poecilosclerida with terminally microspined ectosomal megascleres and up to 5
categories of structural megascleres, most frequently monactinal. Microscleres are palmate chelae,
diverse toxas, but sigmas never present.
Four families.
q FAMILY ACARNIDAE DENDY, 1922.
SYNONYMS: Acarneae Dendy, 1922; Iophoneae Burton, 1929; Acarnidae Boury-Esnault, 1973; (Not
Acarniidae de Laubenfels, 1936); Cornulidae Lvi & Lvi.
DEFINITION: Encrusting, massive, flabellate or digitate growth forms, sometimes burrowing, many
genera producing fistules. Ectosomal skeleton composed of tylotes, strongyles or modified tylote
spicules bearing microspined points forming tangential and/or paratangential tracts, often irregular or
halichondroid in arrangement. Choanosomal megascleres are styles or modified styles (anisoxeas)
forming reticulate structures in massive forms (isodictyal, isotropic, anisotropic or more irregular
skeletons) or plumose or hymedesmioid skeletons in encrusting growth forms. Echinating spicules
present or absent, consisting of acanthostyles and/or unique cladotylotes. Microscleres include palmate
isochelae and toxas of various morphologies (both sometimes lost), with some genera also having other
microscleres such as bipocillae, modified anisochelae, microrhabds-like spicules (modified microxeas or
microstrongyles) and diamond-shaped microxeas (modified toxas).
SCOPE: Iophonidae contains 35 nominal genera of which only 12 are presently considered to be valid.
REMARKS: Fistular poecilosclerid genera like Acheliderma, Cornulum, Damiria, Zyzzya with tylote
ectosomal megascleres have been traditionally included in the family Coelosphaeridae Topsent. Lvi &
Lvi (1983) subsequently split Coelosphaeridae into two groups for genera with palmate isochelae
(Cornulidae Lvi & Lvi) or arcuate isochelae (Coelosphaeridae). Several non-fistular genera had
similar skeletal structure and spiculation, such as Acarnus and Iophon, possessing spined ectosomal
41
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 Lvi, 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); Lvi (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 Lvi
& Lvi, 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), Lvi & Lvi (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
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);
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 Lvi & Lvi, 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).
q FAMILY MICROCIONIDAE CARTER, 1875.
SYNONYMS: Clathriidae Lendenfeld; [Ectyoninae] of authors, in part; Ophlitaspongiidae de
Laubenfels.
DEFINITION: Encrusting, massive, lobate, fan-shaped and branching growth forms; ectosomal skeleton
composed of styles or anisoxeas (exceptionally oxeas), in erect bundles, forming a continuous crust,
lying tangential or sparsely dispersed on the surface; subectosomal skeleton relatively poorly developed;
choanosomal skeleton with well developed spongin fibres forming hymedesmoid, microcionid, plumose,
plumo-reticulate, reticulate or axially condensed tracts; spongin fibres cored by smooth or partially
spined large styles, and echinated by smooth, wholly- or partially-spined small styles or modified forms
(acanthoxeas or acanthostrongyles) embedded perpendicular to fibres; microscleres typically palmate
isochelae, sometimes contort and thickened (pseudo-anchorate, -arcuate, or Isodictya-like isochelae),
and also toxas and occasionally raphides or microxeas; sexual reproduction exclusively viviparous.
SCOPE: About 80 nominal genera have been assigned to the Microcionidae at one time or another,
although only 74 of these are validly included here (under the current concept of the family), but most of
43
these have been merged into well established taxa such that only 8 genera and 14 subgenera are now
recognised.
REVIEWS: Hallmann (1920), Hentschel (1923), Lvi (1960), Simpson (1968), van Soest (1984);
Bergquist & Fromont (1988), Hooper (1988b, 1989a, 1990, 1996), Hooper & Lvi (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.
Clathria Schmidt, 1862 (type species: Clathria compressa Schmidt, 1862).
Seven subgenera are recognised:
Clathria (Clathria) Schmidt, 1862 (type species: Clathria compressa Schmidt, 1862) (syn. Allocia,
Hallmann, 1920; Antherochalina Lendenfeld, 1887; Bipocillopsis Koltun, 1964; Dictyociona
Topsent, 1913; Labacea de Laubenfels, 1936; Ligrota de Laubenfels, 1936; Litaspongia de
Laubenfels, 1936; Marleyia Burton, 1931; Paradoryx Hallmann, 1920; Pitalia Gray, 1867;
Ramoses de Laubenfels, 1936; Thalyseurypon de Laubenfels, 1936) - with single category of
auxiliary styles forming ectosomal and subectosomal skeletons, differentiated from category of
principal styles in the choanosomal skeleton.
Clathria (Wilsonella) Carter, 1885 (type species: Wilsonella australiensis Carter, 1885) (syn.
Clathriopsamma Lendenfeld, 1888; Aulenella Burton & Rao, 1932) - principal or auxiliary
spicules coring fibres may be absent, wholly or partially replaced by foreign debris.
Clathria (Microciona) Bowerbank, 1863 (type species: Microciona atrasanguinea Bowerbank, 1863)
(syn. Abila Gray, 1867; Aaata, sensu de Laubenfels, 1930; Anaata de Laubenfels, 1932;
Axocielita de Laubenfels, 1936; Cionanchora de Laubenfels, 1936; Damoseni de Laubenfels,
1936; Fisherispongia de Laubenfels, 1936; Folitispa de Laubenfels, 1936; Hymantho Burton,
1930; Hymeraphia, in part, Hentschel, 1912; Leptoclathria Topsent, 1928; Paratenaciella
Vacelet & Vasseur, 1971; Pseudanchinoe Burton, 1929; Sophax Gray, 1867; Wetmoreus de
Laubenfels, 1936) - hymedesmoid choanosomal skeleton composed of a basal spongin fibres
lying on substrate, with ascending non-anastomosing fibre nodes ("microcionid" skeleton), and
erect principal, echinating and auxiliary spicules perpendicular to substrate.
Clathria (Dendrocia) Hallmann, 1920 (type species: Clathria pyramida Lendenfeld, 1888) (syn.
Stylotellopsis Thiele, 1905) - single undifferentiated category of auxiliary style forming both
choanosomal and ectosomal skeletons.
Clathria (Axosuberites) Hallmann, 1920 (type species: Esperiopsis cylindrica Ridley and Dendy, 1886)
(syn. Axosuberites Topsent, 1893; Tenaciella Hallmann, 1920) - well differentiated axial
(compressed, reticulate) and extra-axial skeletons (radial, plumose, plumo-reticulate);

echinating acanthostyles absent.
Clathria (Isociella) Hallmann, 1920 (type species: Phakellia jacksoniana Dendy) - choanosomal
skeleton composed of wide-meshed homogenous spongin fibres or spicule tracts, forming
renieroid or subrenieroid reticulation, cored by smooth principal styles, without echinating
megascleres.
Clathria (Thalysias) Duchassaing & Michelotti, 1864 (type species: Spongia juniperina Lamarck,
1813) (syn. Colloclathria Dendy, 1922; Echinonema Carter, 1881; Rhaphidophlus Ehlers,
1870; Tenacia Schmidt, 1870; ? Thalassodendron Lendenfeld, 1888) - - with two distinct
categories of auxiliary styles forming ectosomal and subectosomal skeletons, the smaller styles
producing plumose surface brushes, and differentiated principal styles in the the choanosomal
skeleton.
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:
Echinochalina (Echinochalina) (type species: Ophlitaspongia australiensis Ridley, 1884) monactinal principal spicules echinate fibres.
Echinochalina (Protophlitaspongia) (type species: Siphonochalina bispiculata Dendy, 1895) diactinal principal echinate fibres.
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.
Ophlitaspongia Bowerbank, 1866 (type species Ophlitaspongia papilla Bowerbank, 1866) reticulate, ladder-like, fibre skeleton of parallel ascending fibres with regular cross-connecting isodictyal
reticulate fibres arising from a hymedesmoid basal layer of spongin fibre; only ascending fibres cored by
plumose columns of entirely smooth subtylostyles, with few or no spicules coring transverse connecting
fibres; tracts of spicules protrude through fibres, diverging towards surface, but no true echinating
spicules; mineral skeleton strictly plumose (not reticulate);ectosomal skeleton with protruding
choanosomal styles and single smooth auxilliary subtylostyles, the latter also scattered throughout the
mesohyl; microscleres smooth wing-shape toxas.
? Pandaros Duchassaing & Michelotti, 1864 (type species: Pandaros acanthifolium
Duchassaing & Michelotti, 1864) - ectosome without special spicules but subectosomal skeleton
prominently radial, with very long individual auxiliary subtylostyles protruding through surface,
choanosomal skeleton reticulate, with flattened spongin fibres (trabeculae) cored by slightly rhabdose
principal subtylostyles and echinated by sparse acanthostyles, microscleres absent.
q FAMILY RASPAILIIDAE HENTSCHEL, 1923.
SYNONYM: Euryponidae Topsent.
DEFINITION: Encrusting, massive, lobate, fan-shaped or branching growth forms, usually with a very
hispid surface; specialised ectosomal skeleton consists of brushes of small thin styles or oxeas,
surrounding individual long thick styles or oxeas; choanosomal skeleton varies from a compressed axial
skeleton, to plumo-reticulate or exclusively reticulate structures; spongin fibres usually completely
enclose coring spicules (choanosome styles, oxeas or both); fibres echinated by spined styles, or
modifications to styles; microscleres usually absent, sometimes single raphides or bundles
(trichodragmata); reproduction typically oviparous.
SCOPE: Fifty one nominal genera have been included in the family, but only 19 of these are presently
considered to be valid (of which 2 are presently incertae sedis).
45
REVIEWS: Bergquist (1970), Hooper (1991), Hooper & Lvi (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; extraaxial 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, extraaxial 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.) (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
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 Lvi 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 bulbousswollen, or stellate-swollen heads and spined points; microscleres absent. References: Hooper (1991).
Lithoplocamia Dendy, 1922 (type species: Lithoplocamia lithistoides Dendym 1922) (syn.
Monectyon Lvi & 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 ("pegtop" 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.
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
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.
q FAMILY RHABDEREMIIDAE TOPSENT, 1928.
DEFINITION: Encrusting, massive, bulbous or digitate growth forms; skeleton without axial
compression, usually plumose or plumo-reticulate, composed of light spongin fibres cored by bouquets
of entirely smooth, slightly spined or entirely spined rhabdostyles; microscleres are normal or contort
sigmas, microstyles and thraustoxeas (all with or without microspines).
SCOPE: Four nominal genera are included in this family but only one of these is presently considered to
be valid.
REVIEWS: Hooper (1990), van Soest & Hooper (1993).
GENERA:
Rhabderemia Topsent, 1890 (type species: Microciona pusilla Carter, 1876) (syn.
Rhabdosigma Hallmann, 1916; Nisibaris de Laubenfels, 1936; Hallmannia, [in part], Burton, 1930) encrusting, massive or lobate branching growth forms, skeleton usually plumo-reticulate, although some
species may be hymedesmoid or plumose, composed of smooth, partially or entirely spined rhabdostyles,
microscleres smooth or microspined, with one or more combinations of sigmoid (contort and/or true
sigmas), microstyles (true forms and toxa-like thraustoxeas), and true toxas. References: Hooper
(1990:65); van Soest & Hooper (1993).
q Suborder Myxillina Hajdu, van Soest & Hooper, 1993
DEFINITION: Poecilosclerida with microscleres consisting of tridentate-derived chelae, but never
toxas; ectosomal megascleres are diactinal, although aniso- terminations commonly occur, and terminal
spination of these spicules is absent, or if present they are usually coarse or irregular.
Eight families are included.
q FAMILY ANCHINOIDAE TOPSENT, 1928.
SYNONYM: Phorbasidae de Laubenfels.
DEFINITION: Encrusting, massive and branching sponges; surface with characteristic groups of pores
(areoles); megascleres are smooth diactinal spicules (oxeotes or strongylotes), grouped together or
scattered in the ectosome but never forming a crust, lying perpendicular, tangential, or paratangential to
the surface; choanosomal diactinal spicules (usually identical to those in the ectosome but sometimes
reduced to only acanthostyles), form thick plumose or plumo-reticulate tracts in the choanosome, with
only poorly developed spongin fibres, and echinated by 1-2 sizes of acanthose styles; microscleres
include arcuate isochelae, bipocilli and sigmas, never toxas or raphides.
SCOPE: Seventeen nominal genera are included in this family, of which only six are valid.
REVIEWS: Bergquist & Fromont (1988), Voultsiadou-Koukoura & van Soest (1991), van Soest
(1984), Wiedenmayer (1989).
GENERA:
? Anomolissa de Laubenfels, 1934 (type species: Anomolissa amaza de Laubenfels, 1934) ectosomal oxeas erect on surface; malformed arcuate chelae-like microscleres.
Dendoricella Lundbeck, 1905 (type species: ) - ectosomal tornotes resembling oxeas,
choanosomal oxeas forming dendritic tracts, no echinating spicules, microscleres are arcuate isochelae.
Hamigera Gray, 1867 (type species: Cribrella hamigera Schmidt, 1862) (syn. Amoibodictya
Zahn et al., 1977) - oscules in sieve-plates; ectosomal diactinal spicules form fans around areolate pore
areas; choanosomal skeleton composed of plumose tracts of smooth subtylostyles and/or strongyles or
strongylote tornotes, with echinating subtylostyles; microscleres are arcuate isochelae. References:
Bergquist & Fromont (1988:73); Voultsiadou-Koukoura & van Soest (1991); Wiedenmayer (1989:97);
Shaw (1927:429).
Kirkpatrickia Topsent, 1912 (type species: Tedania variolosa Kirkpatrick, 1907) - flabellate
or digitate; oscules grouped in sieve-plates (areoles); ectosomal skeleton composed of ectosomal
strongylote tornotes or strongyles in both tangential tracts and whispy brushes on the surface (the latter
around pores); choanosomal skeleton composed of loosely agglomerated primary and secondary fibres
cored by choanosomal styles, curving out towards the surface in the peripheral skeleton; megascleres are
choanosomal styles with smooth or spined bases microscleres are absent. References: Kirkpatrick
(1907:279); Desqueyroux-Faundez & Van Soest (1995).
Myxodoryx Burton, 1929 (type species: Lissomyxilla hanitschi Kirkpatrick, 1907) - massive;
ectosomal skeleton composed of smooth tylotes with mucronate bases forming tangential and oblique
tracts at the surface; choanosomal skeleton confused reticulate, with branching fibres cored by smooth
choanosomal styles and echinated by acanthostyles with verticillate spination; microscleres are absent.
References: Kirkpatrick (1907:275).
Phorbas Duchassaing & Michelotti, 1864 (type species: Phorbas amaranthus Duchassaing
and Michelotti, 1864) (syn. Anchinoe Gray, 1867; Clathrissa Lendenfeld, 1888; Grayax de Laubenfels,
1936; Lissopocillon Ferrer-Hernandez, 1916; Merriamium de Laubenfels, 1936; Plumohalichondria
Carter, 1876; Pronax Gray, 1867; Pronaxella Burton, 1931; Suberotelites Schmidt, 1868;
Stylostichon, sensu Topsent, 1892) - oscules grouped in sieve-plates; ectosomal skeleton with a crust of
isochelae and diactinal spicules which form fans disposed at right angles and tangential to the surface;
heavily spined acanthostyles core plumose or plumo-reticulate choanosomal tracts, or these may be
occasionally replaced by smooth diactinal tornotes; echinating acanthostyles also heavily spined;
microscleres are arcuate isochelae and sigmas. References: Bergquist & Fromont (1988:70);
Voultsiadou-Koukoura & van Soest (1991); Lvi (1963:43); Zea (1987:165); van Soest (1984:86); de
Laubenfels (1936:64); Uriz (1988:80).
Plocamionida Topsent, 1927 (type species: Microciona ambigua Bowerbank, 1866) (syn.
Hymendectyon Bakus, 1966) - encrusting; ectosomal skeleton contains paratangential brushes of
tornotes; choanosomal skeleton with hymedesmoid architecture with long smooth styles erect on the
substrate, singly or in bundles, and protruding through the surface, long acanthostyles and short
acanthostrongyles forming a reticulate basal skeleton, and also short echinating acanthostyles erect on
the substrate and clustered around the bases of the long spicules; microscleres are arcuate isochelae.
References: Bowerbank (1866:137; 1874:pl.25); Arneson (1903); Topsent (1927:304).
Podotuberculum Bakus, 1966 (type species: ) - oscules grouped in sieve-plates, smooth
choanosomal strongyles, acanthostyles, no microscleres.
q FAMILY COELOSPHAERIDAE HENTSCHEL, 1923.
DEFINITION: Spherical, club-shaped, tubular, and cushion-shaped growth forms, frequently burrowing
or excavating coralline substrates (with long, open and/or blind fistules on upper surface bearing oscules
and ostia poking above the substrate); ectosomal skeleton a tangential crust of smooth diactinal (usually
tylote) spicules, occasionally secondarily lost; choanosomal skeleton reduced, composed of a delicate
reticulation of smooth or acanthose styles, occasionally oxeas or strongyles, forming plumoreticulate
tracts and cavernous internal chambers; microscleres include sigmas, arcuate isochelae, occasionally
modified to unguiferous, thaumatose or birotulate forms; toxas absent.
SCOPE: Forty eight nominal genera are presently included in this revised concept of Coelosphaeridae,
of which 25 appear to be valid.
REVIEWS: Hentschel (1923), Topsent (1928), Lvi (1973), van Soest (1984), Bergquist & Fromont
(1988).
GENERA:
Abyssocladia Lvi, 1964 (type species: Abyssocladia bruuni Lvi, 1964) - erect; no ectosomal
skeleton; choanosomal skeleton composed of an axial and radial skeleton cored by long, smooth and
polytylote choanosomal styles; microscleres are modified arcuate isochelae (thaumatochelae) and
sigmas. References: Lvi (1964:78).
49
Acanthodoryx Lvi, 1961 (type species: Acanthodoryx fibrosa Lvi, 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: Lvi, 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 & Lvi, 1976 (type species: Coelodischela diatomorpha
Vacelet, Vasseur & Lvi, 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 & Lvi (1976:57); Lvi & Lvi (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)
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 Lvi & Lvi, 1979 (type species: Lepidosphaera hindei Lvi & Lvi, 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: Lvi & Lvi
(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.
q FAMILY CRAMBIIDAE LEVI, 1963.
SYNONYM: Name emended from Crambidae Levi, 1963, a junior homonym of Crambidae Latreille,
1810 (Insecta: Lepidoptera).
DEFINITION: Encrusting or massive growth forms; ectosomal megascleres consist of smooth
subtylostyles, usually standing perpendicular to surface; choanosomal megascleres are smooth or
acanthose styles-tylostyles forming hymedesmoid, plumose or plumoreticulate skeletal structures, with a
secondary interlocking desma ("sublithistid") skeleton common; microscleres anchorate or unguiferous
isochelae.
SCOPE: Nine nominal genera are included here in this family, of which only six are probably valid.
REVIEWS:
GENERA:
Crambe Vosmaer, 1880 (type species: Crambe crambe (Schmidt, 1862)) (syn. Tetranthella
Lendenfeld, 1893) - incrusting; ectosomal skeleton translucent, hispid, conulose, detachable, composed
51
of a tangential layer of styles-subtylostyles; choanosomal skeleton composed of desmas forming a semirigid basal skeleton, and styles or subtylostyles forming a plumose skeleton; megascleres are asteroid
desmas (sometimes absent) and style or subtylostyles; microscleres are pseudasters, spined microrhabds
and anchorate isochelae. References: Vosmaer (1880:135); Lvi (1963:16); Vacelet, Vasseur & Lvi
(1976:54); Uriz (1988:69).
Discorhabdella Dendy, 1924 (type species: ) - smooth ectosomal subtylostyles, long
choanosomal acanthostyles with swollen lumpy bases forming erect hymedesmoid skeleton,
pseudoasterose acanthostyles, microscleres are anchorate and unguiferous isochelae and spined
microrhabds (oxydiscorhabds).
Leptosiopsis Topsent, 1927 (type species: ) - ectosomal styles polytylote, choanosomal styles
of two sizes forming plumose hymedesmoid skeleton, microscleres are anchorate anisochelae.
Lithochela Burton, 1928 (type species: ) - ectosomal styles, large and small desmas forming
basal skeleton, microscleres are unguiferous isochelae.
Monanchora Carter, 1883 (type species: Monanchora clathrata Carter, 1883) (syn. Okadaia
de Laubenfels, 1936; Neofolitispa Bergquist, 1965) - ectosomal skeleton with erect brushes of
subtylostyles arising from the choanosomal skeleton; choanosomal skeleton with a spongin enforced
plumose or plumo-reticulate skeleton of irregular bundles of subtylostyles; megascleres are
subtylostyles of a single category; microscleres are one or more categories of anchorate (unguiferous or
spatulate) isochelae typically with a grooved shaft. References: van Soest (1984:42; 1990:250);
Bergquist & Fromont (1988:37).
Psammochela Dendy, 1916 (type species: Psammochela elegans Dendy, 1916) - ectosomal
skeleton with a fine-meshed reticulation of small sand grains; choanosomal skeleton irregularly
reticulate, composed of thin fibres with poor spongin, cored with sand and styles, and numerous sand
particles and spicules dispersed within the mesohyl; megascleres are styles, strongylote styles, sometimes
vestigial or lost completely; microscleres are anchorate (tridentate) isochelae and sigmas. References:
Dendy (1916:126); Wiedenmayer (1989:67).
q FAMILY CRELLIDAE HENTSCHEL, 1923.
DEFINITION: Encrusting, massive, club-shaped and branching growth forms; choanosomal skeleton
regularly reticulate or plumo-reticulate, composed of bundles of smooth oxeas; ectosomal skeleton with
a thick crust of tangentially placed acanthostyles and/or acanthoxeas; acanthose spicules may also be
embedded perpendicular to skeletal tracts and/or erect on basal spongin (= echinating basal
acanthostyles), and dispersed within the choanosome between the tracts of smooth diactines;
microscleres arcuate isochelae, anisochelae and sigmas.
SCOPE: Twenty five nominal genera are included in this family, of which only 13 are recognised here,
although some of these are not well established and some are possibly synonyms.
REVIEWS: Lundbeck (1909), de Laubenfels (1936), Lvi (1973), van Soest (1984), Lvi and Lvi
(1984), Bergquist & Fromont (1988).
GENERA:
Anisocrella Topsent, 1927 (type species: Anisocrella hymedesmina Topsent, 1927) encrusting growth form; ectosomal skeleton composed of tangential layer of acanthostyles;
subectosomal skeleton composed of small (rudimentary) smooth diactinal spicules in bundles;
choanosomal skeleton composed of an erect basal layer of acanthostyles; microscleres are arcuate
isochelae and unguiferous anisochelae. References: Topsent (1927:234)
Barbozia Dendy, 1921 (type species: Barbozia primitiva Dendy, 1921) - massive subspherical,
with small fistules; choanosomal skeleton reticulate; megascleres are diactinal oxeote or strongylote;
microscleres are palmate anisochelae, amphiaster-like oxydiscorhabds and sigmas. References: Dendy
(1921:132)
Crella Schmidt, 1862 (type species: Cribrella elegans Schmidt, 1862) (syn. Aaaba de
Laubenfels, 1936; Crella Gray, 1867; Crellinspira de Laubenfels, 1936; Crelloxea Hechtel, 1983;
[Cribrella] Schmidt, 1862 [preocc.]; Grella de Laubenfels, 1936; Ramosichela de Laubenfels, 1950) encrusting; ectosomal skeleton composed of a tangential layer of acanthostyles or acanthoxeas;
choanosomal skeleton composed of smooth tornotes or strongyles and echinating acanthostyles, or in
encrusting forms of an erect basal layer of acanthostyles; microscleres may include arcuate isochelae,
sometimes absent. References: van Soest (1984:77); Bergquist & Fromont (1988:75); Vacelet &
Boury-Esnault (1987:163).
Crellina Hentschel, 1914 (type species: Crellina tubifex Hentschel, 1914) - encrusting;
ectosomal skeleton well developed, crust-like composed of acanthostrongyles lying both tangential and
in erect brushes on the surface; choanosomal skeleton poorly developed, with erect basal acanthostyles
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); Lvi & Lvi (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 crisscrossed 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).
q FAMILY HYMEDESMIIDAE TOPSENT, 1928.
DEFINITION: Persistently encrusting growth form; oscules and ostia usually on papillae, or ostia
grouped over subdermal cavities; spined bases of intermingled large choanosomal styles and smaller
acanthostyles embedded in a basal layer of spongin, standing perpendicular to the substrate; smooth,
often polytylote, diactinal (tornotes, anisotornotes or oxeas) or sometimes monactinal spicules (styles),
occur singly or form bundles on the surface; microscleres palmate, arcuate or unguiferous isochelae,
sigmas, forceps, and sometimes also anisochelae; toxas never present.
SCOPE: Twenty two nominal genera are presently included in this family, although only 13 are
recognised here.
REVIEWS: Topsent (1928), Bergquist & Fromont (1988).
GENERA:
Acanthancora Topsent, 1927 (type species: Leptosia schmidti Topsent, 1895) - encrusting;
ectosomal skeleton composed of smooth tornotes or anisotornotes, some with polytylote bases;
choanosomal skeleton composed of acanthostyles erect on the substrate; microscleres are arcuate
isochelae ornamented with small spines, and chiastosigmas. References: Topsent (1927; 1928:274); van
Soest (1984:85)
Arndtanchora de Laubenfels, 1936 (type species: Leptosia sirventi Topsent, 1927) - two sizes
of echinating acanthostyles, microscleres are sigmas and arcuate isochelae, some with bifid alae.
Chiastosia Topsent, 1928 (type species: ) - ectosomal tornotes symmetrical, large
choanosomal acanthostyles, smaller echinating acanthostyles, microscleres are arcuate isochelae and
chiastosigmas.
Dolichacantha Hentschel, 1914 (type species: Dolichacantha macrodon Hentschel, 1914) encrusting; ectosomal skeleton composed of tylostrongyles; choanosomal skeleton with erect basal layer
of echinating monactine rhabds, short acanthostyles; microscleres are palmate isochelae and sigmas.
References: Hentschel (1914:11); Topsent (1928:55).
Hymedesmia Bowerbank, 1863 (type species: Hymedesmia zetlandica Bowerbank, 1863) (syn.
Sclerilla Schmidt, 1868; Hymenamphiastra de Laubenfels, 1930; Ferrerhernandezia de Laubenfels,
1936; Hymoxenia Alander, 1938; Quindesmia de Laubenfels, 1950) - encrusting; ectosomal skeleton
53
composed of smooth ectosomal anisotornotes; choanosomal skeleton with hymedesmoid skeleton,

composed of one or two sizes of acanthostyles erect on the substrate; microscleres are arcuate isochelae
or anisochelae and sigmas. References: Dendy (1924:); van Soest (1984:79); Lundbeck (1910:112);
Bergquist & Fromont (1988:65); Vosmaer (1933:215).
Hymesigmia Topsent, 1927 (type species: Hymesigmia japycina Topsent, 1927) - ectosomal
skeleton consists of smooth anisotylotes; choanosomal skeleton composed of two sizes of acanthostyles
erect on the substrate; microscleres are sigmas in dragmata, no isochelae. References: Topsent (1927;
1928:282); de Laubenfels (1936:87); van Soest (1984).
Jelissima de Laubenfels, 1936 (type species: Hymedesmia tenuissima Thiele, 1905) - no
special ectosomal skeleton, choanosomal skeleton composed of columns of acanthoxeas, acanthostyles
erect on substrate, microscleres are quadridentate (arcuate) isochelae, and sigmas.
Poecilochela de Laubenfels, 1936 (type species: Myxilla decepta Kirkpatrick, 1907) encrusting; ectosomal skeleton with smooth ectosomal strongyles with mucronate bases; choanosomal
skeleton with a single category of acanthostyle erect on basal skeleton; microscleres are arcuate
isochelae, bipocillate chelae and unguiferous isochelae (possibly both the latter are modified arcuate).
References: de Laubenfels (1936:88); Kirkpatrick (1907:278).
Pseudohalichondria Carter, 1886 (type species: Pseudohalichondria clavilobata Carter,
1886) - massive, flabellate; ectosomal skeleton with subtylostyles; choanosomal skeleton composed of
thick fibres cored with sand and subtylostyles; microscleres are spined arcuate isochelae. References:
Carter (1886:454); de Laubenfels (1936:126).
Spanioplon Topsent, 1890 (type species: Hymeniacidon armatura Bowerbank, 1866) (syn.
Acanthoxa Hentschel, 1914) - oscules grouped in sieve plates; ectosomal skeleton composed of smooth
anisotornotes, smooth or microspined choanosomal styles, small acanthostyles or acanthoxeas dispersed
(possibly homologues of echinating spicules or true microscleres), chelae absent. References: Topsent
(1890; 1892:116; 1904:171; 1928:236); de Laubenfels (1936:88).
Spirorhabdia Topsent, 1918 (type species: ) - polytylote ectosomal oxeote tornotes, echinating
acanthostyles, microscleres are spined rhabds.
Stylopus Fristedt, 1885 (type species: Stylopus coriaceus Fristedt, 1885) (syn. Leptosia, sensu
Topsent, 1892) - ectosomal skeleton membraneous, with strongyles forming plumose tracts throughout
the choanosome and at the surfaec they expand into brushes supporting the dermal membrane;
choanosomal skeleton with basally attached acanthostyles orientated vertically to the base and extending
into the ectosome; microscleres absent. References: de Laubenfels (1936:88); Bergquist & Fromont
(1988:69); Bakus (1966:474).
Trachostylea Topsent, 1928 (type species: Trachostylea semota Topsent, 1928) - encrusting,
halichondroid, spongin absent, megascleres styles and subtylostyles completely microspined;
microscleres absent. Review: de Laubenfels(1936:79).
q FAMILY MYXILLIDAE TOPSENT, 1928.
SYNONYMS: [Desmacidinae] Schmidt; [Desmacidonidae] Gray; [Desmacidontidae] de Laubenfels;
Esperiopsidae Hentschel.
DEFINITION: Encrusting, massive, fan-shaped and branching sponges; specialised ectosomal skeleton
composed of diactinal tylotes or tornotes with smooth or microspined, slightly swollen bases, arranged
as bouquets or lying paratangential or perpendicular to the surface; choanosomal skeleton composed of
isotropic, anisotropic or plumose tracts of smooth or partially spined monactinal or diactinal
choanosomal megascleres (or choanosomal spicules replaced within the skeleton by ectosomal
megascleres), sometimes echinated by small acanthose styles; spongin development variable, usually
consisting of light spongin cementing spicule together at their nodes, but sometimes with heavy fibres;
microscleres anchorate isochelae and/or derivatives (spatulate, unguiferous or birotulate chelae,
sometimes anisochelate), sometimes together with palmate isochelae, smooth sigmas and forceps.
SCOPE: Forty nine nominal genera are included in this family, of which 33 are recognised here,
although many of these are very poorly known, many are possible synonyms of more established taxa,
and some are of uncertain placement.
REVIEWS: Hentschel (1923), Lvi (1973), van Soest (1984), Bergquist & Fromont (1988), Hooper
and Lvi (1989), Wiedenmayer (1989), Hajdu et al. (1993).
GENERA:
Amphiastrella Dendy, 1896 (type species: Phloeodictyon birotuliferum Carter, 1886) massive, erect fistulose growth form; choanosomal skeleton consists of diactinal megascleres which are
strongyles or tylotes forming an irregular mesh; ectosomal strongyles or tylotes with smooth bases
(same as in choanosome), lie tangential to surface forming a compact dermal layer; microscleres are
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 Lvi, 1963 (type species: Ectyonancora flabellata Lvi, 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;
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
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); DesqueyrouxFaundez & 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);
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);
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); Lvi
(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.
Myxilla (Ectyomyxilla) Lundbeck, 1909 (type species: Ectomyxilla kerguelensis Hentschel,

