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REVISION OF MICROCIONIDAE (PORIFERA: POECILOSCLERIDA:
DEMOSPONGIAE), WITH DESCRIPTION OF AUSTRALIAN SPECIES.
JOHN N.A. HOOPER
Hooper, J.N.A. 1996 07 01: Revision of Microcionidae (Porifera: Poecilosclerida:
Demospongiae), with description of Australian species. Memoirs of the Queensland Museum
40: 1-626. Brisbane ISSN 0079-8835.
A phylogenetic revision of the poecilosclerid Microcionidae is based on type material, the
worldwide literature, and comprehensive Australian collections. Of 73 available generic
names 7 genera and 12 subgenera are recognised here. Of 561 available species names 459
are considered valid (10 virtually unrecognisable), including 52 new species. The Australian
fauna, including Australian Antarctic Territory, contains 148 species (31 new), many new
records, most are new combinations, and many illustrated for the first time. A synonymy of
world species is provided. Valid taxa include: Clathria (with 7 subgenera: C. (Clathria) (with
31 Australian species, 82 other species worldwide, with new species murphyi, noarlungae,biclathrata, borealis, burtoni , sarai, saraspinifera), C. (Wilsonella) (6, 8, abrolhosensis,
ensiae, lindgreni spp. nov.), C. (Microciona) (5, 91, illawarrae, lizardensis, simae,
brondstedi, campecheae, claudei, hen tscheli, leighensis, stephensae, tunisiae, urizae,
vacelettia spp. nov.), C. (Dendrocia) (7 species endemic to Australia), C. (Axociella) (6, 6,
patula , fromontae, georgiaensis spp. nov.), C. (Isociella) (4, 1, selachia, skia spp. nov.), and
C. (Thalysias) (36, 53, aphylla, craspedia, darwinensis, fusterna, hallmanni, hesperia,
lematolae, phorbasiformis, styloprothesis, tin gens, wesselensis, amiranteiensis, hechteli
spp. nov.); Antho (with 3 subgenera: A. (Antho) (12, 10), A. (Plocamia) (2, 17) and A.
(Isopeneclya) (3,1, punicea, saintvincenti spp. nov.); Echinoclathria (14, 15, bergquistae,
levii, notialis, parkeri, riddlei spp. nov.); Holopsamma (9 species endemic to Australia, 1
indeterminate species); Echinochalina (with 2 subgenera: E. (Echinochalina) (10, 2, felizi
sp. nov.), E. (Protophlitaspongia) (8 species endemic to Australia and New Caledonia,
collata, favulosa, isaaci, tuberosa spp. nov.)); Artemisina (4, 10); and Pandaros (incertae
sedis) (0, 2). Generic keys are provided. Morphometric characters of primary importance
include the origin, geometry and distribution of structural megascleres within the skeleton,
modification of megascleres to monactinal or diactinal forms, the presence or absence of a
specialised ectosomal skeleton, presence of detritus incorporated into spongin fibres, and
overall skeletal structure (including compression of the axial skeleton and differentiation of
axial and extra-axial regions). Brief zoogeographical comparisons are made between continental Australian and adjacent Indo-west Pacific faunas. Australian species comprise about
32% of the world's microcionid diversity; about 75% of species are endemic for the
Australian region, and temperate species (81%) have higher levels of endemism than tropical
species (59%). El Porifera, Demospongiae, Poecilosclerida, M icrocionidae, family revision,
new species, taxonomy, biogeography, Australia.
John N.A. Hooper, Queensland Museum, PO Box 3300, South Brisbane, Queensland, 4101,
Australia; received] December 1995.
INTRODUCTION
Microcionidae is one of the largest families of
Demospongiae, comprising about 8% of all
described (extant) Porifera species (Hooper &
Levi, 1993a). The family has contained at one
time or another about 70 genera and 550 species,
although fewer than these are now recognised as
valid. The family has a worldwide distribution
and it is found from the intertidal zone to depths
exceeding 2000m. It is clearly one of the more
important, ecologically successful groups of
Porifera.
Within the Indo-Australian region microcionids
are particularly abundant, with some species
being dominant components of the shallow water
macrobenthos. Previous works describing this
fauna (and other literature containing extra-limital records of Australian species) include:
Lamarck (1814, 1815, 1816), Gray (1858, 1867,
1869, 1870), Bowerbank (1864, 1875, 1877), Barnard (1879), Kent (1871), Ridley (1884a), Ridley
& Dendy (1886, 1887), Lendenfeld (1888, 1889a),
Kieschnick (1896, 1900), Thiele (1898, 1899,
1900, 1903), Kirkpatrick (1900a, b), Whitelegge,
(1901, 1902), Hentschel (1909, 1911, 1912),
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Hallmann (1912, 1914a-c, 1916a-c, 1920),
Dendy & Frederick (1924), Topsent (1897b,
1930, 1932, 1933), Burton (1934a), Bergquist &
Tizard (1967), Kelly-Borges & Bergquist (1988),
Bergquist & Fromont (1988), Wiedenmayer
(1989), Hooper (1990b), Hooper et al. (1991,
1992), Hooper & Levi (1993a, 1994). A brief
synopsis of the fauna is given by Hooper &
Wiedenmayer (1994), although some of the
nomenclature and synonymies contained in that
earlier work are revised here.
Prior to the present study more than 200 species
of Microcionidae had been described in the
Australian fauna (including its territorial waters),
but many of these were found to be either composite (consisting of several sibling species), or
synonyms of other species. This study 1)
describes 148 species (31 new), many new
locality records for Australia and new taxonomic
combinations; 2) provides an annotated
synonymy for 311 other species worldwide (including 21 new species); 3) revises the morphometric characters used for classification and
population variability for particular species; and
4) determines levels of endemism amongst
provincial faunas.
MATERIALS AND METHODS
COLLECTION AND HISTOLOGICAL TECHNIQUES. Material examined in this study was
predominantly collected using SCUBA (0-40m
depth) or dredging and trawls (30-360m depth).
Seasonal sampling for reproductive periodicity
was conducted over two years in the Darwin and
Cobourg Peninsula regions, NT. Immediately
after collection specimens were either fixed in
80-100% methylated ethanol or frozen (which to
some extent fixes the pigments), and later
preserved in 70% alcohol. Reproductive products
were searched for in fresh or frozen tissue.
Nitric acid spicule preparations, thick-sections
and thin-section mounts were routinely made as
follows. Fragments of each sponge, including
ectosomal and choanosomal regions, were heated
directly on a glass microscope slide in several
drops in nitric acid (the solution was evaporated
rather than boiled, using low heat), and mounted
in Canada balsam once completely dry, and
cooled. Thick, hand-cut sections were made perpendicular to the surface, soaked in a saturated
solution of phenol and xylene (for approximately
24 hours), and mounted in Durcupan (ACM
Fluka Products) using glass slivers or card to raise
the coverslip level. Phenol-xylene precluded the
necessity for dehydration through an alcohol
series. Some microtome sections cut at 30-35m
were made for each species. Fragments were
passed through a dehydration series, cleared in
toluene or Histosol, and wax embedded for at
least 2 hours. Sections were 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),
placed in clearing agent for an adequate period to
dissolve wax and/or dewaxing on a hot plate, then
soaked in ethanol until perfectly clear, floated
onto albumen-coated slides, orientated and flattened, stained with basic fuccsin and mounted.
Fragments of dry specimens (e.g., type material)
were reconstituted in 5% buffered fonnalin for 12
hours, which produced rehydration of the
mesohyl and enabled cleaner histological sections to be made.
MORPHOMETRIC ANALYSES. Spicu les were
measured with a stage micrometer, either directly
through a microscope or computer digitiser.
Twenty five spicules, of each spicule category, in
all specimens were measured. Acanthostyle
width measurements were taken immediately
below the base. Toxa lengths refer to chord
length; isochelae are measured from apex of alae;
width measurements of other spicules refer to
maximum width.
Spicule dimensions were sorted and statistically compared for various parameters (e.g., season,
locality, depth), including one- and two-way
ANOVs with replication, two-way ANOV with
unequal replication, means differentiated using
two-tailed t tests. Line-drawings were made using
a calibrated camera lucida, and microphotographs
were taken with an Olympus microphoto system.
Taxonomic keys were constructed using ordered
multistate, disordered multistate, morphometric
and binary characters, utilising the DELTA system (Dallwitz & Paine, 1986).
SCANNING ELECTRON MICROSCOPY. Sections were prepared as follows:
1) Cut at 1-1.5mm thick, ensuring that both the
ectosome and choanosome were represented.