1914) - echinating strongly spined styles.
Myxilla (Styloptilon) Cabioch, 1968 (type species: Styloptilon ancoratum Cabioch, 1968) with plumose skeleton rather than isotropic skeleton.
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; DesqueyrouxFaundez & 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 Lvi, 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)].
q FAMILY PHORIOSPONGIIDAE LENDENFELD, 1888.
SYNONYMS: Collosclerophoridae Dendy.
DEFINITION: Encrusting, massive, flabellate or digitate growth forms; eEctosomal skeleton frequently
absent, replaced by arenaceous or spicular detritus, but typically with areolate oscular sieve plates on
surface; ectosomal and choanosomal megascleres undifferentiated, usually strongyles (sometimes
secondarily lost); choanosomal spicules are auxiliary megascleres (of ectosomal origin), whereas
principal spicules are absent; microscleres arcuate isochelae, sometimes modified to unguiferous or
birotulate forms.
SCOPE: Seventeen nominal genera are provisionally included in this family, although only 11 of these
are considered here to be valid.
REVIEWS: nil.
GENERA:
Anomomyxilla Burton, 1934 (type species: Desmacidon psammodes Hentschel, 1911) surface with pore sieve-plates; ectosomal skeleton composed of strongyles, of the same form as found in
the choanosome, forming loose dermal brushes supporting a close-meshed tangential skeleton of sand
and foreign spicules (sandy crust); choanosomal skeleton consists of an irregular plumoreticulate
57
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.
q FAMILY TEDANIIDAE RIDLEY & DENDY, 1886.
DEFINITION: Encrusting, massive or digitate sponges; choanosomal skeleton predominantly plumoreticulate or even plumose, composed of tracts of smooth or spined monactinal megascleres, or smooth
diactinal megascleres, enclosed within light or moderate spongin fibres, or with no visible fibres and
spicules merely cemented together at their nodes; ectosomal spicules are diactinal, tylotes or strongyles,
usually with spined bases, lying tangential, paratangential or erect on the surface, although usually not in
bundles; microscleres onychaetes; chelae are absent.
SCOPE: Ten nominal genera and seven probably valid genera are included here, although several of
these are poorly known.
REVIEWS: Burton (1932), van Soest (1984), Bergquist & Fromont (1988); Desqueyroux-Faundez &
Van Soest (1995).
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 Lieberkhn, 1859) (syn. Trachytedania Ridley, 1881; Tedaniopsis Dendy, 1924;
Paratedania Burton, 1929; Oxytedania Sar, 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].
q Suborder Mycalina Hajdu, van Soest & Hooper, 1993
DEFINITION: Poecilosclerida with microscleres consisting of sigmancistra derivatives and megascleres
being subtylostyles, with swollen bases and faintly constricted necks (mycalostyles), usually of a single
smooth category (never echinating).
Five families are included.
q FAMILY CLADORHIZIDAE DE LAUBENFELS, 1936.
DEFINITION: Small symmetrical sponges mostly found in the abyssal zones, with diagonal, radiating
supporting processes and basal root adaptations for living in soft sediments; choanosomal skeleton
consists of an axis composed of monactinal (styles) or occasionally diactinal megascleres (oxeas), from
which radiating extra-axial tracts diverge to the ectosome; microscleres include isochelae, sigmas,
forceps or spear-shaped microstyles.
SCOPE: Although thirteen nominal genera are included in this family, only five of these are recognised
here.
REVIEWS: Lvi (1964, 1973), Bergquist & Fromont (1988), van Soest (1984); Bergquist (1972);
Boury-Esnault & van Beveren (1982); de Laubenfels (1936).
GENERA:
59
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 extraaxial 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); Lvi (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 & Lvi (1989:440); Ridley & Dendy (1887:102).
q FAMILY GUITARRIDAE BURTON, 1929.
DEFINITION: Encrusting, massive or ramose growth forms; special ectosomal spicules absent, but
choanosomal spicules may protrude through surface; choanosomal skeleton is reticulate, isodictyalreticulate or plumoreticulate, with a single category of subtylostyles; microscleres placochelae and
modifications, sometimes also with palmate isochelae and sigmancistras.
SCOPE: Six nominal genera are inlcuded in this family, only four of which are currently recognised:
furthermore, one of these (Hoplakithara) may be a synonym of Guitarra.
REVIEWS: Topsent (1928), Hajdu et al. (1993).
GENERA:
Euchelipluma Topsent, 1909 (type species: Euchelipluma pristina Topsent, 1909) (syn.
Desmatiderma Topsent, 1928) - ramose; ectosomal skeleton with scattered styles and sometimes
desmas; choanosomal skeleton with a dense plumose or plumoreticulate skeleton composed of parallel
fibres cored by plumose tufts of styles, and in some species also with a semi-rigid basal skeleton of
desmas; megascleres are styles, strongyles and sometimes monocrepidial desmas; microscleres are
palmate isochelae, placochelae and two categories of sigmancistras. References: Topsent (1928;
1929:216, 308).
Guitarra Carter, 1874 (type species: Guitarra fimbriata Carter, 1874) (syn. Pocilloguitarra
Topsent, 1928) - massive; ectosomal skeleton composed of erect brushes of oxeas arising from the
choanosomal skeleton; choanosomal skeleton a regular reticulation or isodictyal reticulation of oxeas;
megascleres are oxeas, rarely stylote; microscleres are placochelae, bipocillae (biplacochelae) and small
palmate isochelae with spines in bouquets. References: Bergquist & Fromont (1988:40); Lee
(1987:474); Burton (1929)
Hoplakithara Kirkpatrick, 1907 (type species: Hoplakithara dendyi Kirpatrick, 1907) cushion shaped; ectosomal skeleton with a rigid external armored skeleton produced by adjacent
spheroidal heads of exotyles, which are arranged in radiating bundles in the form of inverted cones
(diverging on the surface); choanosomal skeleton with radiating exotyles and scattered strongyles;
megascleres are strongyles and exotyles (tylotes) with large spherical spined heads; microscleres are
fimbriated placochelae and sigmas. References: Kirkpatrick (1907); Koltun (1964:46).
Tetrapocillon Brondsted, 1924 (type species: Tetrapocillon novaezealandiae Brondsted,

1924) - encrusting; ectosomal skeleton with brushes of styles from protruding choanosomal fibres;
choanosomal skeleton an irregular reticulation of styles; megascleres are styles of only one category;
microscleres are tetrapocilli and small palmate isochelae with spines in bouquets. References: Bergquist
& Fromont (1988:46); Brondsted (1924:457)
q FAMILY DESMACELLIDAE RIDLEY & DENDY, 1886.
SYNONYMS: Biemnidae Hentschel; Sigmaxinellidae Lvi.
DEFINITION: Encrusting, massive, cup-shaped, fan-shaped and branching growth forms; megascleres
usually styles, sometimes also including oxeas or strongyles; spicules typically enclosed within plumose,
reticulate, halichondroid-reticulate or compressed axial fibres; microscleres are diverse, always
consisting of sigmas, and often including microxeas of several sizes, raphides in bundles or individually,
toxas, microstrongyles and spheres.
SCOPE: Sixteen nominal genera are included here in the revised Desmacellidae, of which only seven are
thought to be valid.
REVIEWS: Bergquist (1970), Hooper (1984a), Wiedenmayer (1977), van Soest (1984, 1987),
Bergquist & Fromont (1988), Hooper, Capon & Hodder (1991).
GENERA:
Biemna Gray, 1867 (type species: Desmacidon peachi Bowerbank, 1866) (syn. Allantophora
Whitelegge, 1907; Toxemna Hallmann, 1917) - plumose or plumo-reticulate choanosomal skeleton,
with variable development of spongin fibres cored by styles or subtylostyles of a single size, an erect
ectosomal skeleton and often shaggy surface, and microscleres include sigmas and raphides, and
sometimes microxeas, commas, microstrongyles and spheres. References: Bergquist & Fromont
(1988:29); van Soest (1984:133); Hooper (1984:32)
Desmacella Schmidt, 1870 (type species: Desmacella pumilio Schmidt, 1870) (syn.
Sigmatoxella de Laubenfels, 1936; Sigmotylotella Burton, 1932; Tylodesma Thiele, 1903; Tylosigma
Topsent, 1904;) - reticulate, plumo-reticulate and vaguely halichondroid choanosomal skeleton, poorly
developed spongin fibres, a tangential halichondroid ectosomal skeleton all cored by tylostyles of one or
more sizes, microscleres sigmas and raphides. References: Hooper (1984:15); van Soest (1984:136);
Bergquist & Fromont (1988:34); Wiedenmayer (1977:161).
? Kerasemna Pulitzer-Finali, 1982 (type species: Kerasemna tenuistyla Pulitzer-Finali, 1982)
- reticulate choanosomal skeleton, well developed spongin fibres, tangential ectosomal skeleton, cored
by subtylostyles, microscleres sigmas, raphides and possibly toxas. References: Pulitzer-Finali
(1982:105); Hooper (1984:32).
Merlia Kirkpatrick, 1908 (type species: Merlia normani Kirkpatrick, 1908) (syn. Noronha
Kirkpatrick, 1909) - encrusting, with free spicules consisting of tylostyles, acanthose raphides, commas
and clavidiscs, together with a fused basal calcareous skeleton, composed of chambers with clinogonal
microstructure (of chaetetid grade of construction). The calcareous skeleton is composed principally of
calcite (Vacelet, 1979), but may be absent (= deficient) in specimens growing in deep caves (Pouliquen,
1972). References: Pulitzer-Finali (1972:5; 1983:535); van Soest (1984:24)
? Microtylostylifer Dendy, 1924 (type species: Microtylostylifer anomalus Dendy, 1924) irregularly reticulate choanosomal skeleton, poorly developed spongin fibres cored by larger styles, and a
specialized tangential ectosomal skeleton of small microstyles, microscleres consist of microstyles only.
References: Bergquist & Fromont (1988:34); Dendy (1924:382).
Neofibularia Hechtel, 1965 (type species: Fibularia massa Carter, 1882) (syn. Fibularia
sensu Carter, 1882) - irregular isodictyal reticulate choanosomal skeleton, well defined spongin fibres, a
membraneous or tangential ectosomal skeleton, cored by diactinal spicules (strongyles, oxeas) or
sometimes styles, often vestigial, microscleres sigmas, raphides, microxeas (commas in one species).
Toxic sponges. References: van Soest (1984:141); Hechtel (1965:); Hartman (1967:2).
Sigmaxinella Dendy, 1897 (type species: Sigmaxinella australiana Dendy, 1897) (syn.
Sigmaxia Hallmann, 1916) - condensed axial skeleton, plumose or plumo-reticulate extra-axial
skeleton, well developed spongin fibres, a plumose, erect ectosomal skeleton, all cored by styles of a
single size, microscleres are sigmas and microxeas. References: Hooper (1984:1); Hallmann
(1916:521).
q FAMILY HAMACANTHIDAE GRAY, 1872.
DEFINITION: Encrusting to massive sponges; skeleton consists of a tangential ectosomal and reticulate
choanosomal skeletal components, composed of monactinal (styles), diactinal (oxeas), or both sorts of
megascleres producing multispicular tracts and forming irregular, plumo-reticulate or reticulate
structures, with little or no associated spongin; scattered monactinal or diactinal megascleres, or bundles
61
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), Lvi (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); Lvi (1973:611).
q FAMILY MYCALIDAE LUNDBECK, 1905.
DEFINITION: Encrusting, massive, fan-shaped, cup-shaped and branching growth forms; subectosomal
sculpturing, grooves and ridges often found on the surface, within which are usually found the ostia;
skeleton radially arranged, plumose or plumo-reticulate, composed of styles or oxeas enclosed in
spongin fibres; without specialised ectosomal spicules, although choanosomal spicules may form dense
brushes at the surface; microscleres anisochelae, but may also include many other forms - palmate
isochelae with geometric modifications, sigmas, toxas and raphides.
SCOPE: Twenty nine nominal genera are now included in the Mycalidae, only 11 of which are currently
recognised. Many subgenera are in use for Mycale, but the validity and status of some these is
questionable.
REVIEWS: Hechtel (1965), van Soest (1984); Bergquist & Fromont (1988), Hajdu (in press).
GENERA:
Anomomycale Topsent, 1924 (type species: Desmacidon titubans Schmidt, 1870) - ectosomal
skeleton composed of shorter ectosomal styles; choanosomal skeleton an irregular reticulation of
multispicular fibres cored by larger choanosomal styles; microscleres are arcuate anisochelae and
sigmas. References: Topsent (1928:206); Lundbeck (1905:41).
Arenochalina Lendenfeld, 1887 (type species: Arenochalina mirabilis Lendenfeld, 1887).
Esperiopsis Carter, 1887 (type species: Esperia villosa Carter, 1874) - erect, digitate or
massive; ectosomal skeleton plumose from protruding spicules; choanosomal skeleton plumose or
plumoreticulate with multispicular tracts of styles, ascending, diverging moreso than anastomosing, and
protruding through the surface; in digitate species skeleton may be condensed with a radial extra-axis;
megascleres are styles or subtylostyles, sometimes also monocrepidial desmas; microscleres are palmate
isochelae, sometimes also with sigmas, toxas and raphides. References: Burton (1929:429); Lundbeck
(1905:9); Bergquist & Fromont (1988:25); Hooper & Lvi (1989:437).
Gomphostegia Topsent, 1896 (type species: ).
Mycale Gray, 1867 (type species: Hymeniacidon lingua Bowerbank, 1863) (syn. Acamas, in
part, Duchassaing & Michelotti, 1864; Acamasina de Laubenfels, 1936; Aegogropila Gray, 1867;
Carmia Gray, 1867; Esperella Vosmaer, 1885; Esperia, in part, Nardo, 1833; Grapelia Gray, 1867;
Mycalecarmia de Laubenfels, 1936; Oxycarmia de Laubenfels, 1954; Parisociella Burton, 1952;
Protoesperia Czerniavsky, 1879; Pseudoesperia Carter, 1886; Raphioderma Bowerbank, 1869;
Raphiodesma, in part, Bowerbank, 1874; Rhaphidotheca Saville-Kent, 1872; Sceptrospongia Dendy,
1926). References: Bergquist & Fromont (1988:17); van Soest (1984:9,28); Topsent (1924:85).
Five subgenera availabled:
Mycale (Mycale) (type species: Hymeniacidon lingua Bowerbank, 1863) - ectosomal skeleton is a
multilayered halichondroid tangential skeleton carried by the plumose terminal choanosomal
fibres; choanosomal skeleton largely dendritic and becoming more plumose near periphery,
fibres generally poor in spongin, and also with confused single spicules dispersed within the
mesohyl; megascleres are subtylostyles only; microscleres are anisochelae, some species also
with sigmas, toxas, isochelae, microxeas and raphides in trichodragmata or singly.
Mycale (Carmia) (type species: Hymedesmia macilenta Bowerbank, 1866) - ectosomal skeleton
without specialised structure, although plumose tracts from peripheral choanosomal skeleton
extend into surface as brushes; choanosomal skeleton consists of dendritic-plumose tracts of
subtylostyles; microscleres are anisochelae, sigmas, toxas, raphides and micracanthoxeas.
Mycale (Acamasina) (type species: Acamas laxissima Duchassaing & Michelotti, 1864) - massively
encrusting; without ectosomal skeleton, detachable dermis contains only microscleres and
loose megascleres; choanosomal skeleton consists of a coarse rectangular reticulation of thick
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: Lvi (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.
q FAMILY AXINELLIDAE CARTER, 1875.
SYNONYM: Bubarinae Hentschel.
DEFINITION: Encrusting, massive, branching, fan-shaped and tubular growth forms; encrusting species
may consolidate sedimentary particles at the surface of the substratum; surface usually hispid from
projecting spicules; megascleres styles, oxeas, strongyles (sometimes sinuous) in all combinations;
skeleton typically divided into distinct axial (or basal in encrusting forms) and extra-axial components;
main skeleton typically condensed in axis, consisting of smooth straight spicules in most genera, or
tuberculate or spined, annular, flexuous, U-shaped or vermiform strongyles in some genera; extra-axial
skeleton plumose or plumoreticulate, with tracts of smooth straight spicules, sometimes rhabdose
spicules arising perpendicular to the axis and ascending to the surface; axial and extra-axial
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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 & Lvi
(1993); Alvarez (in press).
GENERA:
Acanthella Schmidt, 1862 (type species: Acanthella acuta Schmidt, 1862) - encrusting, fanshaped, 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 plumoreticulate 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 Lvi & Vacelet, 1958 (type species: Bubaropsis curvisclera Lvi 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
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) - fanshaped, 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,
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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 extraaxial 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 extraaxial 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
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 extraaxial 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, cupshaped 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
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.
q FAMILY DESMOXYIDAE HALLMANN, 1917.
SYNONYM: Higginsiinae de Laubenfels.
DEFINITION: Encrusting, massive or branching sponges; megascleres are monactinal (styles), diactinal
(oxeas), or both, contained within widely spaced multispicular spongin fibres, or with little or no
spongin associated, forming reticulate tracts, with poorly developed or no axial compression and poorly
differentiated axial and extra-axial skeletons (notably disorganised or slightly plumose); ectosomal
skeleton a crust or palisade of smaller oxeotes with spines (occasionally smooth); microscleres smooth
or spined microxeas, often centrangulate or strongly bent at the centre, and sometimes raphides in groups
(trichodragmata) or singly, and in one genus acanthose cladotoxa and birotules are also present.
SCOPE: Fourteen nominal genera, of which eight are recognised here, are presently included in the
family.
REVIEWS: Wiedenmayer (1977), Hooper & Lvi (1993b).
GENERA:
Acanthoclada Bergquist, 1970 (type species: Acanthoclada prostrata Bergquist, 1970) massive to encrusting growth forms; choanosome is not condensed, without an obvious axis; the
skeleton is lax, fibrous, consisting of ascending spongin fibres, reinforced with small quantities of
collagenous spongin, and cored by smooth styles and profusely echinated by smooth rhabdostyles; long
axial styles also protrude through the fibres, and peripheral fibres terminate in brushes of long
centrangulate or toxiform oxeas, which form conulose surface projections; ectosome is packed with
acanthose cladotoxas; microscleres are acanthose cladotoxas and birotules.
? Caesarellum de Laubenfels, 1936 (type species: Bubaris gallica Topsent, 1894) centrotylote strongyle megascleres, microscleres spiny microrhabds.
Didiscus Dendy, 1922 (type species: Didiscus placospongioides Dendy, 1922) - Massiveamorphous to lobate; spicules strewn in confusion and sporadically grouped in spongin-reinforced
directionless tracts; ectosomal skeleton is a gradation from a perpendicular palisade to a tangential or
paratangential arrangement of spicules; ectosomal skeleton also with "discorhabd" microscleres,
apparently derived from centrotylote spirasters, arranged perpendicular to the surface and concentrated
near surface; surface with sculptured grooves and subdermal drainage canals (modified from Diaz et al.,
1993).
Halicnemia Bowerbank, 1864 (type species: Halicnemia patera Bowerbank, 1864) encrusting, disk-shaped, with a very hispid surface; choanosomal skeleton consists of a basal layer of
longitudinal spicule tracts lying on the substrate, including a halichondroid reticulation of styles and
polytylote smooth oxeas; extra-axial styles or tylostyles are embedded and erect on this basal layer,
protruding through and tangential to the surface; specialized ectosomal spicules consist of erect brushes
of acanthose, centrangulate or straight, microxeas.
Heteroxya Topsent, 1904 (type species: Heteroxya corticata Topsent, 1898) - encrusting
growth form; choanosomal skeleton consists of a basal layer of spongin lying on the substrate, without a
complete mineral skeletal structure, containing only acanthoxeas dispersed without any appreciable
order on the basal spongin and strewn about the mesohyl; extra-axial skeleton consists of oxeas or
tornotes perpendicular to the ectosome and protruding through the surface, but these are not embedded
in basal spongin; ectosomal skeleton contains a perpendicular palisade of acanthoxeas, through which
subectosomal oxeas protrude.
Higginsia Higgin, 1877 (type species: Higginsia coralloides Higgin, 1877) (syn. Dendropsis
Ridley & Dendy, 1886; Desmoxya Hallmann, 1917) - erect, lamellate, massive, vasiform or lobate
growth forms; surface conulose, papillose, often silt covered; skeletal structure ranges from
hlaichondrioid with a partially compressed, reticulate axis, and an irregularly plumo-reticulate extraaxial region (Higginsia), a compressed axis and a radial, non-plumose extra-axial region (Dendropsis),
to a lax plumose or plumo-reticulate axial and extra-axial region, without axial compression or regional
differentiation of the skeleton (Desmoxya); spongin fibres usually poorl developed although collagen in
the mesohyl is heavy, usually with numerous megascleres and microscleres scattered between the main
skeletal tracts; all skeletal tracts cored by monactinal and/or diactinal megascleres. Ectosome without
specialised spiculation, but with extra-axial spicule tracts (1 or 2 categories of megascleres) protruding
through the surface. Megascleres are oxeas, strongyles and/or styles of 1-3 sizes. Microscleres are
spined, centrangulate curved or straight microxeas, sometimes also with raphides occurring singly or in
bundles (trichodragmata).
Julavis de Laubenfels, 1936 (type species: Tedania levis Kirkpatrick, 1900) - thinly encrusting,
ectosome with tangential crust of slender acanthostrongyles often contort, choanosome with
hymedesmoid skeleton of erect or scattered long styles and polytylote tornostyles, with dense isotropic
overlaying skeleton of more robust acanthostrongyles with a tendency to unite into bundles;
microscleres raphides scattered singly or in trichodragmata.
? Microxistyla Topsent, 1928 (type species: ) - encrusting growth form and stony texture;
ectosomal skeleton is membraneous, without specialized spiculation; choanosomal skeleton consists of
a confused reticulation of smooth choanosomal principal styles, bonded together with small amounts of
collagenous spongin. Microscleres are centrangulate microxeas.
Myrmekioderma Ehlers, 1870 (type species: Alcyonium granulatum Esper, 1830) (syn.
Acanthoxifer Dendy, 1905; [?] Callites Schmidt, 1868 [virtually unrecognisable]; Anacanthaea Row,
1911; Neoprosypa de Laubenfels, 1954) - massive or encrusting growth forms; surface hispid and with
distinctive canals and grooves forming polygonal tuberculate plates; choanosome is a compressed and
confused mass of acanthoxeas and oxeas, strongyles or less frequently styles, forming irregular
ascending multispicular tracts bound together with sparse collagen; extra-axial skeleton dense
paratangential layer of acanthoxeas, through which project larger choanosomal styles. Ectosomal
skeleton lacks specialized spiculation, but has projecting subectosomal tracts of acanthoxeas forming
closely compacted, adjacent brushes lying perpendicular to the surface. Microscleres are raphides
occurring singly or in bundles (trichodragmata).
Parahigginsia Dendy, 1924 (type species: Parahigginsia phakellioides Dendy, 1924a) lamellate, lobate growth forms; choanosomal skeleton consists of a condensed axial reticulation of
oxeas and acanthoxeas, forming irregular tracts, without distinct fibres or obvious collagenous spongin;
extra-axial skeleton is not clearly defined from the axis, consisting of irregularly branching primary and
secondary tracts of choanosomal oxeas, which do not protrude through the surface but become more
69
ordered and compact near the periphery; ectosome skeleton contains only spongin and scattered, curved
acanthoxea microscleres.
q FAMILY DICTYONELLIDAE VAN SOEST, DIAZ AND POMPONI, 1990.
DEFINITION: Choanosomal skeleton lacking any axial compression or marked differentiation between
axial and extra-axial regions, but has spongin-enforced dendritic or plumose choanosomal spicule tracts
and a fleshy conulose surface; no ectosomal mineral skeleton; megascleres include oxeas, styles or both
in equal proportion; microscleres absent.
SCOPE: Six nominal genera and five probably valid genera are presently included in the family.
REVIEWS: Carballo, Uriz & Garcia-Gomez, 1996.
GENERA:
Dactylella Thiele, 1898 (type species: ) - (Burton 1959:257).
Dictyonella Schmidt, 1868 (type species: Dictyonella cactus Schmidt, 1868) (syn.
Phacanthina Vosmaer, 1912).
Liosina Thiele, 1898 (type species: ) - .
Scopalina Schmidt, 1862 (type species: ) - encrusting growth form, prominently conulose,
ectosomal skeleton with brushes of auxiliary styles, choanosomal skeleton hymedesmoid with heavy
basal spongin fibres on substrate and ascending non-anastomosing fibre nodes, the latter cored by
plumose brushes of auxiliary styles, echinating megascleres absent, microscleres absent.
Tethyspira Topsent, 1890 (type species: Tethea spinosa Bowerbank, 1874) - encrusting and
massive growth forms, choanosomal skeleton consists of a condensed basal layer of spongin with fibre
nodes (microcionid skeleton), without specialized choanosomal spicules, but with plumose columns of
long extra-axial styles; specialized raspailiid ectosome is absent; extra-axial skeletal tracts echinated by
highly modified microscleres resembling acanthoxeas with very large perpendicular spines (Hooper,
1991).
q FAMILY HALICHONDRIIDAE VOSMAER, 1887.
SYNONYMS: Spongosoritidae Topsent; ? Petromicidae Topsent; Hymeniacidonidae de Laubenfels.
DEFINITION: Encrusting to massive growth forms, sometimes with specialised fistules on the upper
surface; principle megascleres are oxeas, sometimes with accessory styles; choanosomal skeleton
consists of a high density of spicules arranged in vague, poorly defined, directionless tracts
("halichondroid" structure), or spicules in complete confusion; there is often marked subectosomal or
vestibular cavities; microscleres usually absent, occasionally raphides.
SCOPE: Fifty three nominal genera are included in the family, of which only 16 appear to be valid.
REVIEWS: Bergquist (1970); van Soest et al. (1990), Diaz et al. (1991, 1992), Hooper et al. (1997).
GENERA:
Amorphinopsis Carter, 1886 (type species: Amorphinopsis excavans Carter, 1886g) (syn.
Migas Sollas, 1908; Tumata de Laubenfels, 1936; Milene de Laubenfels, 1954; Prostylissa Topsent,
1925) - encrusting growth form; surface with stellate spicule patterns; ectosomal skeleton with a distinct
tangential reticulation composed of thick bundles of oxeotes, often with blunt terminations, occasionally
stylotes, "echinated" by small styles; peripheral regions of the skeleton characteristically include smaller
oxeotes and styles than larger choanosomal megascleres; choanosome is halichondroid, without any
compression of the axial region; extra-axial skeleton is poorly developed, verging on halichondroid, but
with some discernible plumose tracts of styles (occasionally oxeas) near the periphery (modified from
van Soest et al. 1990).
Axinyssa Lendenfeld, 1897 (type species: Axinyssa topsenti Lendenfeld, 1897) (syn.
Pseudaxinyssa Burton, 1931; Axinomimus de Laubenfels, 1936) - Massive-amorphous; ectosome
largely organic, tough, with sparsely scattered spicules; choanosomal skeleton disorganised with spicules
strewn in confusion and/or composed of vaguely ascending, widely spaced vertical tracts of large oxeas,
forming loose bundles, with spicule tracts protruding through surface to produce conules; choanosome
with poor or moderate spongin fibres but heavy collagen; spicules oxeas, strongyloxeas or styles usually
of only one size class (modified from van Soest et al. 1990; Hooper & Bergquist, 1992; Diaz et al.,
1993).
Ciocalypta Bowerbank, 1863 (type species: Ciocalypta penicillus Bowerbank, 1863) (syn.
Apatospongia Marshall, 1892; Leucophloeus Carter, 1883; Uritiaia Burton, 1932) - with pointed,
blind fistules; fistules characteristically semi-transparent parchment-like; with distinct ectosomal
tangential reticulation of spicule tracts, occurring as bundles or single spicules; with ectosomal styles
together with predominantly stylote choanosomal megascleres (from van Soest et al. 1990).
Collocalypta Dendy, 1905 (type species: Collocalypta digitata Dendy, 1905) - with fistulose
habit and architecture; with a choanosomal skeleton highly collagenous, with a central column of
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,
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).
q FAMILY CHALINIDAE GRAY, 1867.
SYNONYMS: Renieridae Schmidt; Haliclonidae de Laubenfels; Adociidae de Laubenfels.
DEFINITION: Encrusting, massive, cup-shaped, fan-shaped and branching growth forms, usually with
spongy and delicate consistency; when present ectosomal skeleton consists of a special, tangential,
unilayered, unispicular, isotrophic reticulation of oxeas bound by nodal spongin; choanosomal skeleton
consists of an isodictyal reticulation of uni- or paucispicular primary tracts of oxeas, rarely
multispicular, interconnected by uni- or paucispicular secondary tracts, and spicules are bonded together
at their nodes of junction by small amounts of collagenous spongin, or they may be fully enclosed within
light spongin fibres and form more robust reticulations; microscleres, if present, include only sigmas or
toxas; parenchymella larvae are incubated and are completely and uniformly ciliated or have a bare
posterior cap fringed by longer cilia.
SCOPE: There are 26 nominal genera but only 13 of these are probably valid.
REVIEWS: Lendenfeld (1887a), Burton (1927a, 1934a), Griessinger (1971), Bergquist & Warne
(1980), Van Soest (1980), Desqueyroux-Faundez (1984, 1987), de Weerdt (1986, 1987), Fromont
(1991, 1992).
GENERA:
Acervochalina Ridley, 1884 (type species: Chalina limbata (Montagu, 1818)) - no special
ectosomal skeleton (which is membraneous), and spicules and fibre terminations from ascending
choanosomal skeleton pierce the surface; choanosomal skeleton loosely arranged open reticulation of
paucispicular primary lines interconnected by uni- or paucispicular secondary lines usually more than 1
spicule length; spicules enveloped in abundant collagenous spongin; microscleres absent (de Weerdt &
van Soest, 1986).
Adocia Gray, 1867 (type species: Isodictya simulans Bowerbank, 1866) (syn. Amorphina
Schmidt, 1870; Asychis Gray, 1867; Menanetia Topsent, 1896; Halichondriella Burton, 1931;
Halichoclona de Laubenfels, 1932) - ectosomal skeleton composed of distinct unilayered tangential
reticulation of single oxeas bound at their nodes by spongin; microscleres absent (van Soest, 1980).
Calcifibrospongia Hartman, 1979 (type species: ) - with a hypercalcified basal skeleton of
spherulitic microstructure, together with free spicules producing a reticulation of strongyles.
Cladocroce Topsent, 1892 (type species: Cladocroce fibrosa Topsent, 1892) - choanosomal
skeleton isodictyal unispicular typical of family but further strengthened by dendritic mutispicular tracts,
broad at the base, tapering into thin endings (van Soest, 1980).
Dendroxea Griessenger, 1971 (type species: ) - choanosomal unispicular isotropic skeleton is
strengthened by dendritic multispicular tracts arising from a dense spicular mass at the base of the
sponge (van Soest, 1980).
Gellius Gray, 1867 (type species: ) - no ectosomal skeleton; and ectosomal region
membraneous (although paucispicular tracts from ascending choanosomal skeleton may protrude
through surface); microscleres sigmas and toxas (van Soest, 1980).
Haliclona Grant, 1835 (type species: Spongia oculata Linneaus, 1759) (syn. Chalina
Bowerbank, 1863; Chalinodendron Lendenfeld, 1887; Chalinula Schmidt, 1868; Euchalinopsis
Lendenfeld, 1887; Lessepsia Keller, 1883; Phylosiphonia Lendenfeld, 1887; Rhaphisia Topsent, 1892;
Kallypilidion de Laubenfels, 1954; Nara de Laubenfels, 1954; Veluspa Miklucho-Maclay) - without
ectosomal skeleton; ectosomal region membraneous; choanosomal skeleton unispicular isodictyal;
without microscleres (van Soest, 1980). Recent revision: de Weerdt (1989).
Katiba de Laubenfels, 1954 (type species: Katiba milnei de Laubenfels, 1954) - very thinly
encrusting, pigment literally wipes off the substrate (in the type species turning from greenish to
reddish), with dense, fleshy ectosome, collagenous choanosome; skeleton with spicules in confusion
consisting of very small oxeas, raphides in trichodragmata and singly.
Orina Gray, 1867 (type species: Halichondria angulata Bowerbank, 1866) - ectosomal
tangential unilayered reticulation of single spicules bound at the nodes by spongin; microscleres sigmas
and toxas (van Soest, 1980).
Reniera Nardo, 1847 (type species: Reniera aquaeductus Schmidt, 1862) (syn. Chalinula
Schmidt, 1868) - ectosomal skeleton lying parallel (tangential) to the surface, unispicular, irregular;
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).
q FAMILY NIPHATIDAE VAN SOEST, 1980.
DEFINITION: Encrusting, massive, fan-shaped, vase-shaped and branching growth forms, often with
chimney-like oscular processes; ectosomal skeleton consists of a dense multispicular, three-dimensional,
paratangential reticulation of diactinal spicules (oxeas or strongyles), usually more compact than the
choanosomal skeleton; erect spicule brushes characteristically at the surface; choanosomal skeleton a
reticulation of ascending and transverse-connecting spongin fibres, cored by multispicular tracts of
oxeas; interstitial spicules also common; microscleres, if present, are sigmas or microxeas.
SCOPE: Fourteen nominal genera, seven of which are probably valid, are included here.
REVIEWS:
GENERA:
Aka de Laubenfels, 1936 (type species: Acca insidiosa Johnson, 1899) (syn. [Acca] Johnson
(preocc.); Siphonodictyon Bergquist, 1965) - tangential ectosomal skeleton uniformly meshed
producing a smooth surface; choanosomal reticulation grading into a pulpy mucusy central part; spongin
scarse; microscleres absent; burrowing into dead coral substrate producing short thick fistules (van
Soest, 1980).
Amphimedon Duchassaing & Michelotti, 1864 (type species: Amphimedon compressa
Duchassaing & Michelotti, 1864) (syn. Pachychalina Schmidt, 1868; Hemihaliclona Burton, 1937) ectosomal reticulation is of uniform mesh size; choanosomal primary spicule tracts barely protrude
through the surface producing an optically smooth surface; spongin dominant; microscleres absent (van
Soest, 1980).
Cribrochalina Schmidt, 1870 (type species: Spongia vasulum Lamarck, 1814) - tangential
ectosomal reticualtion is obscured by perpendicular spicule brushes producing an optically smooth but
microscopically hispid surface; spongin dominant; choanosomal meshes long and narrow, condensed
peripherally; microscleres absent (van Soest, 1980); sponge surface typically sticky, slightly mucusy,
growth forms often vases, fans, cups, thin-walled.
Gelliodes Ridley, 1884 (type species: ) - tangential ectosomal reticulation obscured by
protruding tufts of the choanosomal primaries producing a rough conulose surface; spongin present;
secondary fibres largely reduced; microscleres abundant sigmas (van Soest, 1980).
Hemigellius Burton, 1932 (type species: Hemigellius rudis (Topsent, 1901)) - tangential
ectosomal reticulation obscured by protruding tufts of the choanosomal skeleton producing a conulose
surface; spongin absent; microscleres sigmas (van Soest, 1980).
Hoplochalina Lendenfeld, 1887 (type species: Hoplochalina incrustans Lendenfeld, 1887)
(syn. Haliclonissa Burton, 1932) - tangential ectosomal reticulation is obscured by protruding tufts of
the choanosomal primaries, producing a conulose surface, and by perpendicular spicule brushes;
interconnecting choanosomal tracts reduced; spongin absent; no microscleres (van Soest, 1980).
Microxina Topsent, 1916 (type species: Microxina benedeni (Topsent, 1901)) (syn.
Calyxadocia de Laubenfels, 1936; Sigmaxinyssa Kirkpatrick, 1907) - ectosomal tangential reticulation
obscured by protruding tufts of the choanosomal primary spicule tracts, producing a conulose surface;
spongin absent; microscleres microxea (van Soest, 1980).
Niphates Duchassaing & Michelotti, 1864 (type species: Spongia digitalis Lamarck, 1814)
(syn. Dasychalina Ridley & Dendy, 1886) - ectosomal tangential reticulation is obscured by protruding
tufts of the choanosomal primary spicule tracts, producing a rough, conulose surface; spongin dominant;
microscleres rare sigmas (van Soest, 1980).
q FAMILY CALLYSPONGIIDAE DE LAUBENFELS, 1936.
DEFINITION: Encrusting, massive, vase-shaped, tubular, fan-shaped and branching growth forms;
surface characteristically sculptured with conules or ridges, and usually has an optically visible lace-like
73
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).
q FAMILY PHLOEODICTYIDAE CARTER, 1882.
SYNONYM: Oceanapiidae van Soest.
DEFINITION: Encrusting, massive, lobate, or more frequently spherical and tubular growth forms
buried in the substrate, usually with fistules on upper surface bearing apical oscules, occasionally
excavating coralline substrates; ectosomal skeleton multilayered, irregular, tangential reticulation of
diactinal spicules (oxeas or strongyles), forming a distinct, usually detachable. parchment-like crust;
choanosomal skeleton an irregular reticulation of diactinal spicules forming multispicular tracts,
typically producing a pulpy effect, with or without spongin fibres, together with an irregularly dispersed
isotropic reticulation of single spicules scattered between these major tracts; microscleres may include
centrangulate sigmas and toxas.
SCOPE: There are 10 nominal genera, with only five presently recognised, presently included in this
family.
REVIEWS: Dendy (1905, 1922), Wilson (1925), Topsent (1928), van Soest (1980), de Weerdt (1985).
GENERA:
Calyx Vosmaer, 1887 (type species: Calyx nicaensis (Risso, 1826)) (syn. Vagocia de
Laubenfels, 1936; Lieberkuhnia Balsamo-Crivelli, 1860) - thin-walled fans or vase-like growth form;
fistules absent; ectosomal skeleton is multilayered or single-layered spicule reticulation; choanosomal
skeleton dominated by longitudinal multispicular tracts, anastomosing and intertwined, producing a
stringy texture, with single spicules dispersed between major tracts; microscleres may include sigmas
and/or 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 (100-250m) oxeote megascleres; microscleres absent; fistules walls thin,
consisting only of an ectosomal crust and longitudinal supporting spicule tracts (van Soest, 1980).
q FAMILY PETROSIIDAE VAN SOEST, 1980.
SYNONYM: [Nepheliospongiidae] Clarke sensu Weidenmayer, in part.
DEFINITION: Typically massive, vase-shaped or volcano-shaped sponges, sometimes encrusting,
bulbous, and less commonly branching growth forms; texture characteristically stony, brittle, reflecting
that in most species siliceous spicules are clearly dominant over spongin; ectosomal skeleton an
isotropic reticulation of single spicules or spicule tracts forming a crust, giving the surface a smooth
appearance; choanosomal skeleton more-or-less a regular isotropic reticulation of multispicular tracts,
without distinction between primary or secondary tracts, bound together with minimal spongin, forming
oval meshes; microscleres may include microxeas and microstrongyles; reproduction oviparous.
SCOPE: Five nominal genera are included in the family, only three of which are now recognised.
REVIEWS: Wiedenmayer (1977), van Soest (1980), Hartman (1982).
GENERA:
Petrosia Vosmaer, 1885 (type species: Petrosia ficiformis (Poiret, 1789)) (syn. Schmidtia
Balsamo-Crivelli, 1863; Tedaniella Czerniavsky; Acanthostrongylophora Hooper, 1984) - ectosomal
skeleton present as an isotropic spicule crust; choanosomal skeleton is basically lamellate-isotropic
reticulate composed of dense spicule tracts bound with very little spongin giving the sponge a stony
texture; oxeote or strongylote spicules in at least 2 size categories (modified from Bergquist & Warne,
1980; van Soest, 1980).
Strongylophora Dendy, 1905 (type species: Strongylophora durissima Dendy, 1905) lamellate-isotropic choanosomal reticulation of spicule tracts bound by spongin; spicules in at least 2
size categories, ranging from true oxeas, strongyloxeas to strongyles; microscleres kidney-shaped
microstrongyles (van Soest, 1980).
Xestospongia de Laubenfels, 1932 (type species: Xestospongia diprosopata (de Laubenfels,
1930)) (syn. Neopetrosia de Laubenfels, 1949; Prianos Gray, 1867) - ectosomal skeleton indistinct;
choanosomal skeleton confused isotropic reticulation of multispicular tracts, generally lacking spongin,
and sometimes with single spicules scattered throughout mesohyl between major spicule tracts; stony
texture; oxeote spicules in one size category only (Berquist & Warne, 1980; van Soest, 1980).
q FAMILY SPONGILLIDAE GRAY, 1867.
DEFINITION: Freshwater sponges producing asexual resting bodies (gemmules) with a well developed
vesicular coat of spongin, enabling floatation; growth forms generally encrusting, massive or cushionshaped; megascleres diactinal (oxeas or strongyles), smooth or spined, dispersed in regularly renieroid,
irregularly isodictyal or confused tracts, bound together by greater or lesser quantities of spongin fibres,
or only collagen; specialised microscleres (gemmoscleres) present and associated with gemmules and/or
occuring within mesohyl, consisting of smooth or spined microxeas, amphidiscs or birotules; sexually
produced larvae parenchymella have fully developed choanocyte chambers before extrusion from the
parent.
SCOPE: This family includes 24 nominal genera, of which 15 are considered to be valid.
REVIEWS: Penney & Racek (1968), Volkmer-Ribeiro & Ruetzler, 1997.
GENERA:
Anheteromeyenia Schrder, 1927 (type species: ).
Corvospongilla Annandale, 1911 (type species: ).
75
Dosilia Gray, 1867 (type species: ) (syn. Asteromeyenia Annandale, 1912; Astromeyenia
Schrder, 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 Schrder, 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).
q FAMILY METANIIDAE VOLKMER-RIBEIRO, 1986.
DEFINITION: Freshwater sponges producing large asexual resting bodies (gemmules) with well
developed vesicular coat of spongin; growth forms delicate encrusting, bulbous or tubular; megascleres
in two classes: larger entirely smooth, diactinal, structural spicules, typically stout oxeas, strongyloxeas
or strongyles, forming reticulate skeletal tracts, usually enclosed within well developed spongin fibres;
smaller are spined oxeas or strongyles surrounding the gemmule's vesicular coat; microscleres include
minute oxeas, with large central spines and granular spination elsewhere, and small isochelae-like
spicules; gemmoscleres include modified (boletiform) cladotylotes.
SCOPE: Seven nominal genera are included, of which four are presently recognised.
REVIEWS: Nil.
GENERA:
Acalle Gray, 1867 (type species: )
Corvomeyenia Weltner, 1913 (type species: ) (syn. Parameyenia Jewell, 1952).
Drulia Gray, 1867 (type species: ) (syn. Parmula Carter, 1881).
Metania Gray, 1867 (type species: Spongilla reticulata Bowerbank, 1863) (syn. Tubella
Carter, 1881).
q FAMILY POTAMOLEPIDAE BRIEN, 1967.
DEFINITION: Freshwater sponges that produce asexual resting bodies (gemmules) with an atypical
morphology - being larger and lacking the vesicular spongin coat typical of Spongillidae; growth forms
encrusting, often with irregular surfaces or branches; texture characteristically stony; megascleres
typically differentiated into choanosomal strongyles, forming reticulate tracts, with ectosomal oxeas in
periphery of skeleton; megascleres characteristically inflated at ends; spongin poorly developed or absent
from skeleton; microscleres are peculiar for the family; gemmoscleres spherical, often centrangulate
(rather than amphioxeote as in Spongillidae); sexually produced larvae parenchymella lacking well
developed choanocytes, but often have spined larval spicules
SCOPE: Four genera are now included here.
REVIEWS: Volkmer-Ribeiro & De Rosa-Barbosa (1979).
GENERA:
Potamolepis Marshall, 1883 (type species: ).
Potamophloios Brien, 1969 (type species: ).
Sterrastrolepis Volkmer-Ribeiro & De Rosa-Barbosa, 1978 (type species: ).
Uraguaya Carter, 1881 (type species: ).
q FAMILY LUBOMIRSKIIDAE BRIEN, 1969.
DEFINITION: Freshwater sponges lacking gemmules; growth forms encrusting, cushion-shaped or
branching; megascleres exclusively diactinal (oxeas or strongyles), entirely spined or only spined at
points, forming ascending multispicular tracts joined by uni- or paucispicular transverse connecting
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).
q FAMILY SPONGIIDAE GRAY, 1867.
DEFINITION: Encrusting, massive, cup-shaped and branching sponges, including the commercial `bathsponges'; surface typically conulose or with a distinct sand cortex; texture compressible, fibrous,
resilient except where heavily sand encrusted, and interior is rough to touch reflecting the density of
spongin skeleton in relation to soft tissue; choanosomal skeleton consists of reticulate spongin fibres,
usually organised into a heirarchy of sizes: the primary elements are reduced in some genera and
completely absent in one; fibres homogenous in cross section, showing no tendency to fracture around
planes of concentric lamination, lacking a central pith but often incorporating detritus and foreign
spicules into the spongin skeleton; choanocyte chambers small and diplodal.
SCOPE: Twenty nominal genera are presently included in the family, of which 11 are currently
recognised.
REVIEWS: de Laubenfels (1948), Vacelet (1959), van Soest (1978), Bergquist (1980, 1995),
Bergquist et al. (1988), Boury-Esnault et al. (1990: 237).
GENERA:
Carteriospongia Hyatt, 1877 (type species: Spongia foliascens Pallas, 1766) (syn.
Polyfibrospongia Bowerbank, 1877) - variable but lamellate, frondose, cup shpaed spreading habit
most common; surface always heavily sand-encrusted and thrown into a pattern of ridges or prominent
mounds with intervening grooves, characterising oscular and poral faces; oscules flush or elevated on
surface folds or ridges but not marked by any special ectosomal soft tissue location; skeleton extremely
irregular reticulation in qwhich primary elements are heavily cored and subsequently variable in shape
and diameter, with orientation not regular with respect to the surface but they can form marked fascicles
or brushes near the surface; distinguishable secondary elements are present and are either cored or clear;
these are markedly reduced as a skeletal element in favour of long thin vermiform tertiary fibres which
ramify without orientation fo the surface; these fibres branch, anastomose and interwine (Bergquist et
al., 1988).
Coscinoderma Carter, 1883 (type species: Spongia pesleonis Lamarck, 1814) - Flabellate,
pyriform, massive or pedunculate, with apical or marginal oscules; surface with sand armour, but texture
soft, spongy and extremely compressible; primary fibres cored and secondary elements clear, extremely
fine, numerous and intertwined ("whorls of wool") (Bergquist, 1980).
Dactylospongia Bergquist, 1965 (type species: Luffariella elegans Thiele, 1899) - Primary
fibres absent, and secondary fibres make up entire skeleton, relatively regular, with no foreign particles;
surface of fibres covered in refractive granules; sponge texture firm, hard and only slightly compressible
77
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).
Strepsichordaia Bergquist, Ayling and Wilkinson, 1988 (type species: Strepsichordaia