2) Placed in a cavity block and covered with
several drops of sodium hypochlorite to etch the
mesohyl matrix from the skeleton. The etching
process was monitored through a dissecting
microscope in order to prevent the skeleton falling apart. Delicate structures (plumose,
halichondroid, hymedesmoid skeletons) only required a few seconds treatment with bleach;
robust skeletons (reticulate, fibrous, articulated
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skeletons) required several minutes; but generally 30 seconds was adequate.
3) Bleach was pipetted off at the appropriate
time and 70% ethanol immediately added. Sections were left to stand for several minutes to
ensure bleach was completely neutralised.
4) Steps 2-3 were repeated, without removing
section from cavity block, substituting concentrated hydrogen peroxide in place of sodium
hypochlorite, finally rinsing in ethanol. The
hydrogen peroxide step was omitted for very
delicate sections.
5) Sections were placed on clean microscope
slides and let dry completely.
6) Sections mounted on SEM stubs using
double-sided tape, copper dag, or `Supa Glue'
(Supa Glue, Selleys Chemical Company,
Padstow). An alternative method used to fix
samples to stubs was to cover stub with
`Aquadhere' wood glue (Aquadhere, Selleys
Chemical Company, Padstow), let dry completely (usually several days), then prior to use exposed dry glue to vigorous steam (which softened
the set glue), and placed the section on top of the
stub (it would sink in a short way but was bonded
reasonably well to the stub, and had the advantage
of producing a perfectly smooth background).
7) The stub was sputter-coated well to ensure
that all fibres were well coated to reduce
'charging'. In some cases uncoated sections
could be viewed successfully under low accelerator voltage, but better results were generally obtained on coated specimens at higher voltage.
Typical viewing conditions used were 25kV, at
close working distance to provide best depth of
field and focus, and at low magnifications.
Spicule were prepared as follows:
1) Thinly cut sections including both ectosome
and choanosome were placed in a durham tube
(micro-test tube), to which drops of concentrated
nitric acid are added, using drop-by-drop addition
so as to control the oxidation reaction and production of by-product oxides.
2) Upon completion of acid digestion the durham tube was half filled with acid and gently
heated over an alcohol flame, ensuring that only
small bubbles form (low heat, no boiling), for 1-2
minutes.
3) Solution was let stand to cool, then
centrifuged (approximately 4000rpm for 30
seconds).
4) Nitric acid was pipetted off leaving a spicule
mass at the bottom of the tube, undisturbed.
5) Spicules were resuspended in fresh nitric
acid and gently stirred using clean, fine, glass rod.
6) These steps were repeated if any collagen
remained.
7) Spicules were resuspended firstly in
demineralised water, 70% ethanol, then two
series of 100% ethanol solutions, centrifuging
and decanting the supernatant between each
change of solution, finally ending with suspended
spicules in a solution of absolute ethanol.
8) A micro-cover glass was adhered to an SEM
stub using double-sided tap or copper dag, several
drops of suspended spicules placed onto the cover
glass, the alcohol-spicule solution ignited and
spread across the glass with a glass rod or forceps
until all ethanol was vaporised. Spicules bond to
glass relatively firmly, but excess spicules could
be blown off glass using compressed air, or
spread out over the glass by adding further
ethanol and igniting. The distribution of spicules
on the cover glass was monitored under compound or dissecting microscope (magnification
depending on spicule size). More drops of spicule
solution added and this step repeated if too few
spicules were present, ensuring not to overcrowd
field of view for SEM photographic purposes.
9) An alternative method was used to produce
a perfectly smooth background, using an
`Aquadhere' glue-coated stub, dried for several
days then softened with steam, and spicules
placed directly onto soft glue (in this case ethanol
was not burnt but evaporated). Single spicules
would sink into glue too far if it was too soft (i.e.,
left in steam too long).
10) Spicule coated stubs were sputter coated
briefly and viewed at 25kV, minimum working
distance and smallest apperture for best resolution.
ABBREVIATIONS
AAT, Australian Antarctic Territories; ABIP,
Centro de Estudios Avanzados de Blanes, Institut° de Investigaciones Pesqueras Barcelona,
Aquarium de Blanes, Gerona; ABRS, Australian
Biological Resources Survey, Canberra; AFZ,
Australian Fishing Zone; AHF, Alan Hancock
Foundation, University of Southern California,
Los Angeles; AIMS, Australian Institute of
Marine Science, Townsville; AM, Australian
Museum, Sydney; AMNH, American Museum
of Natural History, New York; BMNH, The
Natural History Museum, London; BPBM, Bernice P. Bishop Museum, Honolulu; CP, Cobourg
Peninsula, NT; CPMNP, Cobourg Peninsula
Marine National Park, NT; CSIRO, Commonwealth Scientific and Industrial Research Or-
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ganisation, Marine Laboratories, Hobart,
Cleveland and Perth; DAR, Darwin region, NT;
DELTA, Description Language for Taxonomy
computer software (Dallwitz & Paine, 1986);
EIS, Environmental Impact Study; CSIROEMG,
CSIRO Food Research Laboratory, Division of
Food Processing, North Ryde, Sydney. EMU,
Ensight (formerly Environmental Management
Unit), Water Board (Sydney, Illawarra, Blue
Mountains), Sydney; EPA, Environment Protection Authority, Sydney; EPALR, East Point
Aquatic Life Reserve, Dudley Point, Darw;n Harbour, NT; FNQ, far northern Queensland (Cooktown to Torres Straits); FUB, Freie Universitat
Berlin; GBR, Great Barrier Reef, Queensland;
HNUK, Natural History Museum, Ham Nam
University, South Korea; ICBUC, Instituto
Central de Biologia, Universidad de Concepcion,
Chile; ICZN, International Code of Zoological
Nomenclature (see Anonymous, 1985); IM, Indian Museum (Zoological Survey of India), Calcutta; IMZUB, Istituto e Museo di Zoologia ed
Anatomia Comparata della Universita di Bari,
Bari; IMZUN, Instituto e Museo di Zoologia
dell'Universita di Napoli, Naples; INM, National
Museum of Ireland, Dublin; IZUG, Museo
Civico di Storia Naturale di Genova, Genova;
JCU, James Cook University of North
Queensland, Townsville; KFAU, Zoologischen
Sammlung der Universitat Erlangen-Niirnberg,
Erlangen; LFM, Merseyside County Museums
(formerly Liverpool Free Museum), Liverpool;
LMJG, Abteilung fiir Zoologie am Landesmuseum Joanneum (Landes Museum Jubileum
Graz), Graz; MABA, Museo Argentino de Ciencias Naturales 'Bernardino Rivadavia', Eaenos
Aires; MCNP, Div. Invest. del Museo de Ciencias
Naturales de la Plata, Argentina; MCZN,
Museum of Comparative Zoology, Harvard
University, Cambridge (Mass.); MEQ, mid eastern Queensland (Gladstone to Bowen); MHNG,
Museum d'Histoire NatureIle de Geneve,
Geneve; MLUM, Marine Laboratory of the
University of Miami, Miami; MMBS,
Mukaishima Marine Biological Station, Faculty
of Science, Hiroshima University, Onomichi;
MNHN, Museum National d'Histoire Naturelle,
Laboratoire de Biologie des Invertebres Marins
et Malacologie, Paris (DT, Topsent collections;
DCL, Levi collections; DJV, Vacelet collections;
DNBE, Boury-Esnault collections); MOM,
Musee Oceanographique de Monaco, Monaco;
MRAC, Koninklijk Museum voor MiddenAfrika, Tervuren; MRHN, Musee Royal d'Histoire Naturelle de Belgique, Bruxelles; MTQ,
Queensland Museum, Museum of Tropical
Queensland, Townsville; NCIQ66C-, United
States National Cancer Institute, Australian Institute of Marine Science shallow water collection contract (1984-91), Townsville (primary
voucher samples now lodged in QM, others in
NTM and USNM); NC1OCDN-, United States
National Cancer Institute, Coral Reef Research
Foundation shallow water collection contract,
Chuuk State (voucher samples lodged in QM and
USNM); NEQ, northeast Queensland (Bowen to
Cooktown); NM, Natal Museum, Pietermaritzburg; NMB, Naturhistorisches Museums
zu Basel, Basel; NMCIC, National Museum of
Natural Sciences, National Museums of Canada,
Ottawa; NMNZ, National Museum of New