lendenfeldi Bergquist, Ayling & Wilkinson, 1988) - cup or fan shaped, always with macroscopically
smooth, sand reinforced surface marked by evenly dispersed small flush oscules, each surrounded by
prominent superficially running exhalant canals producing a stellate pattern over the otherwise
microconulose oscular surface; the poral surface is macroscopically smooth; a thick organised sand
cortex is present on both faces; skeleton is irregular with heavy cored primary fibres; simple not
fasciculate, but branching in irregular fashion; secondary elements are not distinct from primaries in
diameter or coring only in disposition; uncored cylindrical tertiary elements dominate the skeleton,
arising from both primary and secondary cored fibres and form a dense mat throughout the sponge; these
fibres meander for considerable distances without branching, have no fixed orientation with respect to
the surface or attachment base, do not form fascicles, and only occasionally interconnect; texture firm,
flexible, but not easily compressible (Bergquist et al., 1988).
q FAMILY IRCINIIDAE GRAY, 1867.
SYNONYM: Hirciniadae Gray, 1867; Hircinidae Lendenfeld, 1888.
DEFINITION: Massive, lobate, spherical, digitate, cup shaped, encrusting growth forms, always with a
conulose surface, except in forms with an organised superficial sand crust where conules may be
reduced to mammiform protruberances; fibres making up anastomosing skeleton laminated in cross
section with a central pith region, often obscured by large quantities of debris incorporated into fibres
and interstitially; skeleton irregularly arranged; primary fibres always fasciculate, often forming very
complex arrays; secondary fibres generally uncored; a third element consists of fine collagen filaments
dispersed in wavy tangled tracts throughout the mesohyl; filaments have terminal knobs, sometimes
studded with lepidocrocite granules, composed of collagen distinct from that found in the mesohyl
matrix or in the fibres; presence of filaments makes the sponge very tough, almost impossible to tear;
choanocyte chambers spherical and diplodal; mmesohyl only lightly infiltrated with collagen (like the
Spongiidae).
SCOPE: Twelve nominal genera are included in the family, 3 of which are valid.
REVIEWS: Lendenfeld (1888), Bergquist (1995), Cook & Bergquist (1996, 1998).
GENERA:
Ircinia Nardo, 1833 (type species: Spongia fasciculata Pallas, 1766) (syn. Dysidicinia
Lendenfeld, 1889; Euricinia Lendenfeld, 1889; Filifera Lieberkhn, 1859; Hircinella Lendenfeld,
1889; Hircinia Nardo, 1833; Polytherses Duchassaing & Michelotti, 1864; Stematumenia Bowerbank,
1845) - sponge surface marked by prominent conules, unarmoured; texture extremely tough, difficult to
cut or tear; primary fibres are cored and with detritus and frequently attain a great size by being woven
into complex fascicles; secondary fibres simple but not cored with detritus; collagenous mesohyl
typically heavy and charged with collagenous filaments bearing a terminal ampule (Bergquist, 1980;
Cook & Bergquist, 1996, 1998).
Psammocinia Lendenfeld, 1889 (type species: Ircinia halmiformis Lendenfeld, 1889) Texture firm, sometimes brittle; surface either pitted or covered with rounded tubercles, and thick sand
cortex present; large quantities of sand also incorporated into fibres and collagenous mesohyl; each sand
grain large, coated with collagen, and linked to the skeletal fibres by spongin strands; fibres only weakly
fasciculate and often obscured by the accumulation of sand; fine interstitial collagen filaments with
terminal ampule also present (Bergquist, 1980; Cook & Bergquist, 1996, 1998).
Sarcotragus Schmidt, 1862 (type species: Sarcotragus spinulosus Schmidt, 1862) (syn.
Dysidicinia Lendenfeld, 1889; Stenospongia Burton, 1928) - Surface conulose and texture tough;
fasciculate primary fibres lack or almost lack foreign inclusions, with characteristic pith clearly visible;
secondary fibres are clear of detritus and also contain pith; knobbed collagen filaments permeate the
matrix, but these are always very fine (Bergquist, 1980).
q FAMILY THORECTIDAE BERGQUIST, 1978.
DEFINITION: Sponge body often tubular organised around a series of long cylindrical canals, and
stalked; yellowish or brick-red internal pigmentation with dark exterior common; surface often
armoured in complex fashion, frequently thrown into ridges and hollows; where unarmoured the surface
is conulose and may resemble Spongiidae; spongin fibres making up the anastomosing skeleton are
laminated in cross-section, with clear zones of disjunction between successive layers; central region with
more diffuse pith, not sharply disjunct from the investing more dense layer (as is the pith in Verongida),
but merges into the outer layer; pith always evident in the primary fibres and may or may not extend into
the secondary elements of the skeleton; fibres often extremely regular with almost perfectly rectangular
meshes; some fibres extremely stout; primary fibres may be greatly reduced in number, absent in one
genus; choanocyte chambers spherical and diplodal.
79
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 Poljaeff, 1884) (syn. Luffaria, in
part, Poljaeff, 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
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.
q FAMILY DYSIDEIDAE GRAY, 1867.
SYNONYM: [Spongeliidae] Vosmaer [nomen oblitum].
DEFINITION: Encrusting, massive or branching growth forms, typically with conulose surface; surface
conulose developed to various degrees, often characteristically sculptured by tangential spongin fibres
cored by sand, giving it a delicate lace-like appearance; texture usually soft and compressible, sometimes
brittle due to interstitial detritus; choanosome a wide reticulation of spongin fibres, concentrically
stratified although to varying degrees; fibres laminated and cored by a central pith, but this may be
obscured by abundant detritus which is frequently incorporated into the spongin fibres; mesohyl contains
only light collagen; choanocyte chambers are large and eurypylous.
SCOPE: Fourteen nominal genera are presently included in this family, of which only three are valid.
REVIEWS: de Laubenfels (1948), van Soest (1978), Bergquist (1980, 1990, 1995), Boury-Esnault et
al. (1990: 237).
GENERA:
Dysidea Johnston, 1842 (type species: Dysidea fragilis Montagu, 1814) (syn. Aulena
Lendenfeld, 1885; Duseideia Johnston, 1842; Duseidea Delage, 1899; Dysidia Agassiz, 1846;
Dyseideia Lieberkhn, 1855; Haastia Lendenfeld, 1888; Halmopsis Lendenfeld, 1885; Sarcocornea
Carter, 1859; Spongelia Schmidt, 1862) - surface always conulose, with distinct sand-tracery (cobweblike) on the surface interconnecting adjacent conules; all fibres are concentrically stratified, filled with
pith, and filled with detritus, but collagen is only light in mesohyl (Bergquist, 1980).
Euryspongia Row, 1911 (type species: Euryspongia lactea Row, 1911) - Surface usually
marked by a delicate tissue network between pore areas; primary fibres forming regular reticulate
81
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).
q FAMILY DARWINELLIDAE MEREJKOWSKY, 1879.
SYNONYM: Aplysillidae Lendenfeld.
DEFINITION: Encrusting, massive, lobate, lamellate and erect columnar growth forms; choanosomal
fibre skeleton, where present, is completely dendritic and sometimes supplemented by spongin spicules
not attached to the primary skeleton; in massive species these fibres always arise from a flat basal
spongin plate; one genus lacks spongin fibres but has the ectosome reinforced with collagenous fibrils;
fibres have laminated bark surrounding the central pith; the pith is usually well developed but in 1 genus
it is replaced by detritus; choanocyte chambers are eurypylous.
SCOPE: Eight nominal genera and six probably valid genera are included in the family.
REVIEWS: Wilson (1925), de Laubenfels (1948), van Soest (1978), Bergquist (1980, 1995).
GENERA:
Aplysilla Schulze, 1878 (type species: Aplysilla rosea Barrois, 1876) (syn. Simplicella
Merejkowsky, 1878) - Always encrusting growth form; often bright yellows, reds, pink colours, with
colour changes; numerous short, almost unbranched fibres; fibres always clear of detritus (Bergquist,
1980).
Chelonaplysilla de Laubenfels, 1948 (type species: Aplysina noevus Carter, 1876) - separable
ectosomal (cortical) skeleton with a delicate reticulation of sand grains; fibre structure is similar to
Aplysilla but in some erect and branching species the supporting skeleton becomes more branched
(Bergquist, 1980).
Darwinella Mller, 1865 (type species: Darwinella mulleri Schulze, 1865) (syn. Darwinia
Schulze, 1865) - Small, encrusting, massive or lobate sponges; slightly ramified dendritic fibre skeleton
is supplemented by diactinal, triactinal or quadriradiate spongin spicules; fibres lack any detritus
(Bergquist, 1980).
Dendrilla Lendenfeld, 1883 (type species: Dendrilla rosea Lendenfeld, 1883) - Large erect
branching or complex lamellate growth forms in which the skeleton is dendritic, branching repeatedly
but not anastomosing; fibres contain no foreign coring material (Bergquist, 1980).
Hexadella Topsent, 1896 (type species: Hexadella racovitzae Topsent, 1896) - Encrusting
growth forms; no fibres or spongin spicules; ectosome is collagen reinforced to form a noticeable skin
which, acting in some measure as an external skeleton, allowing the sponge to attain a thickness up to
5mm (Bergquist, 1980).
Pleraplysilla Topsent, 1905 (type species: Spongelia spinifera Schulze, 1878) - with simple
fibres, sometimes branched only once or twice but in thicker specimens branching more frequently; all
fibres arenaceous, with an axial core of sand grains, with constant diameter from base to surface of the
sponge even though the stratified bark elements diminish (Bergquist, 1980).
q FAMILY DICTYODENDRILLIDAE BERGQUIST, 1980.
DEFINITION: Growth form varies from spreading with digitate projections, to lobate or stalked forms;
texture is delicate and cavernous; choanosomal skeleton has prominently reticulate spongin fibres which
may be augmented by incorporation of free spongin spicules in one genus; fibres often dark purple, red,
or black, and contrasts with the soft tissue which is either pale or densely and uniformly pigmented; fibre
structure is heavy, concentrically laminated and pithed, but pith may be obscured by the incorporation of
detritus into fibres; choanocyte chambers are large, eurypylous.
SCOPE: Three valid genera (four named genera) are included in this family.
REVIEWS: Bergquist (1980, 1995).
GENERA:
Acanthodendrilla Bergquist, 1995 (type species: Acanthodendrilla australis Bergquist, 1995)
- thickly flabellate; surface covered with sharp conules with fibres projecting above surface; ectosomal
skeleton thin collagen layer with cavernous lacunae below ectosome; choanosomal skeleton cavernous
with large volume devoted to canals, fibres forming irregular reticulate meshes, large coarse fibres,
fibres strongly laminate cored with sand and spicule debris, but central pith clearly evident; mesohyl
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 FAMILY HALISARCIDAE VOSMAER, 1885.
DEFINITION: Thin growth form, soft texture, gelatinous surface; fibrous skeleton absent entirely;
choanocyte chambers consist of specialised wide-mouthed, extended tubular and branched eurypylous
forms; parenchymella larvae are simple, lacking long terminal cilia; (often superficially confused with
didemnid ascidians).
DEFINITION: Thin growth form, soft texture and gelatinous surface characterise this family. The
fibrous skeleton is absent entirely. Choanocyte chambers consist of specialised wide-mouthed, extended
tubular and branched eurypylous forms. Parenchymella larvae are simple, lacking long terminal cilia.
SCOPE: One genus is presently included.
REVIEWS: Lvi (1952, 1973), van Soest (1978), Bergquist (1980), Boury-Esnault et al. (1990).
GENERA:
Halisarca Dujardin, 1838 (type species: Halisarca dujardini Johnston, 1842) - fibrous
skeleton absent entirely (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.
q FAMILY APLYSINIDAE CARTER, 1875.
SYNONYM: [Verongiidae] de Laubenfels.
DEFINITION: Encrusting, massive, club-shaped and fan-shaped growth forms; reticulate, anastomosing
spongin fibres produce polygonal meshes, not organised into one plane; fibres have normal bark and pith
elements, without foreign detritus, and the collagenous spongin matrix is dense; choanocyte chambers
small, spherical and diplodal.
SCOPE: Six nominal and two valid genera of aplysinid sponges are presently included.
REVIEWS: Wilson (1925), van Soest (1978), Bergquist (1980).
GENERA:
Aplysina Nardo, 1834 (type species: Aplysina aerophoba Schmidt, 1862) (syn. Aplysia, sensu
Nardo, 1833; Fistularia, in part, Bowerbank, 1844; Verongia Bowerbank, 1845; Luffaria Duchassaing
& Michelotti, 1864) - Marked aerophobic colour change from yellow or green to darker colours; only
one kind of fibre is present, without foreign detritus and with a thick pith component; fibres form a
regular reticulation with large hexagonal meshes and have no specialised surface arrangement
(Bergquist, 1980).
Verongula Verrill, 1907 (type species: Aplysina praetexta Hyatt, 1875) - skeletal
characteristics identical to those of Aplysina except for the elaboration of a radially prismatic
architecture which is particularly evident near the surface, which produces a honeycomb-like appearance
to the surface (Bergquist, 1980).
83
q FAMILY DRUINELLIDAE LENDENFELD, 1889.

SYNONYM: Aplysinellidae Bergquist.
DEFINITION: Lobate and club-shaped sponges; pigmentation frequently sulphur yellow that usually
oxidizes to purple, although some species have superficial pink to purple coloration and a beige to pale
yellow interior; skeleton with dendritic fibres, widely spaced or greatly reduced in relation to the heavy
collagenous spongin matrix, sometimes supplemented by spongin spicules; fibres with strong bark
elements, and fibres have the pith component emphasised over the bark, which is typically reduced or
absent; choanocyte chambers small, spherical and diplodal.
SCOPE: Seven nominal genera have been included in the family, three of which are thought to be valid.
REVIEWS: Bergquist (1980, 1995).
GENERA:
Aplysinella Bergquist, 1980 (type species: Aplysinella strongylata Bergquist, 1980) differential pigmentation in life between surface and interior; fibres of characteristic construction having
both bark and pith components, the latter dominant, and fibre skeleton augmented by fibrous spicules
with strongylote ends; the spicules are composed only of bark and have a clear axial canal (Bergquist,
1980).
Druinella Lendenfeld, 1889 (type species: Psammaplysilla arabica Keller, 1889) (syn.
Psammaplysilla Keller, 1889; Korotnewia Poljaeff, 1889) - Encrusting to complex-ramose, not
massive growth form; texture of sponge compressible and rather rubbery; surface markedly conulose;
skeleton has basic dendritic pattern but also has frequent fasciculation; fibres present in moderate
quantities in relation to the matrix (not sparse or isolated as in Pseudoceratina); pith is present to the
exclusion of bark in fibres, and occasional sand or spicule fragments (Bergquist, 1980).
Porphyria Bergquist, 1995 (type species: Porphyria flintae Bergquist, 1995) - stalked goblet
shaped, thick lamellae, with distinct internal oscular and external ostial faces; interior surface smooth,
exterior roughened by low conules and contoured; ectosomal skeleton well marked with little collagen
but many spherulous cells; choanosomal skeleton made up of abundant, slender, even, branched
(dendritic) fibres branching from central basal point, tapering to sharp points near surface; branched
fibres short, flexuous, sharply pointed; fibres with reduced pith occupying only 25-33% of fibre
diameter, and surrounding bark very dense and tightly laminated producing brittle fibres; mesohyl
strongly collagen reinforced, choanocyte chambers small, spherical, diplodal (Bergquist, 1995).
Pseudoceratina Carter, 1885 (type species: Pseudoceratina durissima Carter, 1885) (syn.
Dendrospongia, sensu Hyatt, 1875; Aiolochroia Wiedenmayer, 1977) - texture firm, often hard and
incompressible; smooth surface of the sponge is interrupted by widely spaced tubercles rather than
conules, or by low raised ridges; sparse fibre skeleton organised on the dendritic plan typical of the
family; pith elements present in the fibres, sometimes to the exclusion of the bark, which is evident only
as occasional peripheral patches; matrix extremely dense and with heavy collagen (Bergquist, 1980).
Suberea Bergquist, 1995 (type species: Suberea creba Bergquist, 1995) - massive, spreading,
irregular or branching shapes; scattered contractile raised oscules; surface smooth but thrown into lumps
and folds appearing tuberculate; ectosomal skeleton strongly collagenous, sharply set off from
choanosome, with abundant spherulous cells and thin external cuticle; choanosomal skeleton sparse,
made up of coarse, thick, irregular, dendritic fibres, circular in cross-section, composed of both bark and
pith elements with pith predominating and accounting for about 75% of fibre diameter; fibre bark
strongly laminated very brittle; texture of sponge interior very rough; mesohyl evenly and densely
collagenous making the sponge barely compressible, choanocyte chambers small, spherical (Bergquist,
1995).
q FAMILY IANTHELLIDAE HYATT, 1875.
SYNONYM: Bajalidae Lvi.
DEFINITION: Lobate and fan-shaped, stalked growth forms common; pigmentation ranges from
typically sulphur yellow, deep orange to deep purple with oxidation to deep purple; fibre skeleton, where
present, is reticulate and frequently compressed into 2 dimensions, radiating from the contracted basal
attachment; fibres typically large, particularly towards the base of the sponge, containing cellular
elements in distinctive concentric annuli occurring mainly in the bark component of each fibre;
choanocyte chambers large and eurypylous, sac-shaped, varying between genera from simply elongate to
occasionally branched.
SCOPE: Five nominal and three valid genera are recognised for the family.
REVIEWS: Bergquist (1980).
GENERA:
Anomoianthella Bergquist, 1980 (type species: Anomoianthella popeae Bergquist, 1980) thickened fan-shaped, club-shaped or cup-shaped growth forms, always with a discrete oscular and ostial
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 DEMOSPONGIAE INCERTAE SEDIS
MyceliospongiaVacelet & Perez, 1997 (type species Myceliospongia araneosa Vacelet &
Perez, 1997) q CLASS CALCAREA (Calcareous Sponges).
DEFINITION: With exclusively calcitic spicules ranging from discrete monactinal, diactinal, triactinal
or tetractinal spicules, to reticulate skeletons composed of fused crystalline calcite spicules; megascleres
and microscleres are not differentiated; skeleton and aquiferous system occurs in three grades of
construction: (1) asconoid, with simple tubular construction (olynthus), without folding of the body
wall, with thin walls pierced externally by ostia, leading to tubular water canals (porocyte canals)
opening onto a central choanocyte-line cavity (choanoderm), connected to the exterior, at the apex of the
sponge, by a single osculum; (2) syconoid construction produced by folding of both the exterior
(pinacoderm) and interior (choanoderm) walls, producing choanocyte chambers to lie within the body
wall rather than only lining the central atrium as in more simple asconoid structures, but these chambers
open directly onto the atrium; (3) leuconoid, found in most sponges (including the Demospongiae), with
complex folding and in which the choanocyte chambers are oval and isolated in a maze of canals within
the body wall, with chambers opening onto branching and complex excurrent canals; sexual reproduction
in Calcarea is exclusively viviparous.
Two subclasses are now recognised.
REVIEWS: Dendy & Row (1913), Hartman (1958), Burton (1963), Borojevic (1966, 1968, 1977),
Vacelet (1970, 1981, 1991), Borojevic & Boury-Esnault (1986), Borojevic, Boury-Esnault & Vacelet
(1990).
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.
q FAMILY CLATHRINIDAE MINCHIN, 1900.
DEFINITION: Essentially tubular organisation, with continuous choanoderm lining all internal cavities;
growth is by longitudinal median divisions and anastomosis of tubes to form large units called the
cormus; neither a common cortex nor a well-defined inhalant and exhalant aquiferous system.
SCOPE: Two genera (of five named genera) are included in the family.
REVIEWS: Bidder (1898), Minchin (1909), Dendy (1913), Hartman (1958), Borojevic (1968),
Borojevic, Boury-Esnault & Vacelet (1990).
GENERA:
85
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).
q FAMILY SOLENISCIDAE BOROJEVIC, BOURY-ESNAULT & VACELET, 1990.
DEFINITION: Essentially tubular organisation; a continuous choanoderm lines all internal cavities;
spicules are regular triradiates and/or quadriradiates, sometimes with tripods or biradiates; sponges grow
in the form of an individual olynthus, with several olynthi growing from the basal stolon-like tubes, or
in the form of distally branching tubes radially arranged around a central olynthus tube without any
special skeletal differentiation.
SCOPE: Two genera are included.
REVIEWS: Nil.
GENERA:
Soleniscus Haeckel, 1870 (type species: Leucosolenia stolonifer Dendy, 1891) - form of an
individual erect olynthus, with several olynthi growing from basal stolon-like tubes (tubular
spongorhiza), or in the form of distally ramified creeping tubes (Borojevic et al., 1990); surface hispid;
apical oscules, firm texture; skeleton of regular to slightly sagittal quadriradiates and oxeas (Burton,
1963).
Dendya Bidder, 1898 (type species: Clathrina tripodifera Carter, 1886) - Cormus solitary,
conical to ovoid, consisting of a large central tube from which smaller radially arranged tubes branch
off; surface even, reticulate; oscules apical; texture firm; skeleton of regular and tripodal triradiates and
sagittal quadriradiates (Burton, 1963; Borojevic et al., 1990).
q FAMILY LEVINELLIDAE BOROJEVIC & BOURY-ESNAULT, 1986.
DEFINITION: Cormus composed of a central tube, sometimes ramified, and diverticuli isolated or
grouped in clusters; skeleton of central and radial tubes composed of regular equiradiate and equiangular
spicules; skeleton of diverticuli composed of regular and/or parasagittal spicules always clearly distinct
from spicules composing the skeleton of the central tube; choanoderm either lines all the central cavity
or is restricted to the diverticuli.
SCOPE: Two genera are included.
REVIEWS: Borojevic et al. (1990).
GENERA:
Levinella Borojevic & Boury-Esnault, 1986 (type species: Levinella thalassae Borojevic &
Boury-Esnault, 1986) - with a cormus divided into a central tube and external diverticuli; central tube
not ramified; choanoderm lines all internal cavities (Borojevic et al., 1990); spicules are equiangular
tetractines in the central tube organised in parallel rows, equiangular and equiradiate tetractines and less
common triactines of the diverticula, and slightly curved diactines (Borojevic & Boury-Esnault, 1986).
Burtonulla Borojevic & Boury-Esnault, 1986 (type species: Burtonulla sibogae Borojevic &
Boury-Esnault, 1986) - with central tube ramified; choanoderm limited to external diverticuli
(Borojevic et al., 1990); spicules are equiangular and equiradiate triactines and tetractines with short
basal actine in the central tube and lateral walls, equiangular and equiradiate triactines and tetractines
with rudimentary apical actine lining the diverticula (Borojevic & Boury-Esnault, 1986).
q FAMILY LEUCALTIDAE DENDY & ROW, 1913.
DEFINITION: Tubular, branching or regularly anastomosing cormus, either with many oscules or with a
large atrium and a single osculum; sponge wall composed of a distinct cortex and choanosome; skeleton
of choanosome and atrial wall absent or composed of small and dispersed triradiates and quadriradiates.
SCOPE: There are four named genera (three valid), and a fourth incertae sedis.
REVIEWS: Hartman (1958), Burton (1963), Borojevic (1968), 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 Poljaeff, 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).
q FAMILY LEUCASCIDAE DENDY, 1893.
DEFINITION: Body differentiated into cortex and choanosome reminiscent of a clathroid body
composed of anastomosed tubes; cortex composed of large triradiates and/or quadriradiate spicules;
choanocyte chambers tubular, often highly ramified and anastomosed; choanoskeleton restricted to walls
of the choanocyte chambers, maintaining a distinctly tubular organisation.
SCOPE: The four nominal and two valid genera included in this family.
REVIEWS: Dendy (1893), Bidder (1898), Dendy & Row (1913), Burton (1963), Borojevic (1968),
Borojevic et al. (1990).
GENERA:
Aulorrhiza Haeckel, 1869 (type species: Leucosolenia lamarckii Haeckel, 1869, sen. syn. of
Aulorrhiza intestinalis Haeckel, 1869) (syn. Ascoleucetta Dendy & Frederick, 1924; Ascaltis, sensu
Borojevic, 1968; Borojevic et al., 1990) - with a massive cormus composed of ramified and
anastomosed tubes covered by a common cortex; the inhalant aquiferous system is represented by spaces
delimited by the cortex and the walls of choanosomal tubes; the exhalant aquiferous system is reduced to
the osculum or to a secondary atrial cavity formed by the calyciform growth of the cormus (Borojevic et
al., 1990); rounded mass of anastomosing tubes; skeleton of equiradiate triradiates and quadriradiates
with regular facial rays (Burton, 1963).
Leucascus Dendy, 1893 (type species: Leucascus simplex Dendy, 1892) - with copiously
branched and anastomosed choanocyte tubes; exhalant aquiferous system is represented by a welldeveloped atrium delinited by a specific wall within the choanoderm (Borojevic et al., 1990); solitary,
irregularly massive to spherical; surface even, smooth, small scattered oscules, firm texure; skeleton of
regular triradiates or occasionally quadriradidiates with incipient apical rays (Burton, 1963).
q FAMILY LEUCETTIDAE DE LAUBENFELS, 1936.
DEFINITION: Solid body; aquiferous system always leuconoid; choanoskeleton well-developed
forming regular network composed of triradiates and/or quadriradiates; cortex thin and composed of
spicules similar to those in the choanoskeleton.
SCOPE: The family includes six generic names, of which two are valid genera and one incertae sedis.
REVIEWS: Borojevic (1968), Borojevic et al. (1990).
GENERA:
87
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 Poljaeff, 1884 (type species: Pericharax heterorhaphis Poljaeff, 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..
q FAMILY MURRAYONIDAE KIRKPATRICK, 1910.
DEFINITION: Choanosomal skeleton composed of a rigid calcareous aspicular network; cortex
composed chiefly of overlapping calcareous scales, with tuning-fork spicules below.
SCOPE: Single genus.
REVIEWS: Dendy & Row (1913), Hartman (1958), Vacelet (1970), Brien et al. (1973), Hartman
(1980), Vacelet (1981), Borojevic et al. (1990).
GENERA:
Murrayona Kirkpatrick, 1910 (type species: Murrayona phanolepis Kirkpatrick, 1910) - with
a definite pore-zone whose ectosomal (cortical) skeleton consists of small triradiates; diapasons (tuningfork shaped spicules) are present beneath the ectosomal scales (Burton, 1963; Borojevic et al., 1990);
subspherical to pyriform; surface even, imbricated; oscules small, lateral, pores in equatorial groove;
texture hard; ectosomal skeleton a layer of subcircular scales, with an ectosomal layer of irregular
triradiates and irregular tuning-fork spicules, regular triradiates of the poral groove, and a main skeleton
composed of a stout reticulation of calcareous fibres (Burton, 1963).
q FAMILY PARAMURRAYONIDAE VACELET, 1967.
DEFINITION: Choanosomal skeleton composed of bundles of diapsons (tuning-fork triradiates)
without any rigid structure; cortical skeleton composed chiefly of a superficial layer of overlapping
calcareous scales and an internal layer of free calcareous plates.
REVIEWS: Borojevic et al. (1990).
GENERA:
Paramurrayona Vacelet, 1967 (type species: Paramurrayona corticata Vacelet, 1967) - as
for family.
q FAMILY LELAPIELLIDAE BOROJEVIC, BOURY-ESNAULT & VACELET, 1990.
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).
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.
q FAMILY LEUCOSOLENIIDAE MINCHIN, 1898.
DEFINITION: Asconoid, erect growth forms, with long, individual, clustered, oscular tubes arising
from stolon-like system of basal tubes; tubes may have diverticular and often arborescent; monaxon
spicules always present; triradiates, if present, typically bilateral, sagittal, inequiangular in form (where
two of the rays form a pair, while the third differs in some way), and with the crystalline optic axis never
vertical but always inclined to the facial plane of the rays; choanocytes with flagellum arising directly
from the pear-shaped nucleus, situated at or near the apex of the cell; choanocytes line central cavity
(spongocoele) of the individual tubes; larvae are amphiblastula.
SCOPE: About 90 generic names are included in this family, but only three of these are now recognised.
REVIEWS: Bidder (1898), Minchin (1909), Dendy & Row (1913), Hartman (1958), Burton (1963).
GENERA:
Ascute Dendy & Row, 1913 (type species: Leucosolenia uteoides Dendy, 1893) - compound,
tubular, sessile, surface even, minutely hispid, striated; oscules apical; firm texture; ectosomal skeleton
has large diacts (oxeotes), of one or two sizes, echinating surface and/or lying longitudinally within the
ectosome; choanosomal skeleton composed of sagittal quadriradiates (Burton, 1963).
Ascyssa; Haeckel, 1872 (type species: Ascyssa acufera Haeckel, 1872) - tubular, colonial;
smooth surface; apical oscules; skeleton a tangential layer of irregularly arranged oxeote and microxeote
spicules (Burton, 1963).
Leucosolenia Bowerbank, 1861 (type species: Spongia botryoides Ellis & Solander, 1786)
(syn. Nardosa Wright, 1868; Olynthium Haeckel, 1870; Olynthus Haeckel, 1870; Prosycum Haeckel,
1870; Leucaria Haeckel, 1870; Leucelia Haeckel, 1870; Leuceria Haeckel, 1870; Leucilia Haeckel,
1870; Leuciria Haeckel, 1870; Leucoria Haeckel, 1870; Tarroma Haeckel, 1870; Tarrus Haeckel,
1870; Nardopsis Haeckel, 1870; Clystolynthus Haeckel, 1870; Sycorrhiza Haeckel, 1870; Auloplegma
Haeckel, 1870; Thecometra Haeckel, 1870; Ascettaga Haeckel, 1872; Ascettopa Haeckel, 1872;
Ascettusa Haeckel, 1872; Ascometra Haeckel, 1872; Nardorus Haeckel, 1872; Solenula Haeckel, 1872;
Tarropsis Haeckel, 1872; Ascilla Haeckel, 1872; Ascillaga Haeckel, 1872; Ascillopa Haeckel, 1872;
Ascaltaga Haeckel, 1872; Ascaltopa Haeckel, 1872; Ascuris Haeckel, 1872; Ascortaga Haeckel, 1872;
Ascortopa Haeckel, 1872; Ascortusa Haeckel, 1872; Asculmis Haeckel, 1872; Ascandraga Haeckel,
1872; Ascandropa Haeckel, 1872; Olynthella Haeckel, 1872; Nardoma Haeckel, 1872; Solenidium
Haeckel, 1872; Olynthaltus Haeckel, 1872; Olynthandrus Haeckel, 1872; Olynthettus Haeckel, 1872;
Olynthortus Haeckel, 1872; Olynthillus Haeckel, 1872; Olynthulmus Haeckel, 1872; Olynthandrium
Haeckel, 1872; Olynthandra Haeckel, 1872; Olynthelletta Haeckel, 1872; Clistolynthus Haeckel,
1872; Clistolynthaltis Haeckel, 1872; Clistolynthandra Haeckel, 1872; Clistolynthetta Haeckel, 1872;
Clistolynthilla Haeckel, 1872; Soleniscetta Haeckel, 1872; Soleniscilla Haeckel, 1872; Soleniscyssa
Haeckel, 1872; Solenicaltis Haeckel, 1872; Solenicortis Haeckel, 1872; Soleniculmis Haeckel, 1872;
Soleniscandra Haeckel, 1872; Solenidandra Haeckel, 1872; Solenulandra Haeckel, 1872; Solenuletta
89
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).
q FAMILY SYCETTIDAE DENDY, 1892.
SYNONYM: [Syconidae] Haeckel [nomen oblitum].
DEFINITION: Tubular, spherical, flask-shaped, ovoid and branching growth forms, either solitary or in
groups; ectosomal cortex is continous and strengthened by tangential spicules, but these do not cover the
choanocyte chamber layer; choanosomal spicules, supporting choanocyte chambers, have an articulated
arrangement of overlapping sagittal triradiates, with the angle between the paired rays larger than the
angles between each paired ray and the long, unpaired ray; sagittal triradiates have the longest ray
pointing to the exterior of the sponge, and form a layer beneath the spongocoele lining (referred to as the
subgastral position); choanocytes usually confined to the radial chambers in the adult, and probably
always with apical nuclei; choanocyte chambers asconoid, arranged radially around a central cavity
(spongocoele), with ends of chambers projecting into ectosomal surface.
SCOPE: There are only four valid genera included in this family although there are approximately 60
other generic names included in synonymy with these.
REVIEWS: Dendy (1893), Dendy & Row (1913), Tanita (1943), Hartman (1958), Burton (1963),
Borojevic (1967).
GENERA:
Sycandra Haeckel, 1870 (type species: Sycandra utriculus Haeckel, 1872) - with an atrial
cavity partially obstructed by a column sustained by parallel diactines (Borojevic, Boury-Esnault &
Vacelet, in prep.).
Sycetta Haeckel, 1872 (type species: Sycetta primitiva Haeckel, 1872) (syn. Sycyssarium
Haeckel, 1872; Sycissa Lendenfeld, 1886) - tubular, stipitate; surface minutely papillate, non-hispid;
oscule simple, apical; texture soft; tubar and choanosomal skeletons composed of regular triradiates;
species lack tufts of oxeas at the distal ends of the radial chambers, and the radial chambers are always
completely separate from one another, otherwise similar to Sycon (Burton, 1963).
Sycodorus Haeckel, 1872 (type species: Sycandra (Sycodorus) hystrix Haeckel, 1872) (syn.
Sycodorussa Haeckel, 1892; Utella Dendy, 1892) - tubular, sessile; even surface prominently hispid;
apical oscule with fringed margin; firm texture; ectosomal skeleton of several tangential layers of
regular triradiates, with oxeas of two sizes projecting beyond the surface; tubar skeleton with several
rows of sagittal triradiates and sagittal quadriradiates and basal rays of subgastral sagittal triradiates and
sagittal quadriradiates; choanosomal skeleton of paired rays of subgastral sagittal triradiates, with
longitudinal oxeas and a tangential layer of sagittal triradiates and sagittal quadriradiates (Burton, 1963).
Sycon Risso, 1826 (type species: Spongia ciliata Fabricius, 1780) (syn. Scypha Gray, 1821;
Calcispongia Blainville, 1830; Calcepongia Lamarck, 1836; Dunstervillia Bowerbank, 1845;
Coniasterium Ehrenberg, 1861; Artynes Gray, 1867; Syconella Schmidt, 1868; Sycum Haeckel, 1870;
Artynella Haeckel, 1870; Sycarium Haeckel, 1870; Artynas Haeckel, 1870; Sycodendrum Haeckel,
1870; Sycidium Haeckel, 1870; Artynium Haeckel, 1870; Sycocystis Haeckel, 1870; Artynophyllum
Haeckel, 1870; Sycophyllum Haeckel, 1870; Sycometra Haeckel, 1870; Sycettopa Haeckel, 1870;
Sycettaga Haeckel, 1870; Sycurus Haeckel, 1870; Sycaltaga Haeckel, 1870; Sycuraltis Haeckel, 1870;
Sycocarpus Haeckel, 1870; Sycocerus Haeckel, 1870; Sycocubus Haeckel, 1870; Sycostrobus Haeckel,
1870; Sycodorilla Haeckel, 1870; Sycodoranna Haeckel, 1870; Leukartea Miklucho-Maclay in
Haeckel, 1872; Leucandropa Haeckel, 1872; Sycuretta Haeckel, 1872; Sycuranda Haeckel, 1872;
Sycurortis Haeckel, 1872; Syconellortis Haeckel, 1872; Sycortarium Haeckel, 1872; Sycandrarium
Haeckel, 1872; Syconellandra Haeckel, 1872; Syconcystandra Haeckel, 1872; Sycocystortis Haeckel,
1872; Sycothamnandra Haeckel, 1872; Sycodenandrum Haeckel, 1872; Sycinulandra Haeckel, 1872;
Sycandrophyllum Haeckel, 1872; Leucandrometra Haeckel, 1872; Sycandrometra Haeckel, 1872;
Sphenophorus Breitfuss, 1898; Sphenophorina Breitfuss, 1898; Hozawaia de Laubenfels, 1936;
Homoderma Lendenfeld, 1885; Sycantha Lendenfeld, 1891; Hypodictyon Jenkin, 1908; Streptoconus
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).
q FAMILY HETEROPIIDAE DENDY, 1893.
SYNONYM: [Djeddeidae] de Laubenfels (nomen vanum).
DEFINITION: Massive, tubular, pear-shaped and branching growth forms, occuring as solitary sponges
or in groups; continous cortex, pierced by ostia and reinforced by asymmetrical triradiate spicules with
unequal angles, covers the entirely choanocyte chamber layer; inarticulated or articulated tubular
skeleton characterised by a distinct subcortical zone formed by pseudosagittal triactines, but articulated
choanosomal spicule skeleton may be present or absent; cortical triradiate spicules probably originate
from articulate chamber skeleton, through reorientation of the spicules, so that one of the paired rays
becomes the sagittal ray and the latter pairs up with the remaining ray; choanocyte chambers asconoid,
elongate and radially arranged, or spherical and irregularly scattered in the choanosome (leuconoid).
SCOPE: Six genera are included in this family (of 8 generic names).
REVIEWS: Dendy (1893), Dendy & Row (1913), Hartman (1958), Burton (1963), Borojevic (1966),
Borojevic, R. (1967).
GENERA:
Sycettusa Haeckel, 1872 (type species: Leucaltis bathybia Haeckel, 1872) (syn. Djeddea
Miklucho-Maclay in Haeckel, 1872; Sycothamnetta Haeckel, 1872) - tubular, substipitate; surface even,
minutely hispid; apical oscule; firm texture; ectosomal skeleton a distinct tangential layer of sagittal
triradiates, with oxeas projecting at right angles to surface; skeleton of chamber layer composed of
centrifugally directed rays of sagittal triradiates and centipetally directed rays of subgastral
pseudosagittal triradiates, supplemented or partially replaced by confused triradiates; choanosomal
skeleton a tangential layer of sagittal tri- or quadriradiates (Burton, 1963).
Grantessa Lendenfeld, 1885 (type species: Grantessa sacca Lendenfeld, 1885) - sac shaped or
cylindrical; surface even but bearing isolated tufts of oxeas; texture firm; ectosomal skeleton a sparse
layer of tangential triradiates with unequal rays, paired rays of subectosomal pseudosagittal triradiates,
and tufts of oxeas projecting from the surface; tubar skeleton of basal rays of subectosomal
pseudosagittal and subgastral sagittal triradiates with numerous intermediate rows of subregular
triradiates; choanosomal skeleton of paired rays of subgastral sagittal triradiates and several layers of
choanosomal quadriradiates (Burton, 1963).
Grantilla Dendy & Row, 1913 (type species: Grantilla quadriradiata Row, 1909) - with
syconoid organisation; the skeleton of the tubes are inarticulate and composed of subatrial triactines and
subcortical pseudosagittal triactines and tetractines (Borojevic, Boury-Esnault & Vacelet, in prep.).
Heteropia Carter, 1886 (type species: Aphroceras ramosa Carter, in Higgin, 1886) - ramose,
sessile; surface even, non-hispid; oscules apical, terminal; firm texture; ectosomal skeleton with a
tangential layer of small sagittal triradiates and large ("colossal") longitudinally-placed oxeas; skeleton
of chamber layer of sagittal triradiates; choanosomal skeleton a tangential layer of small triradiates and
larger sagittal quadriradiates, the latter near the gastral cavity and oscule (Burton, 1963).
Paraheteropia Borojevic, 1966 (type species: Paraheteropia ijimai (Hozawa, 1916) - with
syconoid organisation and with cortical and atrial skeleton sustained by longitudinal diactines
(Borojevic, Boury-Esnault & Vacelet, in prep.).
Vosmaeropsis Dendy, 1893 (type species: Heteropia macera Carter, 1886) - tubular to
sacciform, compound; even hispid or non-hispid surface; apical oscules; firm texture; ectosomal
skeleton of several layers of triradiates together with paired rays of subectosomal pseudosagittal
triradiates and microxeas, with or without oxeas projecting at the surface; skeleton of the chamber layer
of basal rays of subgastral and subchoanosomal triradiates; choanosomal skeleton of paired rays of
sugastral triradiates and several layers of triradiates (Burton, 1963).
q FAMILY GRANTIIDAE DENDY, 1892.
SYNONYM: Leuconiinae Haeckel.
DEFINITION: Encrusting, lobate, tubular, sac-shaped, ovoid, spherical, vase-shaped and many other
growth forms, either solitary or grouped and sessile, substipitate, or stipitate; surface with a distinct
dermal cortex and a proper cortical skeleton of tangential radiates, sometimes supplemented by, and
91
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 Poljaeff, 1884 (type species: Anamixilla torresi Poljaeff, 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 Poljaeff, 1884 (type species: Eilhardia schulzei Poljaeff, 1884) - solitary cupshaped, 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).
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
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;
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 Poljaeff, 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).
q FAMILY AMPHORISCIDAE DENDY, 1892.
DEFINITION: Massive, tubular, ovoid and spherical growth forms, grouped together, never solitary
(one genus (Syculmis) has a root-like tuft of oxeas and anchoring quadriradiates); ectosomal cortex is
distinct and supported by tangentially placed radiates, with or without oxeas; ectosomal radiates may
have the large arm directed inwards, forming the main part of the choanosomal skeleton; no articulated
choanosomal skeleton present, but leuconoid forms may have quadriradiates scattered in the
choanosome and large quadri- or triradiates below the atrium (subgastral spicules); nuclei of
choanocytes probably always apical; choanocyte chambers asconoid, elongate and radially arranged, or
small, spherical and irregularly scattered in the choanosome (leuconoid).
SCOPE: Nineteen generic names, of which only seven are valid, are currently included in this family.
REVIEWS: Dendy (1893a, 1893c), Dendy & Row (1913), Hartman (1958)
GENERA:
Amphoriscus Haeckel, 1869 (type species: Ute chrysalis Schmidt, 1864) (syn. Sycilla Haeckel,
1872; Leuculmis Haeckel, 1872; Dyssycillus Haeckel, 1872; Sycurilla Haeckel, 1872; Dyssyculmus
Haeckel, 1872; Syculmarium Haeckel, 1872; Lipostomella Haeckel, 1872; Leoculmis Ganin, 1879;
Rhabdodermella Urban, 1902; Lenculmis Allemand, 1907) - solitary, tubular, stipitate; smooth surface;
terminal oscule; ectosomal skeleton of tangentially arranged facial rays of large sagittal quadriradiates;
tubar skeleton of apical rays of ectosomal sagittal quadriradiates and subgastral sagittal quadriradiates;
choanosomal skeleton of facial rays of subgastral sagittal quadriradiates and choanosomal sagittal
quadriradiates; skeleton of the chamber layer typically composed of the centripetally and centrifugally
directed apical rays of subgastral and subchoanosomal quadriradiates, but choanosomal sagittal
triradiates and confused chamber layer quadriradiates may be present while choanosomal quadriradiates
may be absent (Burton, 1963).
Baeria Miklucho-Maclay, 1870 (type species: Baeria ochotensis Miklucho-Maclay, 1870) ovate, globular, villose surface, apical oscule; ectosomal skeleton a tangential layer of regular
triradiates, with oxeas and microxeas set at right angles to the surface; skeleton of chamber layer of
scattered quadriradiates; choanosomal skeleton a tangential layer of quadriradiates (Burton, 1963).
Kuarrhaphis Dendy & Row, 1913 (type species: Leucyssa cretacea Haeckel, 1872) encrusting to massive, low growing, surface even, non-hispid, oscules not seen; skeleton of needle-eye
spicules (Burton, 1963).
Leucettaga Haeckel, 1872 (type species: Leucetta pandora var. loculifera ) - leuconoid
aquiferous system, sponge spherical, non-hispid, apical oscule; skeleton composed of a confused mass
of triradiates, which are mostly irregular and which form a cortical skeleton as well as the skeleton of
the chamber layer; gastral cavity traversed by numerous endogastric septa which possess a special
skeleton of their own in the form of minute radiates (Dendy & Row, 1913; Burton, 1963).
Leucilla Haeckel, 1872 (type species: Leucilla amphora Haeckel, 1872) (syn. Polejna
Lendenfeld, 1885) - tubular, sessile; surface even, non-hispid; oscule apical; ectosomal skeleton of facial
rays of ectosomal quadriradiates; skeleton of chamber layer of apical rays of quadriradiates similar to
those in ectosomal layer but more irregular, and basal rays of subgastral sagittal quadriradiates, with
small irregular quadriradiates scattered between them; choanosomal skeleton of paired and apical rays of
subgastral sagittal quadriradiates and the facial rays of a tangential layer of choanosomal sagittal
quadriradiates (Burton, 1963).
Syculmis Haeckel, 1872 (type species: Syculmis synapta Haeckel, 1872) - tubular, sessile, with
root tuft; surface even, non-hispid; apical oscule with marginal fringe; ectosomal skeleton of facial rays
of subregular to sagittal quadriradiates; skeleton of chamber layer of apical rays of ectosomal
quadriradiates and basal rays of subgastral sagittal quadriradiates; choanosomal skeleton of paired and
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 FAMILY LEPIDOLEUCONIDAE VACELET, 1967.
DEFINITION: Minute rounded sponges; ectosomal skeleton (exopinacoderm) consists of several layers
of overlapping (not fused) triangular or rounded scales, derived from triradiate spicules; surface has a
single osculum surrounded by several layers of quadriradiate spicules and diactines, and ostia
surrounded by triradiates and microdiactinal spicules; choanosome lacks megascleres but has
microquadriradiates scattered; basipinacoderm region (at the base of the sponge) has scales and
triactines; tuning fork spicules or sagittal triactines never present; choanocytes with apical nuclei; larvae
are amphiblastula.
REVIEWS: Nil.
GENERA:
Lepidoleucon Vacelet, 1967 (type species: Lepidoleucon inflatum Vacelet, 1967) - as for
family.
q FAMILY STAURORRHAPHIDAE JENKIN, 1908.
DEFINITION: Solitary, tubular sac-shaped growth forms with well-developed spicule fringe around
terminal oscule; continuous cortex covers the choanosome, perforated by ostia; ectosomal tetractines
never present, and tangential atrial skeleton present only in oscular region; subatrial quadriradiates
('chiactines') present and equiangular; symmetrical and asymmetrical triradiates and oxeas scattered
freely within choanosomal skeleton, projecting through cortex; aquiferous system is syconoid or
leuconoid.
SCOPE: Two genera.
REVIEWS: Dendy & Row (1913), Burton (1963) and Borojevic (1968).
GENERA:
Achramorpha Jenkin, 1908 (type species: Achramorpha truncata Topsent, 1907) - thin-walled
tubular, apical oscule with well developed marginal fringe; skeleton of chamber layer composed of basal
rays of choanosomal radiates and, usually of proximal parts of radial oxeas; choanosomal skeleton of
paired and apical rays of subgastral triradiates and quadriradiates, with sometimes a tangential layer of
quadriradiates (or triradiates); ectosomal skeleton, when present, a tangential layer of triradiates or
quadriradiates (Burton, 1963).
Megapodon Jenkin, 1908 (types species: Leuconia crucifera Poljaeff, 1883) - unknown
shape, hispid surface; ectosomal skeleton a tangential layer of triradiates, with oxeas and trichoxeas
projecting from the surface; skeleton of chamber layer of centrifugally-directed basal rays of subgastral
sagittal quadriradiates, together with scattered triradiates; choanosomal skeleton of quadriradiates
(Burton, 1963).
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 FAMILY MINCHINELLIDAE DENDY & ROW, 1913.
DEFINITION: Encrusting and lamellate growth forms; oscules may be supported by di-, tri- and
tetractinal spicules; choanosome with a rigid skeleton of fused quadriradiate spicules cemented together,
or formed by intertwined rays of the triradiate and quadriradiate spicules at the base of the sponge; free
spicules may include tri- and quadriradiates, monactinal or diactinal and tuning fork spicules, some or
all of in the ectosomal skeleton; subgastral sagittal radiates apparently absent; nuclei of choanocytes are
apical; larvae are amphiblastula; canal system in all known forms is leuconoid.
SCOPE: Five extant genera are recognised.
REVIEWS: Hartman (1958), Vacelet (1970, 1981), Brien et al. (1973), Hartman (1980).
GENERA:
Petrostoma Dderlein, 1892 (type species: Petrostoma schulzei Dderlein, 1892) - digitate
processes arising from a common base; surface even, texture hard; oscules not obvious; subregular to
sagittal quadriradiates of the chamber layer fused together laterally by calcareous cement into a network;
ectosomal skeleton of separate quadriradiates and triradiates and bunches of tuning-fork spicules
(Burton, 1963).
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Minchinella Kirkpatrick, 1908 (type species: Minchinella lamellosa Kirkpatrick, 1908) flabellate; even surface, minutely hispid; oscules papillate, pores on fistular processes; texture hard; main
skeleton of fused quadriradiates; ectosomal skeleton a palisade of microxeas, with microxeas of several
sizes, triradiates and quadriradiates with paired rays, and tuning-fork spicules in pore and ocular
processes (Burton, 1963).
Monoplectroninia Pouliquen & Vacelet, 1970 (type species: Monoplectroninia hispida
Pouliquen & Vacelet, 1970) - choanosomal skeleton composed of a basal layer made of one category of
small tetractines cemented together by their basal actines, while their apical actine remains free and
pointed. Cortical skeleton composed of free spicules (Borojevic, Boury-Esnault & Vacelet, in prep.).
Plectroninia Hinde, 1900 (type species: Plectroninia halli Hinde, 1900 [fossil form]) quadriradiates of the main skeleton with their facial rays truncated or expanded terminally and fused at
the end with facial rays of adjacent spicules, while the apical rays remain free and pointed; ectosomal
skeleton of separate radiates, including tuning fork spicules and oxeas (Burton, 1963); P. hindei
Kirkpatrick - encrusting; surface even, granular, oscules not seen; texture hard; main skeleton of fused
quadriradiates, with a basal layer of sagittal triradiates, and with irregular triradiates, spined microxeas,
tuning-fork spicules, spined pin-shaped spicules (Burton, 1963).
Tulearinia Vacelet, 1977 (type species: Tulearinia stylifera Vacelet, 1977) - encrusting; hispid
surface; ectosomal skeleton with long styliform diactines both tangential to and erect on the basal
skeleton; large sagittal triactines form a cortex laying parallel to the surface; within the choanosome are
also numerous microdiactines; basal skeleton composed of fused tetractines having basal actines bound
together by interlamellar cement without suture, and the beds under the rays of triactines have the same
form of union (Vacelet, 1977).
q FAMILY PETROBIONIDAE BOROJEVIC, 1979.
DEFINITION: Hemispherical or conical growth forms; ectosomal skeleton contains sagittal triradiates,
tuning fork spicules and quadrirradiates, some of which also extend into the mesohyl; basal skeleton
composed of spherulitic units of calcite, fused together to form hemispherical mass, each unit with
terminal osculum surrounded by collars of quadriradiates; living tissue penetrates into the calcitic mass
only for short distances, and free spicules in the mesohyl include microdiactines; triradiates, with a
rudimentary fourth ray, also occur at the base of living tissue; nuclei of choanocytes are apical; larvae
are amphiblastula.
REVIEWS: Nil.
GENERA:
Petrobiona Vacelet & Lvi, 1958 (type species: Petrobiona massiliana Vacelet & Lvi, 1958)
- as for family.
q FAMILY LELAPIIDAE DENDY & ROW, 1913.
DEFINITION: Tubular, sessile growth forms; surface even, non-hispid; apical oscule with fringed
margin; ectosomal skeleton of tangential sagittal triradiates and microxeas set at right angles to surface;
skeleton of the chamber layer composed of large scattered oxeotes, loose bundles of tuning-fork spicules
and more rarely bundles of slender oxeas, and subgastral sagittal triradiates; choanosomal skeleton
composed of tangential sagittal triradiates and more rarely sagittal quadriradiates.
SCOPE: Three genera of which two are valid.
REVIEWS: Hartman (1958), Burton (1963), Vacelet (1970), Brien et al. (1973), Vacelet (1977a),
Vacelet (1981), Hartman (1982).
GENERA:
Kebira Row, 1909 (type species: Kebira uteoides Row, 1909) - choanosomal skeleton with
spicular tracts constituted by triactines with rudimenary paired actines; atrial and cortical skeleton
composed of triactines and diactines (Borojevic, Boury-Esnault & Vacelet, in prep.).
Lelapia Gray, 1867 (type species: Lelapia australis Gray, 1867) (syn. Paralelapia Hozowa,
1923) - choanosomal skeleton has spicular tracts constituted by diapasons (tuning-fork spicules)
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
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), Lvi (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.
q FAMILY PHERONEMATIDAE GRAY, 1870.
SYNONYM: Semperellidae Schulze.
DEFINITION: Thick-walled vase-shaped, or columnar and lamellate growth forms; oscules single,
terminal, or grouped and dispersed on opposite sides of lamellae, or grouped into seive-plates and
scattered indiscriminantly; dermal spicules are scepters derived from marginal prostals (i.e. spicules
projecting around the oscules) and pleural prostals (i.e. spicules projecting from the sides of the body);
choanosomal spicules are uncinates and scepters, and hexactine and/or pentactines support the
choanosome; basal spicules have bidentate terminal anchors, and tufts of basal spicules are never twisted
nor do they form axial columns.
SCOPE: Six genera are included.
REVIEWS: Nil.
GENERA:
Pheronema Leidy, 1868 (type species: )
Platylistrum Schulze, 1904 (type species: )
Poliopogon Thomson, 1873 (type species: )
Schulzeviella Tabachnick, 1990 (type species: )
Semperella Gray, 1868 (type species: ) (syn. Meyerina Gray) Sericolophus Ijima, 1894 (type species: )
q FAMILY MONORHAPHIDIDAE IJIMA, 1927.
DEFINITION: Body cylindrical or rounded, with seive-like membranes along one side of the body
covering openings to the aquiferous system; whole body is perched on the end of a huge single basal
anchoring spicule, undoubtedly the largest siliceous structure produced by any animal; most
choanosomal megascleres are stauractines, with a single short ray perpendicular to the long axis of the
spicule, although sometimes this short ray is absent producing a pseudodiactinal spicule; prostals are
absent (except for the elongated basal spicule), and uncinates are moderately common.
REVIEWS: Nil.
GENERA:
Monoraphis Schulze, 1904 (type species: )
q FAMILY HYALONEMATIDAE GRAY, 1857.
DEFINITION: Spheroid or ovoid bodies, although actual shape can be very variable; tufts of long basal
spicules anchor sponges into soft sediments, each bearing a terminal "anchor" (consisting of an invertedconical swelling bearing a circle of several short teeth); basal spicules compactly bundled and twisted
dextrally, forming a single basal tuft extending into sponge body and forming a compact axial columella;
apical end of basal spicules produce a small projection called the gastral cone; exhalant canals open on
97
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 FAMILY FARREIDAE GRAY, 1872.
DEFINITION: Funnel-shaped and tubular, simple or branched growth forms; branching tubes all
approximately equal in width, each branch ending with an oscule; lateral wall of skeleton a single layer
of paratangential dictyonal framework, with no or little space for canals but consisting of a secondary
quadratic skeletal mesh consisting of nodes which are all a hexactin centre; six internodal beams radiate
from these nodes, usually laterally apposed and amalgamated combinations of two rays, each from
adjacent hexactins; this typically regularly-meshed structure may become irregular due to accretion of
dictyonal hexactins, in indefinite orientation, formed after the primary framework is established; dermal
and gastral spicules pentactines; uncinates usually present; microscleres include oxyhexasters,
tylohexaster and discohexasters; sceptules present usually in the form of sarule, lonchiole or clavule,
sometimes including monactinal triaxons, but lacking scopules; flagellated chambers laterally branched.
REVIEWS: Schulze (1887, 1899), Ijima (1927).
GENERA:
Claviscopulia Schulze, 1899 (type species: )
Farrea Bowerbank, 1862 (type species: Farrea occa Bowerbank, 1863)
Lonchiphora Ijima, 1927 (type species: )
Sarostegia Topsent, 1904 (type species: )
q FAMILY EURETIDAE ZITTEL, 1877.
DEFINITION: Funnel-shaped, tubular and vase-shaped growth forms, the latter with tubular branches
opening to the exterior through accessory oscules on the sides of vases and opening internally into a
wide common gastral cavity; lateral wall of skeleton three dimensional dictyonal framework similar to
Farreidae, although meshes always small, triangular, quadrangular or irregular, and certain nodes give
off more than six internodal beams; body wall usually not canalized, and generally unaccompanied by
aporhysis; dermal spicules pentactine or hexactines, with teeth on distal ray, or sometimes secondarily
absent; gastral spicules either pentactines or hexactines, similar in form to dermal spicules; microscleres
include diverse hexasters, scopules and uncinates; clavules absent, sarules rarely present.
SCOPE: Fourteen genera are included.

REVIEWS: Schulze (1887), Ijima (1927)
GENERA:
Bathyxiphus Schulze, 1899 (type species: Bathyxiphus subtulis Schulze, 1899)
Calyptorete Okada (type species: ) Chonelasma Schulze, 1886 (type species: Chonelasma lamella Schulze, 1886)
Conorete Ijima, 1927 (type species: )
Eurete Semper, 1868 (type species: )
Gymnorete Ijima, 1927 (type species: )
Heterorete Dendy, 1916 (type species: )
Iphiteon Bowerbank, 1869 (type species: )
Lefroyella Thomson, 1877 (type species: )
Margaritella Schmidt, 1880 (type species: )
Myliusia Gray, 1859 (type species: )
Pararete Ijima, 1927 (type species: )
Periphragella Marshall, 1875 (type species: )
Pleurochorium Schrammen, 1910 (type species: )
Ptychodesia Schrammen, 1910 (type species: )
q FAMILY CRATICULARIIDAE RAUFF, 1893.
SYNONYM: Laocaetidae Povrel.
DEFINITION: Cup-shaped growth forms; lateral wall skeleton has a dictyonal framework traversed by
two sets of tubular cavities entering radially from opposite sides, running parallel but in opposite
directions, forming an alternating longitudinal series originating either from the dermal or gastral sides
of the framework (epirhyses or aporhyses, respectively); dermal and gastral spicules pentactines with
toothed distal rays; scopules, oxyhexasters and discohexasters present; uncinates absent.
SCOPE: Only one Recent genus is recognised.
REVIEWS: Nil.
GENERA:
Laocoetis Pomrel, 1872 (type species: ) (syn. Leptophragmella Reid, 1963) q FAMILY APHROCALLISTIDAE GRAY, 1867.
DEFINITION: Vase-shaped or branching tubular growth forms; oscules on outpockets on the side walls;
lateral wall dictyonal framework perforated by a system of tubular cavities (diarhyses), running radially
through the skeleton; each cavity occupied by a single lobate flagellated chamber; cavities arranged in
alternating or regularly hexagonal groups; dictyonal framework between cavities forms irregular meshes;
choanosomal hexactines regular or compressed laterally so that all six rays lie in one plane; dermal
spicules are hexactines or pentactines, with teeth on the distal ray; gastral spicules similar to dermals, or
they are rhabdodiactines (curved diactinals); hexaster microscleres include oxy-, tylo- onycho- or
discohexasters; scopules and uncinates always present.
SCOPE: Two Recent genera are included here, one dubiously so.
REVIEWS: Nil.
GENERA:
Aphrocallistes Gray, 1858 (type species: )
Heterochone Ijima, 1927 (type species: )
q FAMILY TRETODICTYIDAE SCHULZE, 1886.
DEFINITION: Thick-walled cup-shaped, funnel-shaped or plate-like growth forms, or reticulate masses
of branching tubes or solid cylinders; lateral wall dictyonal framework composed of irregular tri- or
quadrangular meshes with multiradiate nodes, marked by development of extensive canal system
(schizorhyses) containing labyrinths of flagellated chambers; chambers interconnecting and/or dividing
and branching, running from gastral side (covered by a membrane) to dermal side of dictyonal
framework (the latter covered by seive-like membrane); diverse hexaster microscleres present; uncinates
and scopules also present.
SCOPE: Ten genera are included in this family.
REVIEWS: Nil.
GENERA:
Anomochone Ijima, 1927 (type species: )
Auloplax Schulze, 1904 (type species: )
Cyrtaulon Schulze, 1887 (type species: )
99
Hexactinella Carter, 1885 (type species: )

Psilocalyx Ijima, 1927 (type species: )
Sclerothamnopsis Wilson, 1904 (type species: )
Sclerothamnus Marshall, 1875 (type species: )
Tretocalyx Schulze, 1900 (type species: )
Tretodictyum Schulze, 1886 (type species: )
Tretorete Ijima, 1927 (type species: )
q FAMILY AULOCALYCIDAE IJIMA, 1927.
DEFINITION: Vasiform, spherical or tubular growth forms; main osculum above a number of lateral
oscules on the sides; skeletal framework with irregular meshes, lacking canals in the dictyonal
framework; hexactines inserted individually into dictyonal framework, with spicule rays frequently
elongated and curved, intersecting one another at various angles, and fused together at points of their
intersection (the rays often connected by synapticulae), or where they are laterally apposed or terminate
by abutting on others; dermal and gastral spicules always pentactines; discohexasters always present,
with or without oxyhexasters; scopules rarely present; uncinates present or absent.
SCOPE: Five genera are included.
REVIEWS: Reiswig & Tsurumi (1996)
GENERA:
Aulocalyx Schulze, 1886 (type species: Aulocalyx irregularis Schulze, 1886)
Euryplegma Schulze, 1886 (type species: )
Fieldingia Saville Kent, 1870 (type species: )
Leioplegma Reiswig & Tsurumi, 1996 (type species: Leioplegma polyphyllon Reiswig &
Tsurumi, 1996)
Rhabdodictyum Schmidt, 1880 (type species: )
Tretopleura Ijima, 1927 (type species: )
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 FAMILY AULOCYSTIDAE SCHULZE, 1886.
SYNONYM: Diapleuridae Ijima.
DEFINITION: Ovoid and bulbous stalked growth forms, with branching and rejoining aquiferous tubes
and interconnected canals; external surface with secondarily produced layer consisting of a feltwork of
fine rays from projecting stauractines and pentactines; dermal and gastral pentactines present;
microscleres include oxy-, disco- and graphiohexasters.
SCOPE: Only two genera are presently included in this family.
REVIEWS: Schulze (1887, 1904), Ijima (1927), Reiswig (1991).
GENERA:
Neoaulocystis Rezvoy, Zhuravleva & Koltun, 1962 (type species: ) (syn. [Aulocystis] Schulze,
1886) Diapleura Ijima, 1927 (type species: )
q FAMILY DACTYLOCALYCIDAE GRAY, 1867.
DEFINITION: Vasiform growth forms with folded walls; lychniscs of dictyonal framework of lateral
wall secondarily fused producing a "pseudohexactinosidan" skeleton; dermal pentactines and hexactines
present, and free hexactines also occur as parenchymal spicules; microscleres include oxy- and
discohexasters; uncinates and scopules absent.
SCOPE: Only one Recent genus is included in this family.
REVIEWS: Reid (1957), Reiswig (1991)
GENERA:
Dactylocalyx Stutchbury, 1841 (type species: )
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
of small dermal spicules overlying larger hypodermal pentactines, with the unpaired ray extending
inwards.
Four extant families are recognized.
q FAMILY LEUCOPSACASIDAE IJIMA, 1903.
SYNONYMS: Leucopsacinae Ijima; Leucopsacidae Ijima; Leucopsacasidae Ijima.
DEFINITION: Thick-walled cup-shaped or ovoid, stalked growth forms; body anchored to substratum
by basal spicules; dermal skeleton has large dermal pentactines with unpaired ray directed inwards,
without hypodermal spicules; parenchymal spicules hexactines and rhabdodiactines; microscleres
hexasters including discohexasters, sigmatocomes, floricomes, but not oxyhexasters.
SCOPE: Five Recent genera are included in the family.
REVIEWS: Ijima (1927).
GENERA:
Caulocalyx Schulze, 1886 (type species: )
Chaunangium Schulze, 1904 (type species: )
Chaunoplectella Ijima, 1896 (type species: )
Leucopsacus Ijima, 1898 (type species: )
Placoplegma Schulze, 1896 (type species: )
q FAMILY EUPLECTELLIDAE GRAY, 1867.
DEFINITION: Tubular, massive or cup-shaped growth forms ("venus' flower baskets"), often with many
open oscules; bases either stalked, firmly attached to substratum, or with tufts of monactinal or
anisodiactinal basal spicules; dermal skeleton has large hexactinal spicules (dermalia) with proximal ray
longest; hypodermal spicules absent; parenchymal spicules hexactines with two to six rays; hexasters
diverse, including floricomes, graphio-, oxy- and onychohexasters.
SCOPE: Euplectellids were split into two subfamilies by Ijima (1903), (Euplectellinae Ijima, 1903,
Corbitellinae Ijima, 1903), containing 18 nominal genera although it is not known how many of these
are really valid.
REVIEWS: Schulze (1886, 1887, 1899), Lendenfeld (1915), Ijima (1903, 1927).
GENERA:
Subfamily Euplectellinae Ijima, 1903
Acoelocalyx Topsent, 1910 (type species: )
Docosaccus Topsent, 1910 (type species: )
Euplectella Owen, 1841 (type species: )
Holascella Lendenfeld, 1915 (type species: )
Holascus Schulze, 1886 (type species: Holascus stellatus Schulze, 1886)
Malacosaccus Schulze, 1886 (type species: )
Subfamily Corbitellinae Ijima, 1903
Bolosoma Ijima, 1903 (type species: )
Corbitella Gray, 1867 (type species: )
Dictyaulus Schulze, 1895 (type species: )
Dictyocalyx Schulze, 1886 (type species: )
Heterotella Gray, 1867 (type species: )
Hertwigia Schmidt, 1880 (type species: )
Hyalostylus Schulze, 1886 (type species: )
Regadrella Schmidt, 1880 (type species: )
Rhabdopectella Schmidt, 1880 (type species: )
Saccocalyx Schulze, 1895 (type species: )
Trachycaulus Schulze, 1886 (type species: )
Walteria Schulze, 1886 (type species: )
q FAMILY ROSSELLIDAE GRAY, 1872.
SYNONYM: Rossellidae Schulze.
DEFINITION: Cup-like or sac-shaped growth forms, often with a stalk, attached directly to substratum,
or with basal processes, or with tufts of pentactinal basal spicules; secondary oscules may be present in
addition to main terminal oscule; dermal skeleton with small roughened pentactines, stauractines or
rhabdodiactine spicules, having similar rays and not markedly spined; distal rays of dermal spicules, if
developed, similar to remaining rays and not markedly spined; hypodermal spicules pentactines or
rhabdodiactines or both, sometimes protruding through surface so that spicule rays form veil-like
101
covering over the sponge; parenchymal spicules hexactines and/or rhabdodiactines; microscleres oxyand 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: )
Subfamily Rossellinae Schulze, 1872 - without discoasters, without strobiloplumicomes.
Anaulosoma Kirkpatrick, 1907 (type species: )
Anoxycalyx Kirkpatrick, 1907 (type species: Anoxycalyx ijimai Kirkpatrick, 1907)
Aphorme Schulze, 1899 (type species: )
Asconema Kent, 1870 (type species: ) (syn. Askonema; de Laubenfels, 1936 [lapsus]) Aulochone Schulze, 1886 (type species: )
Aulorossella Kirkpatrick, 1907 (type species: Aulorossella levis Kirkpatrick, 1907)
Aulosaccus Ijima, 1896 (type species: )
Bathydorus Schulze, 1886 (type species: Bathydorus fimbriatus Schulze, 1886)
Crateromorpha Gray, 1872 (type species: )
Gymnorossella Topsent, 1916 (type species: )
Hyalascus Ijima, 1896 (type species: )
Rossella Carter, 1872 (type species: Rossella antarctica Carter, 1872)
Schaudinnia Schulze, 1900 (type species: )
Scolymastra Topsent, 1916 (type species: Scolymastra joubini Topsent, 1916)
Scyphidium Schulze, 1900 (type species: )
Trichasterina Schulze, 1900 (type species: )
Vitrollula Ijima, 1898 (type species: )
Subfamily Acanthascinae Schulze, 1897 - with discoctasters.
Acanthascus Schulze, 1886 (type species: )
Rhabdocalyptus Schulze, 1886 (type species: ) (syn. Acanthosaccus Schulze)
Staurocalyptus Ijima, 1897 (type species: )
q FAMILY CAULOPHACIDAE SCHULZE, 1886.
DEFINITION: Solitary or branching, cup-shaped and mushroom-shaped growth forms, with the stalk
firmly attached to substrata; dermal skeleton with small hexactines, sometimes pentactines bearing
spined proximal rays; hypodermal spicules pentactines and sometimes rhabdodiactines; parenchymal
spicules hexasters and rhabdodiactines; microscleres include disco-, onycho- or oxyhexasters, sometimes
with strobiloplumicomes.
REVIEWS: Schulze (1887), Ijima (1903, 1927), Lendenfeld (1915)
GENERA:
Caulodiscus Ijima, 1927 (type species: Caulophacus lotifolium Ijima, 1903)
Caulophacella Lendenfeld, 1915 (type species: )
Caulophacus Schulze, 1886 (type species: Caulophacus latus Schulze, 1886)
Sympagella Schmidt, 1870 (type species: ) (syn. Calycosilva Lendenfeld; de Laubenfels, 1936)
q PORIFERA INCERTAE SEDIS
Names listed here cannot presently be placed in any known family or higher taxon with any confidence.
Some genera listed are unrecognisable because their types are species inquirendae. For species names
listed where type material is extant, they are considered to be species inquirendae. For species names
listed where type material could not be found, and must be presumed to be lost, these names are classed
as nomina dubia. Many of de Laubenfels (1936) genera are included here as incertae sedis because they
were usually erected without re-examination of type material, even in cases where original descriptions
were poorly known or virtually unrecognisable.
Acanthostylotela Burton & Rao - with acanthose megascleres; possibly Myxillidae

Adreissa Topsent - papillate, only megascleres styles with larger ones forming tracts; possibly
Suberitidae
Alcyoncellum Quoy & Gaimard (type species: ) (syn. Eudictyon Marshall, Eudictyum Schulze,
Habrodictyon Wyville Thomson, Taegeria Schulze) - possibly Euplectellidae (de Laubenfels, 1936).
Allantella Hallmann (according to de Laubenfels, 1936: 84; not found in his publications)
Ammoconia Haeckel, 1889 (type species: Ammoconia auloplegma Haeckel, 1889) unrecognisable calcarean
Ammolynthus Haeckel, 1889 (type species: Ammolynthus haliphysema Haeckel, 1889) unrecognisable calcarean
Amphoridium Haeckel, 1864 (type species:) - unrecognisable calcarean
Ancorella Lendenfeld (type species: ) - microxea-like streptaster microscleres without spines;
possibly Ancorinidae (de Laubenfels, 1936)
Annandalia Topsent (type species: ) - possibly Clionidae.
Arcesios Duchassaing & Michelotti, 1864 (type species: Arcesios prominula Duchassaing &
Michelotti, 1864).
Astromimus Lendenfeld - possible synonym of Epipolasis
Aulorhipis Ehlers - probably not a sponge (de Laubenfels, 1936:213)
Badiaga Girtanner - unrecognisable freshwater sponge (de Laubenfels, 1936)
Baicalolepis Makushok - unrecognisable freshwater sponge (de Laubenfels, 1936)
Bethia de Laubenfels, 1936 (type species: Dictyocylindrus laciniatus Carter, 1879) subradiate, hemispherical architecture reminiscent of Suberitidae, with styles and oxeas, no microscleres.
Bursalina Schmidt - virtually unrecognisable, possibly synonym of Suberites (de Laubenfels)
Cacochalina Schmidt, 1870 (type species: Cacochalina subtilis Schmidt, 1870).
Callisphaera Gray, 1870 - unrecognisable (de Laubenfels, 1936)
Callites Schmidt - unrecognisable (de Laubenfels, 1936)
Carterias Svarchevsky - unrecognisable freshwater sponge (de Laubenfels, 1936)
Cartilospongia Bowerbank - unrecognisable (de Laubenfels, 1936)
Cavochalina Carter, 1882 (type species: Cavochalina digitata var. arenosa Carter, 1882)
Cerelasma Haeckel, 1889 (type species: Cerelasma gyrosphaera Haeckel, 1889) unrecognisable
Cerelpemma de Laubenfels, 1936 (type species: Psammopemma radiolarium Haeckel, 1889) unrecognisable
Cerelpsamma de Laubenfels, 1936 (type species: Holopsamma argillaceum Haeckel, 1889) unrecognisable
Chalinoraphis Lendenfeld, 1887 (type species: Chalinoraphis armata Lendenfeld, 1887)
Collospongelia Ferrer-Hernandez - possibly a Dysideidae (de Laubenfels, 1936).
Corythophora Ferrer-Hernandez, 1932 (type species: Corythophora ijimai Ferrer-Hernandez,
1932) - pentacts as an important skeletal element like Amphidiscophora; with pentacts and uncinates like
Pheronematidae; has basal spicules that terminate in cones but the rhabdodiactines do not form the
principal skeleton as in Hyalonemidae; in addition with peculiar "xylostyles" that are twisted and bent
like the letter "j" at one end (de Laubenfels, 1936); possibly a conglomerate of several species.
Coscinospongia Bowerbank - "lithistid" with desmas and dichotriaenes with peculiarly twisted
rhabds (de Laubenfels, 1936)
Crellastrina (type species: ).
Cryptax de Laubenfels, 1954 (type species: Cryptax orygmi de Laubenfels, 1954) - excavating
into coralline substrate; megascleres exclusively tylostrongyles dispersed in confusion within the
skeleton, microscleres absent; possibly a synonym of Cliona.
Cryptospongia Burton, 1928 - a hydroid (Berquist, 1980).
Cryptotethya Dendy (type species: ) - with only oxeas as megascleres and asters that are
chiasters; possibly Coppatiidae.
Cyathella Schmidt - unrecognisable (de Laubenfels, 1936)
Cystispongia Roemer - unrecognisable (de Laubenfels, 1936)
Dactylochalina Lendenfeld, 1885 (type species: Chalina cervicornis Bowerbank, 1866).
Daedalopelta Sollas, 1888 (type species: Daedalopelta nodosa Sollas, 1888) - "lithistid" with
discotriaenes as principal spicules, and very much deformed desmas, microscleres spirasters to
streptasters (de Laubenfels, 1936)
Damo Gray - unrecognisable (de Laubenfels, 1936)
Deanea Bowerbank, 1875 - unrecognisable hexactinellid (de Laubenfels, 1936)
Delectona de Laubenfels, 1936 (type species: Alectona higgini Carter, 1880) - with rhabds
subdivided by spiny or lumpy subspherical swellings; possibly not a sponge at all (de Laubenfels, 1936).
Dercitancorina Topsent (type species: ) - with some calthrops having dichotomous
modifications, microscleres straight streptasters; possibly Plakinidae (de Laubenfels, 1936).
Desmacodes (type species: )
Diaretula Schmidt, 1880 (type species: Diaretula cornu Schmidt 1880) - supposed
relationship to Euretidae (de Laubenfels, 1936).
103
Diplodemia Bowerbank, 1862 (type species: Diplodemia vesicula Bowerbank, 1862).

Diplacodium Schmidt, 1880 (type species: Diplacodium mixtum Schmidt, 1880) - barely
recognisable but possibly related to Diapleura (de Laubenfels, 1936).
Distomus Oken - probably not a sponge (de Laubenfels, 1936).
Doconesthes Topsent - uncertain placement within Hexactinellida.
Dotona Carter - with only oxea megascleres and 2 peculiar sorts of microscleres reminiscent of
Donotella and Scantilletta (de Laubenfels, 1936); possibly Clionidae.
Dotonella Dendy - tylostyle megascleres, spiral spiraster microscleres and others having a
straight shaft with spines arranged spirally (de Laubenfels, 1936); probably Spirastrellidae.
Dymnus Gray - unrecognisable although with inferred relationship to Chalinidae (de
Laubenfels, 1936).
Ectyobatzella Burton & Rao (type species: Ectyobatzella enigmatica Burton and Rao, 1932) conulose with a conspicuous fleshy dermis; with only monactinal megascleres being more-or-less
malformed styles; possibly Phoriospongiidae.
Ectyonopsis Carter - principal spiculesacanthostrongyles but some modified to acanthostyles
(de Laubenfels, 1936); possibly Myxillidae.
Euryades Duchassaing & Michelotti, 1864 (type species Euryades notabilis Duchassaing &
Michelotti, 1864) - unrecognisable (de Laubenfels, 1936).
Euthymus. (type species: )
Evenor Duchassaing & Michelotti, 1864 - unrecognisable, probably not a sponge (de
Laubenfels, 1936)
Fasubera de Laubenfels, 1936 (type species: Hymedesmia lipochela Dendy, 1922) - thinly
encrusting, with tylostyles of 2 sizes, the smaller pseudo-echinating, but lacking ectosomal diactines (de
Laubenfels, 1936); possibly Hymedesmiidae.
Fistula Oken (type species: Spongia fulva Pallas, 1766) - unrecognisable (de Laubenfels,
1936).
Gilchristia Burton (type species: ) - with larger styles and smaller tylostyles, occurring in
symbiosis with another sponge (de Laubenfels, 1936); probably a synonym of Suberites.
Halina Bowerbank, 1858 (type species: Halina bucklandi Bowerbank, 1858) - with calthrops
occasionally with dicho- modification, streptasters straight, and toxiform microscleres (de Laubenfels,
1936); possibly Plakinidae.
Halinastra de Laubenfels, 1936 (type species: Pachastrella exostotica Schmidt, 1868) - with
both euasters and straight streptaster microscleres, megascleres oxeas and calthrops only; possibly
Pachastrellidae.
Halyphysema Bowerbank - unrecognisable (de Laubenfels, 1936).
Heterophymia Pomel - unrecognisable (de Laubenfels, 1936).
Hezekia de Laubenfels, 1934 (type species: Hezekia demera de Laubenfels, 1934) - only spiny
microrhabds as microscleres; possibly Ancorinidae.
Holorodesmia Topsent, 1928 (type species: Hymedesmia flaccida Topsent, 1928) - polytylote
dermal spicules, palmate isochelae (de Laubenfels, 1936); possibly Iophonidae
Hyalocaulus Marshall - possibly Euretidae (de Laubenfels, 1936).
Janulum de Laubenfels, 1936 (type species: Isodictya spinispiculum Carter, 1876) - peculiar
twice-bent strongyles, heavily spined at the middle but not at the ends; possibly Myxillidae.
Jasplakina de Laubenfels, 1954 (type species: Jasplakina nux de Laubenfels, 1954) - with
oxeas of 2 or more distinct types, microscleres microxeas, eusasters and calthrops-like asters with few
rays; possibly Coppatidae.
Joannella Schmidt, 1880 (type species: Joannella compressa Schmidt, 1880) - "almost
certainly a hylospongid (de Laubenfels, 1936); possibly Hyalonematidae.
Kallispongia Wright - unrecognisable (de Laubenfels, 1936).
Keratylum de Laubenfels, 1936 (type species: Suberites capillitium Topsent, 1892) - only
tylostyles embedded in fibres, generally forming reticulate architecture; possibly Suberitidae, close to
Caulospongia.
Kowalewskyella Svartschevsky - with basal horizontal layer of acanthoxeas, no microscleres;
possibly Myxillidae.
Lepidothenea de Laubenfels, 1936 (type species: ) (syn. [Lepidospongida] Dendy (preocc.)) few desmas, phyllotriaenes present instead of dichotriaenes.
Leptochelastra de Laubenfels, 1936 (type species: Hymeraphia toureti Topsent, 1894) principal tylostyles with peculiar double heads, accessory spicules acanthostyles with pronouncedly
recurved spines; also allegedly with isochelae and asters but these are very poorly described and possibly
contaminants; possibly Anchinoidae.
Leptosastra Topsent (type species: ) - tylostyles, acanthostyles with recurved spines,
microscleres "odd spiny pyriform" (de Laubenfels, 1936); possibly Anchinoidae.
Lipastrotethya de Laubenfels, 1954 (type species: Lipastrotethya ana de Laubenfels, 1954) irregular mass, cartilagenous consistency; surface tuberculate, hispid; ectosome corticate; choanosome
radiate with spicules more organised at the surface, more confused below, with stongyloxeas of varying
sizes; microscleres absent; uncertain placement in Tethyidae.
Lissomyxilla (type species: )
Litamena Nardo - unrecognisable (de Laubenfels, 1936)
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
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).
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
Stannoma Haeckel, 1889 (type species: Stannoma dendroides Haeckel, 1889); virtually
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).
8. PREFERRED FORMAT FOR SPONGE SAMPLES SENT FOR IDENTIFICATION
q 1.1. Specimen data.
In situ colour photograph/slide (preferrably of live specimen).
Field notes on colouration, overall shape, surface characteristics, mucus exudates, and any
peculiar habits (e.g. burrowing, encrusting other fauna, etc.).
q 1.2. Locality data.
Precise locality, including latitude and longitude.
Depth of collection.
Date of collection.
Collectors name(s).
Mode of collection (SCUBA, dredge, etc.).
Type of habitat (e.g. coral reef, mud fauna, mangrove, rocky shore, etc.).
q 1.3. Fixation and preservation.
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 7080% 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.
Specimens should be posted to:
The Director
Attn. Senior Curator, Sessile Marine Invertebrates
Queensland Museum
P.O. Box 3300
SOUTH BRISBANE QLD 4101
AUSTRALIA
CITES Institution AU005
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].
Boury-Esnault, N. & Ruetzler, K. (Eds) (1997). Thesaurus of sponge morphology. Smithsonian
Contributions to Zoology (596): 1-55.
Brien, P., Lvi, C., Sar, M., Tuzet, O & Vacelet, J. (eds) (1973). Spongiaires. Trait de Zoologie.
Anatomie, Systmatique, 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].
107
10. GLOSSARY.
acantho- - with spines.
accessory spicules - old term referring to echinating megascleres (cf. principal, auxiliary spicules).
actine - see monactine.
amphiblastula larva - larval form associated with viviparous development, with an internal cavity and
at least some cytological development, with flagella on anterior hemisphere only.
anastomosing - reticulated, rejoining, referring to cross connections between fibres of tracts.
aniso- - asymmetrically-ended spicule (e.g. anisoxea).
aphodal - (a) a small canal which joins the choanocyte chambers to an exhalant canal (cf. prosodal); (b)
a condition of the aquiferous system in which the choanocyte chambers are joined to the
inhalant system directly by a pinacocyte prosopyle, lacking prosodal canals (cf. diplodal,
eurypylous).
apopyle - an aperture through which water leaves a choanocyte chamber (cf. prosopyl).
aquiferous system - the water conductive (circulatory) system extending from the inhalant pores (ostia)
to exhalant pores (oscula).
archaeocytes - a motile amoeboid cell roaming within the mesohyl of the sponge, with at least 1
nucleolus and many phagosomes, capable of phagocytosis and able to develop into any other
sponge cell type; also known as nucleolate cells.
arenaceous - condition of skeletal architecture in which sand and/or foreign spicule debris partly or
completely replaces native spicules within the sponge skeleton.
asconoid construction - simple tubular body-plan, without folding of the body wall, with the central
cavity (atrium) lined by choanocyte chambers, and a single osculum at the apex (characteristic
of primitive Calcarea, also seen in a few homoscleromorph demosponges) (cf. syconoid,
leuconoid).
atrium - exhalant water cavity leading to 1 or more exhalant canals (oscula); also known incorrectly as
cloaca.
auxiliary spicules - old term referring to second and third categories of megascleres usually found
outside the fibres, dispersed between tracts or on the surface; incorrectly interchanged with the
terms "ectosomal" and "subectosomal spicules" (cf. accessory, principal spicules).
axial canal or filament - central lumen of a spicule, occupied in life by an organic filament.
axial skeleton - organic and/or inorganic skeleton found in the centre or axis of the sponge (cf. extraaxial).
axial construction - see condensed skeleton.
-axon or -actine - number of geometric axes of a spicule.
auto-dermal, auto-gastral spicules - tetractinal spicules lying on or below the surface of the exterior
(dermis) or interior (gastral) cavity, with the asymmetrical free ray directed outwards
(Hexactinellida).
basopinacocytes - outer layer of epithelial cells covering the basal surface, at the point of contact with
the substrate (cf. exopinacocytes).
benthic - living on the bottom of the sea; benthos refers to life forms on the sea bed.
bioerosion - chemical and physical degradation of an organic substrate caused by an association with
another organism.
body plan - grade of construction of sponge body based on the complexity of the aquiferous system and
disposition of choanocyte chambers and interconnecting canals (see asconoid, syconoid,
leuconoid, aphodal, diplodal, prosodal); (not to be confused with the terms body shape or
growth form).
centrangulate - sharp bend or angular curve at the centre.
choanocyte - collar-cell bearing a flagellum surrounded by a collar of cytoplasmic microvilli, used to
produce a water current system and entrapment of small food particles or colloidal material.
choanocyte chamber - cavity lined by spherical clusters of choanocytes with flagella directed into the
water-filled lumen (Demospongiae, Calcarea) (cf. flagellated chamber of Hexactinellida).
choanoderm - strictly a continuous layer of choanocytes lining a single internal cavity (found only in
some Calcarea); also in a generalized sense to include all internal surfaces not bound by exo- or
basipinacocytes (24).
choanosome - region of the sponge containing choanocyte chambers, which includes everything
bounded by the pinacoderm; also known as endoderm or endosome (cf. ectosome).
choanosomal spicules - condition referring to localization of megascleres within (coring) spongin
fibres or tracts (cf. ectosomal and subectosomal spicules).
coeloblastula larva - heterogenous assemblage of larval forms associated with oviparous development,
with a cytologically undifferentiated central region, and a an even distribution of small flagella.
collagen - proteinaceous material forming (a) the intercellular connective "tissue"/ ground substance/
matrix within the mesohyl, formed of minute fibrils which are visible only through electron
microscopy, and collectively called spongin type A; and (b) the macroscopic spongin fibres
producing the organic skeleton; collagen fibrillogenesis is universal within the Porifera.
collagenous filaments - (or collagen fibrils) intercellular fibrils produced by cellular secretion
(fibrillogenesis) forming the ground substance or spongin type A matrix within the mesohyl; 2
forms are recognized, rough and smooth, both requiring electron microscopy to elucidate their
structure.
condensed or compressed - condition of skeletal architecture in which there is a compressed central

axis of fibres and/or spicules, from which arises plumose or plumoreticulate columns of fibres
and/or spicules; also known as axinellid or axinelloid.
coring spicules - spicules found inside (coring) spongin fibres (cf. echinating spicules).
dendritic - condition of skeletal architecture whereby spongin fibres branch but do not rejoin with each
other (i.e. non-reticulate).
dermal - strictly refers to any association with the pinacoderm; also used to refer to structures lying on
or just below the ectosomal membrane or dermis (correct usage is "ectosomal").
diactinal - spicules with 2 diverging rays, representing growth in 2 directions, usually with bilateral
symmetry; diacts may include monaxonic spicules (e.g. oxeas of demosponges), and tetraxonic
derivatives (e.g. "oxea" of Calcarea; uncinate of Hexactinellida) (cf. monactinal).
dictyonal framework - rectangular mesh formed by fused spicules or lychniscs joined together
(Hexactinellida).
dioecious - with the cells of each sex occurring in different individuals (also known as gonochoristic),
but in sponges it is usual for one individual to produce eggs at one time and sperm at another
times (strictly successive or temporal hermaphroditism) (cf. hermaphroditic).
diplodal - a condition of the aquiferous system, where some sponges possess both prosodal and aphodal
canals between choanocyte chambers (cf. aphodal, eurypylous).
echinating spicules - spicules with one end, usually the head, implanted in a spongin fibre or a spicule
column and standing more-or-less perpendicular to the column (found in Poecilosclerida) (cf.
coring spicules).
ectosome - peripheral region of the sponge lacking choanocyte chambers; the term strictly refers to the
unicellular surface layer (pinacocytes), but it is also used to refer to the mineral skeleton found
in the periphery; also known as cortex or dermis (cf. choanosome).
ectosomal spicules - condition referring to localization of megascleres to the ectosomal skeleton (cf.
subectosomal, choanosomal spicules).
extra-axial - organic and/or inorganic skeleton arising from the centre (or axis) and ascending towards
the periphery of the sponge (cf. axial).
eurypylous - condition of the aquiferous system in which there are wide mouthed sac-like choanocyte
chambers without any aphodal canals present (cf. aphodal, diplodal conditions).
exhalant - part of aquiferous system related to the expelling of water from the sponge, and includes all
water vascular structures between the apopyles and the oscula; also known as excurrent system
(cf. inhalant).
exopinacocytes - outer layer of pinacocyte cells covering the free surface of the sponge (cf.
basopinacocytes).
fibres - (macroscopic) discrete column of spongin, forming the organic skeleton (see spongin fibres).
fibrils - (sub-light microscopic) collagenous filaments forming the spongin ground substance (see
collagen fibrils).
filter feeding - process of obtaining food by pumping water through a series of sieves of decreasing size,
finally obtaining small food particles for ingestion usually less than 0.1 m in diameter.
fistule - tubular structure, on the upper surface of some sponges, on which the oscule is situated, used to
exhale water; frequently found on species that burrow into mud or excavate coral.
flagella - the central organelles of choanocyte cells, used to set up a water current via rhythmic beating
and aid in entrapment of food particles.
flagellated chambers - cylindrical chambers lined by choanocytes, which are not embedded within a
cellular matrix, and also lack any connecting canals (Hexactinellida only) (cf choanocyte
chambers of Demospongiae and Calcarea).
fusiform - spicule tapering regularly towards one or both ends (cf. hastate).
gastral - associated with the central atrium or gastral cavity (Hexactinellida) (cf. dermal, parenchymal).
gonochoristic - male and female cells produced by different individuals (cf. hermaphroditic).
ground substance - see collagen.
habit - term used to describe external shape of a sponge, also known as ecophenotype.
halichondroid - condition of skeletal structure, with megascleres arranged in vague tracts which may be
reticulate or scattered in a disorganised criss-cross within the mesohyl.
hastate - spicules with abruptly tapering ends (cf. fusiform).
hermaphroditic - male and female cells occur together in one individual at the same time (cf.
gonochoristic).
hexact - spicule with 6 rays (Hexactinellida).
hymedesmoid - condition of skeletal architecture found in encrusting sponges, in which there is a
flattened basal layer of spongin lying on the substrate, and spicules stand erect on (echinate) this
basal layer.
hypo-dermal, hypo-gastral spicules - tetractinal spicules lying on or below the exterior surface or the
gastral cavity, with the asymmetrical free ray directed inwards (Hexactinellida).
inhalant - part of the aquiferous system related to bringing water into the choanocyte chambers,
including all structures between the ostia and prosopyles; also known as incurrent system (cf.
exhalant).
isodictyal - condition of skeletal architecture in which the reticulation is triangular in 3-dimensions,
produced by single spicules joined together at their ends (nodes) by an accretion of collagenous
spongin.
109
isotropic - condition of skeletal architecture in which there is a disoriented or seemingly random

reticulation of spicules or fibres without distinction between primary and secondary columns.
junior synonym - used in the scientific sense, refers to a proper scientific name given to a particular
species that already has a valid (older) scientific name, in which case the junior synonym is not
regognised as being valid by convention.
keratose sponges - collective term referring to sponges which lack a native mineral skeleton (usually
only includes the orders Dictyoceratida, Dendroceratida, Verongida).
leuconoid - body-plan construction produced by complex folding of the body wall, forcing choanocyte
chambers to become oval and isolated in a maze of canals within the body wall, and chambers
open onto branching excurrent canals (most Demospongiae, some Calcarea) (cf. asconoid,
syconoid).
matrix - see collagen.
marginal prostals - spicules supporting and projecting around the oscula (Hexactinellida).
megascleres - larger size of spicules forming the sponge skeleton; also known as structural spicules (cf.
microscleres).
mesohyl - intercellular compartment in sponges, equivalent to the mesenchyme of other metazoans,
including the region bounded by the pinacoderm and choanoderm.
microcionid - condition of skeletal architecture found in encrusting sponges, in which nonanastomosing (plumose) spongin fibre nodes arise from a basal layer of spongin lying on the
substrate, and these are echinated by (usually perpendicular) plumose tracts of spicules.
microscleres - smaller category of spicules forming the sponge skeleton; also known as flesh spicules
(cf. megascleres).
microvilli - extensions of the choanocyte cell which form an upright tube or collar surrounding the
central flagellum, and used as in food entrapment and water flow.
monactinal - a spicule with 1 ray, growing from 1 end only, usually asymmetrical in geometry; monacts
include monaxonic spicules (e.g. styles in demosponges), derivatives of tetraxonic spicules (e.g.
"needle-eye" microxea in Calcarea) and hexactinal spicules (e.g. basal bidentates); (cf.
diactinal).
monaxonid - linear spicule with no more than 2 rays along 1 axis, including both monactinal and
diactinal spicules; also known as monaxonic (cf. triaxonid, tetraxonid).
multispicular - more than 1 row of spicules in a fibre, tract or reticulation; also known as polyspicular
(cf. unispicular).
oscula - exhalant pores, through which water leaves the sponge; usually represented as the larger pores
(cf. ostia).
ostia - inhalant pores, through which water enters the sponge (cf. oscula).
ovipary - method of sexual reproduction in which eggs develop within the female sponge, are broadcast
into the water, often becoming attached to a mucous layer on the external surface, they are
fertilized externally, larvae are subsequently released, are free for varying periods and
eventually settle on the substrate (cf. vivipary).
parenchymal - associated with the interior of the skeletal framework (trabeculae) (Hexactinellida).
parenchymella larva - larval form associated with viviparous development, with an even covering of
minute cilia or flagella, often with considerable cytological differentiation including juvenile
spicules.
pentact - radiate spicule with 5 rays (found in most Hexactinellida, but also occurring in some
demosponges as modified calthrops).
peripheral skeleton - see ectosome.
pinacocyte - outer layer of highly flattened anucleolate epithelial cell forming the external
(exopinacocyte) and basal (basopinacocyte) surfaces.
pinacoderm - outer (ectosomal) layer of the sponge formed by pinacocytes (Demospongiae, Calcarea)
or syncytial "tissue" (Hexactinellida).
pith - central diffuse region within a spongin fibre.
pleural prostals - spicules projecting laterally from the sponge (Hexactinellida).
plumoreticulate - condition of skeletal architecture whereby main tracts of spicules and/or fibres
diverge in plumose fashion and secondary tracts are interconnected.
plumose - condition of skeletal architecture in which spicules and/or fibres diverge, usually in ascending
tracts with the points of spicules projecting outwards, but do not rejoin.
polyact - spicule with more than 6 rays (microscleres).
Porifera - Phylum of animals, known collectively as sponges, in the subkingdom Parazoa, lying between
the single-celled (Protozoa) and multicellular animals (Metazoa).
porocyte - modified pinacocyte cell lining the pores of the inhalant and exhalant canal systems.
primary fibre or tract - major fibre or spicule tract usually ascending to the surface at right angles (cf.
secondary).
principal spicules - old term referring to the largest or main category of structural megascleres often
(but not exclusively) found within (coring) fibres or primary tracts (cf. accessory, auxiliary
spicules).
prosopyle - an aperture through which water enters the choanocyte chamber.
prosodal - a condition of the aquiferous system in which a small canal joins the choanocyte chambers to
an inhalant canal (cf. aphodal).
prostals - projecting spicules (see pleural and marginal prostals) (Hexactinellida).
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 welldifferentiated 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
15 club-shaped acanthostrongyle
16 cladotylote
Monactinal (monaxonic) Megascleres
17 style
18 hastate style
19 anisostyle
20 subtylostyle
21 tylostyle
22 acanthostyle
23 stellate acanthotylostyle
24 acanthostyle with clavulate spines
25 acanthostyle with recurved spines
26 rhabdostyle
27 verticillate acanthostyle
28 sagittal triact (acanthoplagiotriaene)
Calthrops and Derivatives
29 undifferentiated calthrops
30 tetrapod calthrops
31 centrangulate diact
32 monoloph (lophotetractine)
33 candelabrum
34 amphimesodichotriaene
Tetractinal (tetraxonic) Megascleres
35 short shaft triaene
36 plagiotriaene
37 anatriaene
38 protriaene
39 promonaene
40 orthotriaene
41 dichotriaene
42 trichotriaene
43 trichodal or heterocladal protriaene
44 discotriaene
45 phyllotriaene
46 oxytylote
Desmas
47 ophirhabd (monocrepidial);
48,49,50 dendroclones
51 tripodal dicranoclone (tricrepidial)
52,53 rhizoclones (monocrepidial)
54 rhizoclone with spinose zygomes;
55 heloclone (monocrepidial)
56 irregular heloclone
57 megaclone (monocrepidial)
58 tetraclone (tetracrepidial)
59 rhabocrepid (monocrepidial)
60 sphaeroclone (anacrepidial)
61 typical sphaeroclone
62 modified sphaeroclone (astroclone)
63 hypersilicified sphaeroclone
Collagenous Spongin Spicules
64 tylotiform
65 triaeniform
FIGS 66 - 145:
Geometric forms of
microscleres
and
gemmoscleres
in
Demospongiae
Meniscoid Microscleres
66 palmate isochela
67 arcuate isochela
68 anchorate isochela
69 unguiferous isochela
70 birotulate isochela
71,72 bipocilli
73 palmate anisochela
74 placochela
75 sphaerancora
76 canonochela
77 clavidisc (compound diancistra)
111
78 diancistra
79 c-sigma
80 s-sigma
81 serrate sigma
82 croca
83 centrangulate sigma
84 tetrapocilla
85 cleistochela
86 spined isancora
87 spined chela
88 tuberculate isochela
Monaxonic Microscleres
89 toxa
90 spined toxa
91 forceps
92 discorhabd (Didiscus)
93 oxydiscorhabd
94 dentate "sigmata"
95 sanidastoid discorhabd (Latrunculia)
96 sanidastoid discorhabd (Sigmosceptrella)
97 anisodiscorhabd
98 young sigmodiscorhabd
99 spiraster
100 spinispira
101 spirula
102 toxaspire
103 sigmaspire
104 selenaster
105 microxea
106 microstrongyle
107a centrotylote microxea
107b centrotylote microstyle (microtylostyle)
108 comma
109 raphide
110 trichite (bundle of raphides)
111 onychaete
112 spined centrotylote rod
113 spear-shaped microstyle
114 thraustoxea
115 sanidaster
116 "eailles" (monocrepidial disc)
Astrose Microscleres
117 plesiaster streptaster
118 amphiaster streptaster
119 metaster streptaster
120 spiraster streptaster
121 oxyaster euaster
122 oxyaster
123 oxyspheraster euaster
124 pycnastereuaster
125 strongylaster euaster
126 tylaster euaster
127 anthaster euaster
128 anthospheraster euaster
129 sterrospheraster euaster
130 sterraster euaster
131 aspidaster euaster
Gemmoscleres
132-136
acanthoxeas
and
transitional
pseudospherasters
137,138 acanthoxea
139 gemmule amphidisc or birotule
140 gemmule amphidisc
141 acanthoxeote
142-144 pseudasters
145 gemmule amphidisc
CALCAREA
FIGS 147 - 182:

Geometric forms of
calcareous spicules.
147 triradiate symmetrical rays
148 tripod
149 sagittal triradiate paired rays
150 triradiate curved ray
151 triradiate fused rays
152 triradiate sinuous rays
153 triradiate spined rays
154 triradiate vestigial rays
155 sagittal triradiate
156 triradiate tuberculate rays
157 sagittal quadriradiate
158 quadriradiate
159 sagittal quadriradiate with paired rays
160 sagittal quadriradiate equal rays
161 quadriradiate cladote rays
162 quadriradiate of apopyle
163 acanthose quadriradiate
164 - 174 calcitic oxeas ("monacts")
175 - 176 trichoxeas
177 - 178 needle-eye microxeas
179 diapason or 'tuning fork' spicule
180 tuning fork with paired rays
181 sinuous oxea
182 irregular tuning fork spicule
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
220 amphidisc
221 abnormal macramphidisc
222 onychohexaster
223 spherical discohexaster (discospiraster)
224 strobiloplumicome
Megascleres
225 part of dictyonal mesh with two lychniscs
joined
226 young lychnisc before inclusion in
framework
Microscleres
227 centrotylote rhabd
228 sphere
229 ring "sigma"
230 hemioxyhexaster
113
12. INDEX TO EXTANT SPONGE GENERA AND FAMILIES

Acanthoclada ................................................68
[
Acanthodendrilla..........................................82
Acanthodoryx................................................49
[Acca] ............................................................73
Acanthoplakina ............................................24
[Astraxinellidae]............................................31
Acanthorhabdus ...........................................41
[Aulocystis] .................................................101
Acanthostrongylophora ...............................75
[Choanitidae].................................................33
Acanthostylotela .........................................103
[Corticella] ...................................................27
Acanthotetilla................................................25
[Crambidae] .................................................51
Acanthotriaena .............................................28
[Cribrella].....................................................52
Acanthoxa......................................................53
[Damira] .......................................................41
Acanthoxifer..................................................69
[Desmacidinae]..............................................54
Acarneae.........................................................40
[Desmacidonidae]..........................................54
Acarnia ..........................................................43
[Djeddeidae] ..................................................91
Acarnidae .......................................................40
[Ectyonidae]...................................................39
ACARNIDAE...............................................40
[Ectyoninae]...................................................42
Acarniidae......................................................40
[Epipolasidae]................................................26
Acarnus..........................................................41
[Halinidae ......................................................28
Acervochalina ...............................................72
[Halme] .........................................................44
Acheliderma ..................................................41
[Jereopsis].....................................................37
Achinoe..........................................................24
[Lepidospongida] ......................................105
Achramorpha ................................................95
[Neocoela] ....................................................39
Aciculites .......................................................26
[Neocoeliidae]...............................................39
Acoelocalyx .................................................101
[Nepheliospongiidae]....................................75
Adocia............................................................72
[Plocamia] ..............................................43, 46
Adociidae.......................................................71
[Podospongiidae] ..........................................31
Adreissa .......................................................103
[Polyrhabdus] ............................................105
Adreus............................................................31
[Raspelia] .....................................................46
Aegogropila ..................................................62
[Scantilla]......................................................30
Agelas ............................................................40
[Spirophorellinae] .........................................36
AGELASIDA ...............................................39
[Spongeliidae]................................................81
AGELASIDAE ............................................39
[Streptasteridae].............................................26
Aiolochroia ...................................................84
[Syconidae] ....................................................90
Aka .................................................................73
[Tetracladidae]...............................................36
Alcyoncellum...............................................103
[Verongiidae].................................................83
Alcyospongia.................................................31
Alebion...........................................................42
Alectona.........................................................29
Cornulacantha...........................................42
Alemo .............................................................35
LITHISTIDA .............................................36
Algol...............................................................26
A
Allantella .....................................................103
Allantophora.................................................60
Aaaba.............................................................52
Allocia............................................................43
Aaata..............................................................43
Alloscleria
.....................................................71
Aaptos ............................................................34
Ammoconia..................................................103
Abila.........................................................43, 46
Ammolynthus...............................................103
Abilana ..........................................................46
Amniscos........................................................35
Abyssocladia .................................................49
Amoibodictya ................................................48
Acalle .............................................................76
Amorphilla ....................................................70
Acamas...........................................................62
Amorphina...............................................70, 72
Acamasina.....................................................62
Amorphinopsis ..............................................70
Acanthacarnus ..............................................41
Amphiastrella................................................54
Acanthancora................................................53
Amphibleptula...............................................26
Acanthascinae ..............................................102
AMPHIDISCOPHORA .............................97
Acanthascus ................................................102
Amphilectus...................................................54
Acanthella .....................................................63
Amphimedon .................................................73
Acanthellina ..................................................67
Amphinomia ..................................................45
Acantheurypon..............................................46
Amphitethya ..................................................25
Acanthochaetetes..........................................33
Amphiute........................................................92
Acanthochaetetidae........................................33
Amphoricortis ...............................................93
Acanthocinachyra.........................................25
Amphoridium ..............................................103
Amphoriidandra ...........................................93
Amphoriscandra ...........................................93
AMPHORISCIDAE....................................94
Amphoriscus..................................................94
Amphoriscyssa ..............................................94
Amphoriulandra ...........................................93
Anaata ...........................................................43
Anacanthaea .................................................69
Anaderma ......................................................37
Anamixilla .....................................................92
Anaulosoma ................................................102
Anchinoe........................................................48
ANCHINOIDAE..........................................48
Ancorella .....................................................103
Ancorina ........................................................26
ANCORINIDAE..........................................26
Anheteromeyenia ..........................................75
Anisocrella ....................................................51
Anisotylacanthaea ........................................41
Anisoxya ........................................................71
Annandalia..................................................103
Anomochone................................................100
Anomoclathria ..............................................43
Anomodoryx ..................................................49
Anomoianthella ............................................84
Anomolissa....................................................48
Anomomycale................................................61
Anomomyxilla ...............................................57
Anoplina........................................................38
Anoxycalyx...................................................102
Anthastra .......................................................27
Antherochalina .............................................43
Antheroplax...................................................81
Antho..............................................................43
Antho (Acarnia) ............................................43
Antho (Antho)................................................43
Antho (Isopenectya)......................................43
Anthoarcuata ................................................43
Anthosigmella ...............................................33
Apatospongia................................................70
Aphorme ......................................................102
Aphrocallistes ...............................................99
APHROCALLISTIDAE ............................99
Aphrocerandra..............................................93
Aphroceras ....................................................92
Aphrocerortis ................................................93
Aphroceryssa.................................................94
Aphrodite.......................................................77
Aplysia ...........................................................83
Aplysilla.........................................................82
Aplysillidae....................................................82
Aplysina.........................................................83
Aplysinella.....................................................84
Aplysinellidae ................................................83
APLYSINIDAE ...........................................83
Aplysinopsis ..................................................79
Aponastra ......................................................71
Arcesios........................................................103
Archaeocliona...............................................29
Arenochalina.................................................62
Arenosclera ...................................................73
Arndtanchora................................................53
Artemisina .....................................................43
Artynaltis .......................................................87
Artynandrium................................................93
Artynandrus...................................................92
Artynas...........................................................90
Artynella ........................................................90
Artynellandra................................................93
Artynes ...........................................................90
Artynium ........................................................90
Artynophyllum...............................................90
Artynortus......................................................93
Asbestopluma................................................59
Ascaltaga.......................................................89
Ascaltis.....................................................85, 87
Ascaltometra .................................................90
Ascaltopa.......................................................89
Ascandra........................................................87
Ascandraga ...................................................89
Ascandrometra..............................................90
Ascandropa ...................................................89
Ascetta............................................................85
Ascettaga .......................................................89
Ascettometra..................................................90
Ascettopa .......................................................89
Ascettusa........................................................89
Ascilla ............................................................89
Ascillaga........................................................89
Ascillopa........................................................89
Ascoleucetta ..................................................87
Ascometra......................................................89
Asconema.....................................................102
Ascortaga ......................................................89
Ascortopa ......................................................89
Ascortusa.......................................................89
Asculmis.........................................................89
Ascuris ...........................................................89
Ascute.............................................................89
Asteromeyenia...............................................75
Asteropus.......................................................26
Astromeyenia.................................................75
Astromimus..................................................103
Astropeplus....................................................26
ASTROPHORIDA......................................26
Astroplakina..................................................24
Astrosclera.....................................................40
ASTROSCLERIDAE .................................40
Astrospongia .................................................64
Astrostreptidae...............................................28
Astylinifer ......................................................41
Asychis ...........................................................72
Atergia ...........................................................32
Athnacama ....................................................61
Aulaxinia .......................................................36
Aulena......................................................44, 81
Aulenella........................................................43
Auletta............................................................63
AULOCALYCIDAE.................................100
Aulocalyx.....................................................100
Aulochone....................................................102
AULOCYSTIDAE.....................................100
Auloplax ......................................................100
Auloplegma ...................................................89
Auloplegmaltis ..............................................90
Auloplegmandra ...........................................90
Auloplegmetta ...............................................90
Auloplegmilla................................................90
Auloplegmortis..............................................90
Aulorhipis....................................................103
Aulorhiza .......................................................90
Aulorossella ................................................102
Aulorrhiza .....................................................87
Aulosaccus...................................................102
Aulospongiella..............................................45
Aulospongus..................................................45
Aurora............................................................27
Aurorella .......................................................27
Axechina ........................................................45
Axiamon.........................................................66
Axidragma.....................................................67
Axinectya .................................................46, 63
Axinella..........................................................64
AXINELLIDAE...........................................63
Axinomimus...................................................70
Axinosia.........................................................64
Axinyssa.........................................................70
Axocielita.......................................................43
Axoniderma ...................................................59
Axoplocamia .................................................46
Axos................................................................31
Axosuberites ..................................................44
Azorica...........................................................38
AZORICIDAE .............................................38
B
Badiaga .......................................................103
Baeria ............................................................94
Baicalolepis ................................................103
Baicalopis .....................................................76
Baikalospongia ............................................76
Bajalidae ........................................................84
Bajalus...........................................................85
Barbozia ........................................................52
Basiectyon .....................................................46
Basta ..............................................................85
Bathydorus ..................................................102
Bathyxiphus...................................................99
Batzella..........................................................57
Bernatia.........................................................29
Bethia...........................................................103
Biemna...........................................................60
Biemnidae......................................................60
Biminia ..........................................................74
Bipocillopsis .................................................43
Bolosoma.....................................................101
Brachiaster....................................................28
Brondstedia...................................................43
Bubarinae.......................................................63
115
Bubaris ..........................................................64
Bubaropsis ....................................................64
Bursalina.....................................................103
Burtonanchora .............................................56
Burtonella .....................................................42
Burtonispongia.............................................54
Burtonitethya ................................................35
Burtonulla .....................................................86
C
Cacochalina................................................103
Cacospongia .................................................79
Calcabrina ....................................................28
CALCAREA ................................................85
CALCARONEA..........................................89
Calcepongia..................................................90
Calcifibrospongia ........................................72
CALCINEA..................................................85
Calcispongia.................................................90
Callipelta.......................................................37
Callisphaera ...............................................103
Callites.................................................. 69, 103
Callyspongia.................................................73
CALLYSPONGIIDAE...............................73
Calthropella..................................................27
CALTHROPELLIDAE..............................27
Calycosilva..................................................103
Calycosoma.................................................102
Calyptorete....................................................99
Calyx..............................................................74
Calyxadocia ..................................................73
Caminella ......................................................27
Caminus.........................................................27
Camptisocale ................................................49
Cantabrina....................................................45
Carmia...........................................................62
Carnleia.........................................................34
Carterella ......................................................75
Carterias......................................................103
Carteriospongia ...........................................77
Carterius........................................................75
Cartilospongia............................................103
Caulocalyx ..................................................101
Caulodiscus.................................................103
Caulophacella ............................................103
CAULOPHACIDAE.................................102
Caulophacus ...............................................103
Caulospongia ...............................................34
Cavochalina................................................103
Cellulophana ................................................31
CERACTINOMORPHA............................38
Ceraochalina ................................................73
Ceratodendron..............................................77
Ceratoporella................................................40
Ceratopsion...................................................45
Ceratopsis .....................................................45
Cerbaris.........................................................64
Cercidochela.................................................54
Cerelasma....................................................103
Cerelpemma ................................................103
Cerelpsamma ..............................................103
Chaetodoryx..................................................49
Chalaronema ................................................98
Chalina..........................................................72
Chalinella......................................................73
CHALINIDAE .............................................71
Chalinissa .....................................................64
Chalinodendron ...........................................72
Chalinopora..................................................73
Chalinopsilla ................................................74
Chalinopsis ...................................................74
Chalinoraphis.............................................103
Chalinula ......................................................72
Characella.....................................................28
Chaunangium .............................................101
Chaunoplectella .........................................101
Chelonaplysilla.............................................82
Chelotropaena ..............................................27
Chelotropella ................................................27
Chiastosia .....................................................53
Chondrilla.....................................................31
Chondrilladae ................................................30
Chondrillastra ..............................................31
CHONDRILLIDAE ....................................30
Chondrillina .................................................33
Chondrocladia..............................................59
Chondropsis..................................................57
Chondrosia ...................................................31
Chondrosiidae................................................30
Chonelasma ..................................................99
Choristida ......................................................26
Chrotella .......................................................25
Cinachyra......................................................25
Cinachyrella..................................................25
Ciocalapata ..................................................70
Ciocalypta .....................................................70
Cionanchora .................................................43
Cioxeamastia ................................................70
Cladochalina ................................................73
Cladocroce ....................................................72
CLADOPELTIDAE....................................37
Cladorhiza ....................................................59
CLADORHIZIDAE ....................................59
Cladothenea..................................................28
Clathria .........................................................43
Clathria (Axosuberites)................................44
Clathria (Clathria) .......................................43
Clathria (Dendrocia) ...................................44
Clathria (Isociella).......................................44
Clathria (Microciona)..................................43
Clathria (Thalysias) .....................................44
Clathria (Wilsonella) ...................................43
Clathriella .....................................................43
Clathriidae .....................................................42
Clathrina .......................................................85
CLATHRINIDA ..........................................85
CLATHRINIDAE........................................85
Clathriodendron...........................................46
Clathriopsamma ...........................................43
Clathrissa ......................................................48
Claviscopulia ................................................99
Clinolabis......................................................50
Cliona ............................................................29
CLIONIDAE ................................................29
Clionopsis .....................................................29
Cliothosa .......................................................29
Clistolynthaltis..............................................89
Clistolynthandra...........................................89
Clistolynthetta...............................................89
Clistolynthilla ...............................................89
Clistolynthus .................................................89
Clystolynthus.................................................89
Coelectys........................................................49
Coelocalypta .................................................71
Coelocarteria................................................49
Coelodischela ...............................................49
Coelosphaera................................................49
Coelosphaerella............................................41
Coelosphaericon...........................................49
COELOSPHAERIDAE .............................48
Coenostomandium........................................93
Coenostomandra ..........................................93
Coenostomella ..............................................93
Coenostomellium ..........................................93
Coenostomium ..............................................93
Coenostomortis.............................................93
Collinella.......................................................36
Collocalypta..................................................70
Colloclathria.................................................44
Collosclerophora..........................................57
Collosclerophoridae......................................57
Collospongelia ...........................................103
Collospongia ................................................80
Columnitis .....................................................35
Cometella ......................................................34
Compsocalyx.................................................98
Coniasterium.................................................90
Conorete ........................................................99
Coppatias ......................................................26
COPPATIIDAE...........................................26
Corallistes .....................................................37
CORALLISTIDAE .....................................37
Corbitella ....................................................101
Corbitellinae................................................101
Cornulella .....................................................41
Cornulidae .....................................................40
Cornulotrocha ..............................................41
Cornulum ......................................................41
Corticella.......................................................24
Corticellopsis................................................27
Corticiidae .....................................................24
Corticium.......................................................24
Cortispongilla...............................................76
Corvomeyenia ...............................................76
Corvospongilla .............................................75
Corybas .........................................................54
Corynonema..................................................98
Corythophora .............................................103
Coscinoderma...............................................77
Coscinonema.................................................98
Coscinospongiidae ........................................37
Costifer ..........................................................37
Cotylina .........................................................59
Crambe ..........................................................51
CRAMBIIDAE ............................................51
Craniella .......................................................25
Craniellopsis.................................................25
Crateromorpha ...........................................102
CRATICULARIIDAE ................................99
Crella.............................................................52
Crellancistra .................................................61
Crellastrina .................................................104
CRELLIDAE ...............................................51
Crellina..........................................................52
Crellinspira ...................................................52
Crellomima....................................................52
Crellomyxilla.................................................56
Crelloxea .......................................................52
Cribrochalina ...............................................73
Crinorhiza .....................................................59
Cryptax ........................................................104
CRYPTOCOELIIDAE...............................39
Cryptospongia ............................................104
Cryptosyringa ...............................................26
Cryptotethya................................................104
Cyamon..........................................................45
Cyathella .....................................................104
Cyathiscus .....................................................92
Cyliconema....................................................98
Cymbastela....................................................64
Cyrtaulon ....................................................100
Cystispongia ...............................................104
D
Dactylella ......................................................69
Dactylia .........................................................74
DACTYLOCALYCIDAE ........................101
Dactylocalyx................................................101
Dactylochalina ...........................................104
Dactylospongia.............................................77
Daedalopelta ..............................................104
Damiria .........................................................41
Damiriana .....................................................50
Damiriella .....................................................49
Damirina .......................................................42
Damiriopsis...................................................54
Damo ...........................................................104
Damonilla .....................................................52
Damoseni.......................................................43
Darwinella ....................................................82
DARWINELLIDAE....................................82
Darwinia .......................................................82
Dasychalina ..................................................73
Deanea.........................................................104
Delectona ....................................................104
DEMOSPONGIAE .....................................23
Dendoricella .................................................48
Dendoryx .......................................................56
Dendrilla .......................................................82
DENDROCERATIDA................................81
117
Dendropsis ....................................................69
Dendrospongia .............................................84
Dendroxea.....................................................72
Dendya...........................................................86
Densa.............................................................70
Dercitancorina............................................104
Dercitopsis ....................................................24
Dercitus .........................................................28
Dermatreton..................................................92
Desmacella....................................................60
DESMACELLIDAE ...................................60
Desmacidon...................................................54
Desmacodes.................................................104
DESMANTHIDAE......................................38
Desmanthus...................................................38
Desmapsamma..............................................54
Desmatiderma...............................................60
Desmophorida ...............................................36
Desmoxya ......................................................69
DESMOXYIDAE ........................................68
Diacarnus......................................................31
Diactinolopha...............................................24
Diapleura ....................................................101
Diapleuridae ................................................100
Diaretula .....................................................104
Diastra...........................................................27
Dictyaulus ...................................................102
Dictyocalyx..................................................102
DICTYOCERATIDA..................................76
Dictyociona ...................................................43
Dictyoclathria ...............................................43
Dictyocylindrus.......................................45, 46
Dictyodendrilla.............................................82
DICTYODENDRILLIDAE........................82
Dictyonella....................................................69
DICTYONELLIDAE..................................69
Didiscus.........................................................68
Diplacodium ...............................................104
Diplastrella ...................................................36
Diplodemia..................................................104
Dirrhopalum .................................................43
Discodermia..................................................36
Discodermiidae..............................................36
Discorhabdella .............................................51
Distomus......................................................104
Disyringia......................................................26
Ditela .............................................................78
Djeddea .........................................................91
Doconesthes ................................................104
Docosaccus .................................................101
Dolichacantha ........................................42, 53
Donatia..........................................................35
Donatiidae......................................................35
Donotella.......................................................30
Dorydermidae ................................................37
Dorypleres.....................................................26
Dosilia ...........................................................75
Dotona.................................................. 29, 104
Dotonella.....................................................104
Dragmacidon ................................................64
Dragmatella ..................................................64
Dragmatyle....................................................64
Dragmaxia ....................................................65
Druinella .......................................................84
DRUINELLIDAE........................................83
Drulia ............................................................76
Dunstervillia .................................................90
Duriella .........................................................80
Duseidea........................................................81
Duseideia.......................................................81
Dymnus........................................................104
Dyscliona.......................................................29
Dyseideia.......................................................81
Dysidea..........................................................81
DYSIDEIDAE..............................................81
Dysideopsis ...................................................80
Dysidia...........................................................81
Dysidicinia ....................................................79
Dysscaltella...................................................93
Dyssicarium...................................................93
Dyssiconella..................................................93
Dyssycandrella..............................................93
Dyssycandrium..............................................93
Dyssycandrus ................................................93
Dyssycarium..................................................93
Dyssycillus.....................................................94
Dyssyconella .................................................93
Dyssycortus ...................................................93
Dyssyculmus..................................................94
Dyssycum.......................................................93
Dyssycyssus ...................................................94
E
Ebnerella .......................................................92
Echinaxia ......................................................46
Echinochalina...............................................44
Echinochalina (Echinochalina)..................44
Echinochalina (Protophlitaspongia) .........44
Echinoclathria ..............................................44
Echinodictyum ..............................................45
Echinonema ..................................................44
Echinoplocamia............................................43
Echinospongia..............................................31
Echinostylinos...............................................55
Ecionemia......................................................26
Ectoforcepia..................................................50
Ectyobatzella...............................................104
Ectyodoryx.....................................................49
Ectyomyxilla..................................................56
Ectyon ............................................................40
Ectyonancora................................................55
Ectyonilla ......................................................45
Ectyonopsis .................................................104
Ectyoplasia....................................................46
Eilhardia .......................................................92
Emplocus .......................................................56
Endectyon......................................................46
Epallax ..........................................................31
Ephydatia ......................................................75
Epicles ...........................................................46
Epipolasis......................................................70
Erylidae..........................................................27
Erylus.............................................................27
Esperella........................................................62
Esperia...........................................................62
Esperiopsidae.................................................54
Esperiopsis....................................................62
Euastrosa .......................................................26
Euchalina ......................................................73
Euchalinopsis ...............................................72
Euchelipluma ................................................60
Eudictyon ....................................................103
Eudictyum....................................................103
Euhyalonema ................................................98
Eumastia........................................................70
Eunapius .......................................................75
Euplacella .....................................................74
Euplectella ..................................................101
EUPLECTELLIDAE ...............................101
Euplectellinae..............................................101
Eurete.............................................................99
EURETIDAE ...............................................99
Euricinia........................................................79
Euryades......................................................104
Euryphylle .....................................................29
Euryplegma .................................................100
Eurypon .........................................................46
Euryponidae...................................................44
Euryspongia..................................................81
Euspongia .....................................................78
Euspongilla...................................................75
Euthymus .....................................................104
Evenor..........................................................104
Evomerula .....................................................61
Exaxinata ......................................................59
F
Fangophilina ................................................25
Farrea............................................................99
FARREIDAE...............................................98
Fascaplysinopsis ..........................................80
Fasciospongia ..............................................80
Fasubera .....................................................104
Fenestraspongia ...........................................80
Ferrerhernandezia........................................53
Fibularia .......................................................61
Fibulia ...........................................................55
Ficulina .........................................................34
Fieldingia....................................................100
Filifera...........................................................79
Fisherispongia..............................................43
Fistula..........................................................104
Fistularia.......................................................83
Foliolina........................................................74
Folitispa ........................................................43
Fonteia ..........................................................41
Forcepia ........................................................50
Forcepina ......................................................50
Fusifer............................................................41
G
Gapoda..........................................................29
Gastrophanella.............................................38
Geelongia ......................................................81
Gelliodes........................................................73
Gellius............................................................72
Geodia ...........................................................27
Geodia (Cydonium) ......................................27
Geodia (Isops)...............................................27
Geodia (Sidonops)........................................27
GEODIIDAE................................................27
Geodinella.....................................................27
Gilchristia ...................................................104
Gomphostegia...............................................62
Goreauiella ...................................................40
Grantessa ......................................................91
Grantia ..........................................................92
GRANTIIDAE .............................................92
Grantilla........................................................91
Grantiopsis....................................................92
Grapelia ........................................................62
Grayax ...........................................................48
Grayella.........................................................52
Grella.............................................................52
Guancha ........................................................86
Guitarra.........................................................60
GUITARRIDAE ..........................................60
Guminida .......................................................30
Gummina .......................................................31
Gummineae....................................................30
Gumminidae ..................................................30
Gymnorete .....................................................99
Gymnorossella ............................................102
H
Haastia ..........................................................81
Habrodictyon ..............................................103
Haddonella ...................................................85
HADROMERIDA .......................................29
Halichoclona ................................................72
Halichondria.................................................70
HALICHONDRIDA....................................63
Halichondriella ......................................70, 72
HALICHONDRIIDAE ...............................70
Haliclona.......................................................72
Haliclonidae...................................................71
Haliclonissa ..................................................73
Halicnemia ....................................................68
Halicometes...................................................35
Halina..........................................................104
Halinastra ...................................................104
Halisarca.......................................................83
HALISARCIDAE........................................83
Halispongia ..................................................81
Hallmannia ...................................................47
Halmopsis......................................................81
Halyphysema...............................................104
Hamacantha..................................................61
HAMACANTHIDAE..................................61
119
Hamigera.......................................................48
HAPLOSCLERIDA. ..................................71
Hastatus.........................................................56
Helophloeina ................................................59
Hemectyon.....................................................46
Hemectyonilla ...............................................45
Hemiasterella................................................31
HEMIASTERELLIDAE ............................31
Hemigellius ...................................................73
Hemihaliclona ..............................................73
Hemimycale...................................................57
Hemitedania..................................................58
Herengeria ....................................................37
Hertwigia.....................................................102
Heterectya......................................................45
Heterochone................................................100
Heteroclathria...............................................43
Heterocliona .................................................30
Heterocornulum............................................41
Heteromeyenia ..............................................75
Heteronema...................................................80
Heteropegma.................................................87
Heterophymia..............................................104
Heteropia.......................................................91
HETEROPIIDAE........................................91
Heterorete......................................................99
Heterorotula..................................................75
Heterotella...................................................102
Heteroxya ......................................................68
Hexactinella ................................................100
HEXACTINELLIDA..................................97
HEXACTINOSIDA.....................................98
Hexadella ......................................................82
HEXASTEROPHORA...............................98
Hezekia ........................................................104
Hiattrochota..................................................55
Higginsia.......................................................69
Higginsiinae...................................................68
Hiltonus .........................................................50
Hippograntia ................................................92
Hippospongia ...............................................77
Hircinella ......................................................79
Hircinia .........................................................79
Hirciniadae.....................................................79
Hircinidae ......................................................79
Hispidopetra .................................................40
Histioderma...................................................49
Histoderma....................................................49
Histodermella................................................50
Histodermion ................................................50
Histodermopsis .............................................42
Holascella ...................................................101
Holascus ......................................................101
Holoplocamia ...............................................43
Holopsamma .................................................44
Holorodesmia ...................................... 42, 104
Holoxea .........................................................26
Homandra .....................................................90
Homaxinella..................................................65
Homoderma...................................................91
Homoeodictya ...............................................55
Homophymia.................................................37
HOMOSCLEROMORPHA ......................23
HOMOSCLEROPHORIDA......................24
Hoplakithara ................................................60
Hoplochalina ................................................73
Hozawaia ......................................................91
Hyalascus ....................................................102
Hyalocaulus ................................................104
Hyalonema ....................................................98
HYALONEMATIDAE ...............................98
Hyalostylus..................................................102
Hyattella........................................................77
Hymantho ......................................................43
Hymedesanisochela......................................42
Hymedesmia ..................................................53
HYMEDESMIIDAE ...................................52
Hymenamphiastra ........................................53
Hymenancora................................................55
Hymendectyon...............................................48
Hymeniacidon ...............................................70
Hymeniacidonidae.........................................70
Hymenotrocha...............................................55
Hymenotrochota ...........................................55
Hymeraphia.............................................43, 46
Hymerhabdia ................................................65
Hymesigmia...................................................53
Hymoxenia.....................................................53
Hypodictyon ..................................................91
Hypograntia..................................................92
Hypsispongia ................................................61
Hyrtios ...........................................................80
I
Ianthella ........................................................85
IANTHELLIDAE........................................84
Ichnodonax ...................................................49
Idomon...........................................................29
Igernella ........................................................83
INCERTAE SEDIS ...................................103
Inflatella ........................................................50
Ingallia ..........................................................42
Inodes ............................................................80
Iophon ...........................................................42
Iophoneae.......................................................40
Iophonopsis...................................................42
Iophonota......................................................42
Iotaota ...........................................................55
Iotroata..........................................................55
Iotrochopsamma...........................................55
Iotrochostyla .................................................55
Iotrochota......................................................55
Iouea ..............................................................37
Iphiteon .........................................................99
Ircinia ............................................................79
IRCINIIDAE ................................................79
Isociona .........................................................43
Isodictya ........................................................55
Isopenectya....................................................43
Isops...............................................................27
Isoraphiniidae ................................................37
J
Janulum .......................................................104
Jaspinae..........................................................26
Jaspis .............................................................26
Jasplakina ...................................................105
Jelissima ........................................................53
Jenkina ..........................................................93
Jereicopsis.....................................................38
Jereidae ..........................................................36
Jia...................................................................43
Joannella.....................................................105
Jones ..............................................................50
Joyeuxia.........................................................50
Julavis............................................................69
K
Kaira..............................................................25
Kalastrella.....................................................31
Kaliapsidae ....................................................36
Kaliapsis........................................................36
Kallispongia ...............................................105
Kallypilidion .................................................72
Kalykenteron.................................................45
Kaneohea ......................................................57
Kapnesolenia ................................................26
Katiba ............................................................72
Kebira ............................................................97
Kerasemna.....................................................61
Keratylum....................................................105
Ketosus ..........................................................65
Kieplitela .......................................................45
Kirkpatrickia.................................................48
Korotnewia....................................................84
Kowalewskyella ..........................................105
Kuarrhaphis..................................................94
L
Labacea .........................................................43
Labisophlita..................................................50
Lamellomorpha ............................................28
Laminospongia.............................................71
Lamontia .......................................................93
Lanuginella.................................................102
Lanuginellinae .............................................102
Lanugonychia .............................................102
Laocaetidae....................................................99
Laocoetis .......................................................99
Latrunculia ...................................................32
LATRUNCULIIDAE..................................31
Laxosuberella ...............................................34
Laxosuberites................................................34
Lefroyella ......................................................99
Leiodermatiidae.............................................38
Leiodermatium..............................................38
Leioplegma..................................................100
Leiosella ........................................................78
Lelapia...........................................................97
Lelapiella ......................................................89
LELAPIELLIDAE......................................88
LELAPIIDAE..............................................96
Lenculmis ......................................................94
Lendenfeldia .................................................78
Leoculmis ......................................................94
Lepidoleucon ................................................95
LEPIDOLEUCONIDAE............................95
Lepidosphaera ..............................................50
Leptochelastra ............................................105
Leptoclathria ................................................43
Leptolabis......................................................50
Leptonema.....................................................98
Leptophragmella ..........................................99
Leptosastra..................................................105
Leptosia .........................................................53
Leptosiopsis ..................................................51
Lessepsia .......................................................72
Leucaltaga ....................................................93
LEUCALTIDAE..........................................86
Leucaltis ........................................................87
Leucaltusa .....................................................87
Leucandra .....................................................93
Leucandraga.................................................93
Leucandrena .................................................92
Leucandrometra ...........................................91
Leucandropa.................................................90
Leucandrusa .................................................93
Leucania........................................................93
Leucaria ........................................................89
LEUCASCIDAE..........................................87
Leucascus ......................................................87
Leucelia .........................................................89
Leuceria.........................................................89
Leucetta .........................................................88
Leucettaga.....................................................94
LEUCETTIDAE..........................................87
Leucettusa .....................................................87
Leucilia..........................................................89
Leucilla..........................................................95
Leuciria .........................................................89
Leuclathrina..................................................87
Leucogypsia ..................................................92
Leucomalthe..................................................88
Leuconia........................................................93
Leuconiinae ...................................................92
Leucophloeus ................................................70
Leucopsacasidae..........................................101
LEUCOPSACASIDAE ............................101
Leucopsacidae .............................................101
Leucopsacinae .............................................101
Leucopsacus................................................101
Leucopsila .....................................................93
Leucopsis.......................................................85
Leucoria ........................................................89
Leucortis........................................................93
Leucortmetra.................................................93
Leucosolenia .................................................89
LEUCOSOLENIIDA..................................89
LEUCOSOLENIIDAE ...............................89
Leuculmis ......................................................94
121
Leucyssa ........................................................93
Leukartea ......................................................90
Levinella........................................................86
LEVINELLIDAE ........................................86
Lieberkuhnia.................................................74
Ligrota...........................................................43
Liosina...........................................................69
Lipastrotethya.............................................105
Lipostomaltis ................................................93
Lipostomandra .............................................93
Lipostomella .................................................94
Lipostomortis ................................................93
Lissodendoryx ...............................................50
Lissomyxilla ................................................105
Lissoplocamia...............................................43
Lissopocillon.................................................48
Litamena......................................................105
Litaspongia ...................................................43
Lithobactrum ..............................................105
Lithobubaris .................................................65
Lithochela .....................................................51
LITHONIDA. ...............................................95
Lithoplocamia...............................................46
Lithospongia...............................................105
Lonchiphora .................................................99
Lophacanthus ...............................................38
Lophocalyx..................................................102
Lophophysema..............................................98
Lovenia ..........................................................32
Lubomirskia ..................................................76
LUBOMIRSKIIDAE ..................................76
Luffaria....................................................80, 83
Luffariella .....................................................80
Luffariospongia............................................77
LYCHNISCOSIDA...................................100
Lycopodina ...................................................59
Lydium ...........................................................37
Lyncuria ........................................................35
LYSSACINOSIDA....................................101
M
Macandrewia ................................................37
Malacosaccus .............................................101
Manawa.........................................................50
Manon .........................................................105
Marcusoldia..................................................50
Margaritella..................................................99
Marleyia ........................................................43
Mauricea .......................................................78
Medon............................................................93
Megaciella.....................................................42
Megalopastas................................................82
Megalopstas..................................................81
Megapodon ...................................................95
Meliiderma ....................................................59
Mellonympha ..............................................102
Melonanchora ..............................................56
Melonchela....................................................41
Melophlus......................................................27
Menanetia ...............................................70, 72
Menyllus ........................................................42
Merlia ............................................................61
Merriamium ..................................................48
Mesapos.........................................................46
Metania .........................................................76
METANIIDAE.............................................76
Metrastrosa ....................................................28
Meyenia .........................................................75
Meyerina........................................................97
MICROCIONIDAE.....................................42
Microcionina ................................................40
Microcordyla ................................................34
Microscleroderma ........................................26
Microstylifer..................................................65
Microtylostylifer ...........................................61
Microtylotella ...............................................41
Microxina ......................................................73
Microxistyla ......................................... 69, 105
Migas .............................................................70
Milene ............................................................70
Minchinella ...................................................96
MINCHINELLIDAE ..................................95
Mlea ...............................................................93
Monanchora .................................................51
Monanthus ....................................................71
Monectyon.....................................................46
Monocrepidium ............................................65
Monoplectroninia.........................................96
Monoraphis...................................................98
MONORHAPHIDIDAE.............................97
Monosyringa.................................................27
Monotria .....................................................105
Murrayona ....................................................88
MURRAYONIDA .......................................88
MURRAYONIDAE.....................................88
Mycale ...........................................................62
Mycale (Acamasina).....................................62
Mycale (Aegogropila) ..................................62
Mycale (Carmia)...........................................62
Mycale (Grapelia) ........................................62
Mycale (Mycale) ...........................................62
Mycalecarmia ...............................................62
MYCALIDAE..............................................61
Mycalina .......................................................59
Mycalopsis ..................................................105
Myceliospongia ............................................85
Myle ...............................................................29
Myliusia.........................................................99
Myriastra.......................................................27
Myrmekioderma............................................69
Myxichela ......................................................42
Myxilla...........................................................56
Myxilla (Burtonanchora).............................56
Myxilla (Ectyomyxilla) .................................56
Myxilla (Myxilla) ..........................................56
Myxilla (Stelodoryx).....................................56
Myxilla (Styloptilon) ....................................56
MYXILLIDAE.............................................53
Myxillina........................................................47
Myxodoryx.....................................................48
N
Naauna ..........................................................50
Nailondra ......................................................71
Nailondria...................................................105
Naniupi..........................................................52
Nara ...............................................................72
Nardoma........................................................89
Nardomandrum ............................................90
Nardopsandra...............................................90
Nardopsetta...................................................90
Nardopsis ......................................................89
Nardopsortis .................................................90
Nardoraltis....................................................90
Nardorandra.................................................90
Nardoretta.....................................................90
Nardorilla .....................................................90
Nardoris ........................................................90
Nardorortis ...................................................90
Nardorus .......................................................89
Nardosa .........................................................89
Naviculina .....................................................43
Negombata ....................................................32
Negombo .......................................................32
Neoadocia .....................................................72
Neoaulocystis..............................................101
Neocladia ......................................................59
Neoesperiopsis..............................................55
Neofibularia..................................................61
Neofolitispa...................................................51
Neopelta ........................................................37
NEOPELTIDAE..........................................37
Neopetrosia ...................................................75
Neoprosypa ...................................................69
Neosiphonia ..................................................37
Neothenea....................................................105
Nethea............................................................28
Niphates.........................................................73
NIPHATIDAE..............................................72
Nisella..........................................................105
Nisibaris ........................................................47
Normania ......................................................28
Noronha ........................................................61
Nudospongilla ..............................................75
O
Oceanapia .....................................................74
Oceanapiidae..................................................74
Ochridaspongia............................................76
Octavella .......................................................24
Okadaia.........................................................51
Oligoceras.....................................................80
Olynthaltus....................................................89
Olynthandra..................................................89
Olynthandrium..............................................89
Olynthandrus ................................................89
Olynthella......................................................89
Olynthelletta..................................................89
Olynthettus ....................................................89
Olynthillus.....................................................89
Olynthium......................................................89
Olynthortus ...................................................89
Olynthulmus ..................................................89
Olynthus ........................................................89
Ommatosa .....................................................64
Onconema .....................................................98
Onychomyxilla ..............................................56
Oonema .........................................................98
Oopsacas .....................................................105
Ophiraphidites............................................105
Ophistospongia ..........................................105
Ophlitaspongia.............................................44
Ophlitaspongiidae..........................................42
Orina..............................................................72
Orthorachis .................................................105
Oscarella .......................................................24
Oscarellidae ...................................................24
Oscaria ..........................................................24
Osculina ........................................................29
Oxeosarcodea .............................................105
Oxeostilon .....................................................71
Oxycarmia .....................................................62
Oxycordyla ....................................................34
Oxyheteromeyenia ........................................75
Oxylatrunculia ..............................................32
Oxymycale .....................................................62
Oxytedania ....................................................58
P
Pachamphilla................................................28
Pachastrella ..................................................28
PACHASTRELLIDAE ..............................27
Pachastrissa..................................................27
Pachataxa .....................................................27
Pachaxinella .................................................65
Pachychalina ................................................73
Pachydictyum................................................76
Pachymatisma...............................................27
Pachypellina .................................................74
Pachyrotula...................................................75
Pandaros .......................................................44
Papillella.......................................................29
Papillina........................................................29
Papillissa.......................................................29
Papyrula........................................................28
Paracordyla ..................................................26
Paracornulum...............................................42
Paradisconema .............................................98
Paradoryx......................................................43
Parafieldingia.............................................105
Paragrantia ..................................................92
Paraheteropia...............................................91
Parahigginsia ...............................................69
Paralelapia ...................................................97
Paraleucilla ..................................................93
Parameyenia .................................................76
Paramurrayona ............................................88
PARAMURRAYONIDAE.........................88
Paramyxilla...................................................50
Pararete.........................................................99
123
Pararhaphoxya.............................................65
Parasyringella ..............................................46
Paratedania ..................................................58
Paratenaciella ..............................................43
Paratetilla .....................................................25
Paratimea......................................................31
Paresperella..................................................62
Paresperia.....................................................56
Parisociella ...................................................62
Parmula.........................................................76
Patuloscula ...................................................73
Pectispongilla ...............................................75
Pellina ...........................................................74
Pellinula ......................................................105
Penares..........................................................27
Pericharax.....................................................88
Periphragella................................................99
Perissinella....................................................65
Petrobiona ....................................................96
PETROBIONIDAE.....................................96
Petromica ......................................................71
Petromicidae..................................................70
Petrosaspongia.............................................80
Petrosia .........................................................75
PETROSIIDAE ...........................................75
Petrostoma ....................................................96
Phacanthina............................................66, 69
Phakellia .......................................................66
Pharetronema .............................................105
Phelloderma..................................................50
Pheronema ....................................................97
PHERONEMATIDAE ...............................97
Phialonema ...................................................98
PHLOEODICTYIDAE...............................74
Phloeodictyon ...............................................74
Phlyctaenopora ............................................52
Phorbas .........................................................48
Phorbasidae....................................................48
Phoriospongia ..............................................57
PHORIOSPONGIIDAE.............................57
Phycopsis.......................................................66
Phyllospongia...............................................78
Phylosiphonia...............................................72
Phymaraphinidae ...........................................36
Phymatellinidae .............................................36
Physcaphora .................................................32
Pilochrota .....................................................27
Pione..............................................................30
Pitalia ............................................................43
Pityrete.........................................................105
Placinastrella................................................24
Placinolopha ................................................24
Placochalina.................................................74
Placoplegma ...............................................101
Placospongia ................................................32
PLACOSPONGIIDAE...............................32
Plakidium ......................................................38
Plakina ..........................................................24
Plakinastrella ...............................................24
PLAKINIDAE .............................................24
Plakoosa........................................................24
Plakortis ........................................................24
Platychalina............................................55, 73
Platylistrum...................................................97
Plectispa........................................................44
Plectrodendron .............................................34
Plectronella...................................................47
Plectroninia ..................................................96
Pleiomeyenia.................................................75
Pleorhabdus................................................105
Pleraplysilla..................................................82
Pleroma .........................................................37
PLEROMIDAE ...........................................37
Pleurochorium ..............................................99
Plicatella .......................................................66
Plicatellopsis.................................................66
Plocamiancora .............................................56
Plocamilla .....................................................43
Plocamione ...................................................46
Plocamionida................................................48
Plocamiopsis.................................................43
Plocamissa ....................................................56
Plumocolumella............................................55
Plumocolumetta............................................56
Plumohalichondria ......................................48
Pocilloguitarra .............................................60
Pocillon .........................................................42
Podospongia.................................................32
Podotuberculum ...........................................48
Poecillastra ...................................................28
Poecilochela..................................................53
POECILOSCLERIDA ...............................40
Polejna ..........................................................95
Poliopogon ...................................................97
Polyfibrospongia..........................................77
Polymastia.....................................................32
POLYMASTIIDAE.....................................32
Polytherses ....................................................79
Poritella.........................................................38
Porphyria ......................................................84
POTAMOLEPIDAE...................................76
Potamolepis ..................................................76
Potamophloios..............................................76
Poterion.........................................................34
Pozziella ........................................................61
Prianos ..........................................................75
Prionema .......................................................98
Pronax .....................................................29, 48
Pronaxella.....................................................48
Propachastrella ............................................28
Proraspailia..................................................46
Prostylissa.....................................................70
Prosuberites ..................................................34
Prosycum.......................................................89
Proteleia........................................................33
Protoclathria ..............................................105
Protoesperia..................................................62
Protophlitaspongia ......................................44
Protoraspailia ..............................................46
Protoschmidtia ...........................................105
Protosuberites...............................................34
Psammaplysilla.............................................84
Psammascus..................................................57
Psammastra...................................................27
Psammina....................................................105
Psammochela ................................................51
Psammocinia.................................................79
Psammoclemma ............................................57
Psammodoryx................................................58
Psammopemma.............................................57
Psammophyllium ........................................105
Psammotoxa..................................................57
Psetalia..........................................................25
Pseudanchinoe..............................................44
Pseudaxinella................................................66
Pseudaxinyssa...............................................70
Pseudobasta..................................................83
Pseudoceratina .............................................84
Pseudochalina ............................................105
Pseudoclathria..............................................52
Pseudoesperia...............................................62
Pseudohalichondria.....................................53
Pseudomyxilla...............................................56
Pseudosuberites............................................34
Pseudotrachya ..............................................66
Psilocalyx ....................................................100
Pteronema .....................................................98
Ptilocaulis .....................................................66
Ptychodesia ...................................................99
Pyloderma ...............................................50, 70
Pytheas ..........................................................52
Pytheilla ........................................................52
Pyxitis ............................................................27
Q
Qasimella ......................................................43
Quasillina......................................................33
Quepanetsal ................................................105
Querciclona...................................................66
Quindesmia ...................................................53
Quintoxilla ..................................................106
Quixilla........................................................106
Quizciona ......................................................43
R
Rachella.........................................................25
Racodiscula...................................................36
Radiella .........................................................33
Radiospongilla .............................................75
Ramoses.........................................................43
Ramosichela..................................................52
Raoa...............................................................59
Raphidotethya...............................................25
Raphioderma ................................................62
Raphiodesma ................................................62
Raphyrus .......................................................29
Raspaciona ...................................................46
Raspaigella ...................................................70
Raspailia .......................................................46
Raspailia (Clathriodendron).......................47
Raspailia (Hymeraphiopsis)........................47
Raspailia (Raspailia) ...................................47
Raspailia (Raspaxilla) .................................47
Raspailia (Syringella) ..................................47
RASPAILIIDAE..........................................44
Raspailopsis..................................................46
Raspaxilla .....................................................46
Raspeloplocamia ..........................................46
Regadrella...................................................102
Reidispongia .................................................37
Reniera ..........................................................72
Renieridae......................................................71
Reniochalina.................................................66
Rhabdastrella ...............................................27
Rhabderemia.................................................47
RHABDEREMIIDAE.................................47
Rhabdeurypon ..............................................47
Rhabdocalyptus ..........................................102
Rhabdodermella ...........................................94
Rhabdodictyum...........................................100
Rhabdodragma...........................................106
Rhabdopectella...........................................102
Rhabdoploca.................................................67
Rhabdosigma ................................................47
Rhabdosina...................................................37
Rhabdostauridium......................................106
Rhacodiscula ..............................................106
Rhaphidectyon ..............................................45
Rhaphidhistia .............................................106
Rhaphidophlus .............................................44
Rhaphidorus .................................................33
Rhaphidostyla...............................................71
Rhaphidotheca..............................................62
Rhaphisia ......................................................72
Rhaphoxiella.................................................71
Rhaphoxya ....................................................67
Rhizaxinella ..................................................34
Rhizochalina .................................................74
Rhizoniera .....................................................72
Rhopaloeides ................................................78
Ridleia .........................................................106
Rimella...........................................................36
Rinalda ..........................................................32
Roosa .............................................................24
Rossella .......................................................102
ROSSELLIDAE ........................................102
Rossellinae...................................................102
Rotuloplocamia ............................................56
S
Saccocalyx...................................................102
Samidae..........................................................25
Samus.............................................................25
Sanidastra .....................................................32
Sanidastrella.................................................26
Sapline...........................................................29
Sarcocornea ............................................57, 81
Sarcomella ..................................................106
Sarcotragus...................................................79
Sarostegia .....................................................99
125
Scantilletta ....................................................30
Sceptrella.................................................32, 62
Sceptrintus............................................ 33, 106
Sceptrospongia .............................................62
Schaudinnia ................................................102
Schmidtia.......................................................75
Schulzeviella .................................................97
Sclerilla .........................................................53
Scleritoderma................................................26
SCLERITODERMIDAE............................25
Sclerochalina ..............................................106
Scleroplegma ..............................................106
Sclerothamnopsis .......................................100
Sclerothamnus ............................................100
Scolymastra.................................................102
Scopalina ......................................................69
Scutastra......................................................106
Scypha ...........................................................90
Scyphia ........................................................106
Scyphidium..................................................102
Seliscothon ..................................................106
Semisuberites ................................................62
Semperella.....................................................97
Semperellidae ................................................97
Seriatula ........................................................70
Sericolophus .................................................97
Setidium.......................................................106
Sideroderma..................................................49
Siderodermella..............................................49
Sigmadocia ...................................................72
Sigmarotula ..................................................56
Sigmatella .....................................................57
Sigmatoxella .................................................60
Sigmaxia........................................................61
Sigmaxinella .................................................61
Sigmaxinellidae .............................................60
Sigmaxinyssa.................................................73
Sigmeurypon .................................................47
Sigmosceptrella ............................................32
Sigmotylotella ...............................................60
Siliquariospongia.........................................28
Simplicella.....................................................82
Siphonella .....................................................73
Siphonidiella...............................................106
Siphonidiidae.................................................37
Siphonidium..................................................38
Siphonocalypta.............................................67
Siphonochalina ............................................74
Siphonodictyon.............................................73
Siphydora ....................................................106
Skeizia............................................................67
Smenospongia...............................................80
Solenicaltis....................................................90
Solenicortis ...................................................90
Soleniculmis ..................................................90
Solenidandra ................................................90
Solenidium ....................................................89
Soleniscandra ...............................................90
Soleniscetta ...................................................89
SOLENISCIDAE.........................................86
Soleniscilla....................................................89
Soleniscus......................................................86
Soleniscyssa ..................................................89
Solenula.........................................................89
Solenulandra ................................................90
Solenuletta ....................................................90
Solina...........................................................106
Sollasella.......................................................26
Sollasellidae...................................................26
Somatispongia ............................................106
Sophax ...........................................................44
Spanioplon....................................................53
Sphaerotylus .................................................33
Spheciospongia ............................................30
Sphenophorina .............................................91
Sphenophorus ...............................................91
Sphincterella .................................................28
Sphinctrella...................................................28
Spinosella......................................................74
Spinularia .....................................................33
Spirasigmidae ................................................28
Spirastrella....................................................33
SPIRASTRELLIDAE.................................33
Spiretta ..........................................................25
Spirophora ....................................................36
Spirophorella................................................36
SPIROPHORIDA .......................................25
Spirorhabdia.................................................53
Spiroxya.......................................................106
Spongelia ......................................................81
Spongia .........................................................78
SPONGIIDAE..............................................77
Spongilla .......................................................75
Spongilla (Spongilla)...................................76
Spongilla (Stratospongilla).........................76
SPONGILLIDAE........................................75
Spongionella.................................................81
Spongocardium ..........................................106
Spongosorites ...............................................71
Spongosoritidae.............................................70
Spuma ............................................................70
Stannarium..................................................106
Stannoma ....................................................106
Stannophyllum............................................106
Staurocalyptus ............................................102
STAURORRHAPHIDAE ..........................95
Stegxella ........................................................56
Stelletta..........................................................27
Stellettidae .....................................................26
Stellettinopsis................................................26
Stelligera .......................................................31
Stellitethya.....................................................35
Stellogeodia ................................................106
Stelodoryx......................................................56
Stelospongia .................................................80
Stelospongos .................................................80
Stelospongus .................................................80
Stelotrochota.................................................56
Stematumenia................................................79
Stenospongia ................................................79
Sterrastrolepis...............................................76
Sterrastrosa ....................................................27
Stoeba ............................................................28
Strepsichordaia ............................................78
Streptoconus .................................................91
Stretastrosa ....................................................28
Stromatospongia ..........................................40
Strongylacidon .............................................58
Strongylamma...............................................58
Strongylodesma ............................................32
Strongylophora.............................................75
Stryphnus.......................................................27
Stylaxinella....................................................62
Stylinos ..........................................................62
Stylissa...........................................................67
Stylocordyla ..................................................34
STYLOCORDYLIDAE..............................33
Stylohalina ....................................................71
Styloptilon .....................................................56
Stylopus .........................................................53
Stylorhiza ......................................................34
Stylospira ....................................................106
Stylostichon...................................................48
Stylotella........................................................67
Stylotellopsis .................................................44
Stylotrichophora...........................................62
Suberanthus ..................................................34
Suberea..........................................................84
Suberella .......................................................34
Suberites........................................................34
SUBERITIDAE ...........................................34
Suberotelites..................................................48
Sulcastrella .................................................106
Swartschewskia.............................................76
Sycaltaga.......................................................90
Sycandra........................................................90
Sycandrarium................................................90
Sycandrometra..............................................91
Sycandrophyllum..........................................91
Sycantha ........................................................91
Sycarium........................................................90
Sycetta............................................................90
Sycettaga .......................................................90
SYCETTIDAE .............................................90
Sycettopa .......................................................90
Sycettusa........................................................91
Sycidium ........................................................90
Sycilla ............................................................94
Sycinula .........................................................93
Sycinulandra.................................................91
Sycissa ...........................................................90
Sycocarpus ....................................................90
Sycocerus.......................................................90
Sycocubus......................................................90
Sycocystis.......................................................90
Sycocystortis..................................................90
Sycodenandrum ............................................91
Sycodendrum.................................................90
Sycodoranna .................................................90
Sycodorilla ....................................................90
Sycodorus ......................................................90
Sycodorussa ..................................................90
Sycolepis........................................................94
Sycometra......................................................90
Sycon..............................................................90
Syconcystandra.............................................90
Syconella .......................................................90
Syconellandra ...............................................90
Syconellortis..................................................90
Sycophractus.................................................93
Sycophyllum ..................................................90
Sycorrhiza .....................................................89
Sycortarium...................................................90
Sycortis ..........................................................92
Sycortusa .......................................................92
Sycostrobus ...................................................90
Sycothamnandra...........................................91
Sycothamnetta...............................................91
Sycothamnus .................................................88
Syculmarium .................................................94
Syculmis.........................................................95
Sycum.............................................................90
Sycuraltis.......................................................90
Sycuranda .....................................................90
Sycurandra....................................................93
Sycuretta........................................................90
Sycurilla ........................................................94
Sycurortis ......................................................90
Sycurus ..........................................................90
Sycute.............................................................93
Sycyssa...........................................................93
Sycyssarium...................................................90
Sympagella..................................................103
Symplectella ................................................106
Sympyla .......................................................106
Synops............................................................27
Synute ............................................................93
Syringella ................................................34, 46
T
Tablis .............................................................44
Taboga ..........................................................35
Tabulospongia..............................................33
Tabulospongiidae ..........................................33
Taegeria ......................................................103
Taonura.........................................................81
Taprobane.....................................................26
Tarraltis.........................................................90
Tarrandra......................................................90
Tarretta..........................................................90
Tarrilla ..........................................................90
Tarroma.........................................................89
Tarromandra ................................................90
Tarropsandra................................................90
Tarropsetta....................................................90
Tarropsis .......................................................89
Tarrus ............................................................89
Technitella...................................................106
Tectitethya.....................................................35
Tedandoryx ...................................................57
127
Tedania..........................................................58
Tedania (Tedania)........................................58
Tedania (Tedaniopsis) .................................58
Tedania (Trachytedania) .............................58
Tedaniella .....................................................75
TEDANIIDAE..............................................58
Tedanione......................................................59
Tedaniopsamma ...........................................59
Tedaniopsis ...................................................58
Tedaniphorbas..............................................42
Teichaxinella.................................................67
Teichenopsis..................................................94
Teichonella....................................................88
Teichonopsis .................................................92
Tenacia ..........................................................44
Tenaciella......................................................44
Tenthrenodes.................................................91
Tentorina.......................................................33
Tentorium ......................................................33
Tereus ............................................................56
Terpios...........................................................34
Terpiosella ....................................................34
Tethya ............................................................35
Tethycometes.................................................35
Tethycordyla ...........................................34, 35
TETHYIDAE ...............................................35
Tethyophaena ...............................................34
Tethyopsilla...................................................25
Tethyopsis......................................................26
Tethyorrhaphis .............................................35
Tethyspira......................................................69
Tethytimea.....................................................35
Tetilla.............................................................25
TETILLIDAE...............................................25
Tetracladina ...................................................36
TETRACTINOMORPHA..........................24
Tetranthella...................................................51
Tetrapocillon ................................................60
Textiliforma...................................................55
Thalassodendron..........................................44
Thalassomora ...............................................26
Thalyseurypon ..............................................43
Thamnonema ................................................98
Thecometra....................................................89
Thecophora ...................................................33
Thenea ...........................................................28
THENEIDAE ...............................................28
Theonella.......................................................37
THEONELLIDAE ......................................36
Thieleia....................................................67, 71
Thielia............................................................70
Thoosa ...........................................................30
Thorecta ........................................................81
Thorectandra ................................................81
THORECTIDAE .........................................79
Thorectopsamma ..........................................80
Thrinacophora..............................................47
THROMBIDAE...........................................28
Thrombus ......................................................28
Thymosia .......................................................31
Thymosiopsis.................................................31
Timea .............................................................36
TIMEIDAE...................................................35
Tisrone...........................................................52
Topsentia.......................................................71
Toxadocia......................................................72
Toxemna ........................................................60
Toxiclona.......................................................72
Toxochalina ..................................................74
Trachostyla ...................................................71
Trachostylea..................................................53
Trachya..........................................................34
Trachycaulus...............................................102
TRACHYCLADIDAE................................36
Trachycladus.................................................36
Trachyforcepia..............................................50
Trachygellius ................................................29
Trachyopsilla ................................................70
Trachyopsis ...................................................71
Trachyspongilla............................................75
Trachytedania...............................................58
Trachytelia ....................................................34
Tragium .......................................................106
Tragosia ........................................................67
Trefortia ........................................................41
Tremaulidium..............................................106
Tretocalyx....................................................100
TRETODICTYIDAE ................................100
Tretodictyum ...............................................100
Tretolophus .................................................106
Tretopleura .................................................100
Tretorete ......................................................100
Triaenosa........................................................36
Triaenosina...................................................36
Tribrachion ...................................................26
Trichasterina...............................................102
Tricheurypon.................................................46
Trichogypsia .................................................94
Trichostemma................................................33
Trikentrion ....................................................47
Triptolemus ...................................................28
Trochoderma.................................................59
Trochospongilla ...........................................76
Trypespongia ................................................77
Tuba...............................................................74
Tubella...........................................................76
Tuberella .......................................................34
Tubulodigitatus ............................................73
Tubulodigitus..............................................106
Tulearinia......................................................96
Tumata...........................................................70
Tupha.............................................................75
Tylexocladus .................................................33
Tylodesma .....................................................60
Tylosigma ......................................................60
U
Uliczka.........................................................106
Ulosa..............................................................62
Umborotula...................................................76
Uncinatera ..................................................106
Uplexoa .........................................................68
Uraguaya ......................................................76
Uritaia ...........................................................71
Uritiaia ..........................................................70
Ute............................................................93, 94
Utella .............................................................90
Uteopsis.........................................................94
V
Vaceletia........................................................39
Vagocia .........................................................74
Valedictyum...................................................46
Valentis........................................................107
Vazella .........................................................107
Velinea...........................................................81
Veluspa ..........................................................72
Verongia........................................................83
VERONGIDA ..............................................83
Verongula......................................................83
VERTICILLITIDA......................................38
Verticillitidae.................................................39
Vetulina .........................................................38
VETULINIDAE...........................................38
Vibulinus .......................................................31
Viles ...............................................................68
Vioa................................................................29
Vitrollula .....................................................102
Volzia.............................................................30
Vomerula .......................................................61
Vosmaeria .....................................................92
Vosmaeropsis................................................91
Vulcanella .....................................................28
W
Waldoschmittia .............................................50
Walteria.......................................................102
Waltherarndtia .............................................68
Weberella.......................................................33
Wetmoreus.....................................................44
X
Xenospongia .................................................35
Xestospongia.................................................75
Xylospongia ................................................107
Xytopsaga......................................................55
Xytopsene ......................................................41
Xytopsihis ......................................................50
Xytopsoocha................................................107
Xytopsoocho..................................................49
Xytopsues.......................................................58
Xytosiphum....................................................58
Y
Yodomia.........................................................28
Ysila ...............................................................51
Ysiphonia.....................................................107
Yvesia.............................................................52
Yvesiella.........................................................52
Yvesiorbas .....................................................52
Z
Zetekispongia................................................50
Zetekopsis......................................................50
129
Zottea.............................................................50
Zygherpe........................................................61
Zygomycale ...................................................63
Zyzzya ............................................................42

Guide To Sponge Identification

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Guide To Sponge Identification

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SPONGUIDE.

GUIDE TO SPONGE COLLECTION AND IDENTIFICATION

3.2.11.2. Spicules. These are classified according to five major criteria:

19. hollow, several tubular digits attached to

6. soft, spongy, compressible/

7. membraneous, hispid, with erect spicules from

8. compressed, with more-or-less disorganized

fibre nodes, cored by plumose tufts of radiating

13. only primary (ascending) skeletal tracts present,

5. poorly developed, lightly invested with spongin,

4. heavily microspined bases/

(large, strongly recurved hooks) present on both

6. v-shaped toxas, sharp angular central curvature,

5. KEY TO THE EXTANT ORDERS OF PORIFERA.

Body plan syconoid or leuconoid ............................................... [Order Leucosoleniida, other

SCOPE: Two genera are recognised.

REVIEWS: Lendenfeld (1903).

(compressed, reticulate) and extra-axial skeletons (radial, plumose, plumo-reticulate);

composed of smooth ectosomal anisotornotes; choanosomal skeleton with hymedesmoid skeleton,

Myxilla (Ectyomyxilla) Lundbeck, 1909 (type species: Ectomyxilla kerguelensis Hentschel,

Tetrapocillon Brondsted, 1924 (type species: Tetrapocillon novaezealandiae Brondsted,

Strepsichordaia Bergquist, Ayling and Wilkinson, 1988 (type species: Strepsichordaia

q FAMILY DRUINELLIDAE LENDENFELD, 1889.

SCOPE: Fourteen genera are included.

Hexactinella Carter, 1885 (type species: )

Acanthostylotela Burton & Rao - with acanthose megascleres; possibly Myxillidae

Diplodemia Bowerbank, 1862 (type species: Diplodemia vesicula Bowerbank, 1862).

condensed or compressed - condition of skeletal architecture in which there is a compressed central

isotropic - condition of skeletal architecture in which there is a disoriented or seemingly random

FIGS 147 - 182:

12. INDEX TO EXTANT SPONGE GENERA AND FAMILIES

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