Zealand (formerly Dominion Museum), Wellington; NMV, Museum of Victoria (formerly
National Museum of Victoria), Melbourne;
NSM, National Science Museum, Tokyo; NSW,
New South Wales; NT, Northern Territory; NTM,
Northern Territory Museum of Arts and Sciences,
Darwin; NTU, Northern Territory University,
Darwin; NWS, Northwest Shelf region, Western
Australia; PAUP, Phylogenetic Analysis Using
Parsimony (see Swofford, 1991); PIBOC, Pacific
Institute of Bio-organic Chemistry, Far East
Scientific Centre, Academy of Sciences of the
USSR, Vladivostok; PMJ, Phyletisches Museum,
Jena; PNG, Papua New Guinea; QFS,
Queensland Fisheries Service, Department of
Primary Industries, Brisbane and Cairns; QLD,
Queensland; QM, Queensland Museum, Brisbane; QVML, Queen Victoria Museum and Art
Gallery, Launceston; RMBS, Roscoff Marine
Biological Station, Roscoff, France; RMNH,
Rijksmuseum van Natuurlijke Historie, Leiden;
RRIMP, Roche Research Institute of Marine
Pharmacology, Sydney (discontinued; sponge
collections now held in AM); RSME, Royal Scottish Museum, Edinburgh; SA, South Australia;
SAM, South Australian Museum, Adelaide; SEQ,
southeast Queensland (Tweed River to
Gladstone); SM, Musee Zoologique, Strasbourg;
SME, Station Marine d'Endoume, Marseille;
SMF, Natur-Museum und Forschungsinstitut
Senckenberg, Frankfurt; TAS, Tasmania; TM,
Museo e Istituto di Zoologia Sistematica dell'Universita di Torino, Torino; TMAG, Tasmanian
Museum and Art Gallery, Hobart; UAZD,
University of Auckland, Zoology Department,
Auckland; UB, Ubersee-Museum, Bremen;
UCT, South African Museum of Natural History,
Cape Town; UQ, University of Queensland, Brisbane; USC, University of Southern California,
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Los Angeles; USNM, National Museum of
Natural History, Smithsonian Institution,
Washington DC; UZM, Zoologisk Museum,
Universitetsparken, Copenhagen; VIC, Victoria;
WA, Western Australia; WAM, Western
Australian Museum, Perth; YPM, Peabody
Museum of Natural History, Yale University,
New Haven (Conn.); ZIL, Zoological Institute of
Leningrad, Academy of Sciences Museum of
Zoology, St Petersburg; ZMA, ZoOlogisch
Museum, Universiteit van Amsterdam, Amsterdam; ZMB, Museum ftir Naturkunde an der
Humboldt-Universitat zu Berlin, Berlin; ZMC,
Zoologisk Museum, Copenhagen; ZMH,
Zoologisches Institute und Zoologisches
Museum der Universitat Hamburg, Hamburg;
ZMUU, Uppsala Universitets Zoologiska
Museet, Zoologiska Institutet, Uppsala; ZSN,
Aquarium e Museo della Stazione Zoologica di
Napoli, Naples; ZRS, Zoologiska Rijkmuseum,
Stockholm.
ACKNOWLEDGEMENTS
Patricia Bergquist (UAZD) encouraged this
study and provided inspiration to persevere with
the long learning curve associated with sponge
taxonomy. Felix Wiedenmayer (NMB) provided
access to his numerous unpublished personal
notes on museum collections (Sponge Archives,
NMV; Wiedenmayer, 1989; Hooper & Wiedenmayer, 1994). Rob Van Soest (ZMA) provided
numerous discussions on sponges, alternative
views on the diagnostic importance and polarity
of characters, and possible relationships between
sponge groups. Michelle Kelly-Borges (BMNH)
and Peter Jell (QM) provided many positive comments on the manuscript.
I am particularly grateful to Claude Levi
(MNHN) for providing a post-doctoral fellowship at the MNHN and giving me access to its
type collections, including all the Lamarck
material; the Sir Winston Churchill Memorial
Trust (Canberra), the Australian Biological
Resources Study (Canberra), Klaus Riitzler
(USNM), Willard Hartman (YPM), Joachim
Reitner (FUB, now Gottingen) and the Trustees
of the Queensland Museum (QM) for providing
grants at various times, enabling me to examine
major Museum collections and to interact with
colleagues at several international forums (from
which many of the ideas in this present volume
matured). The Northern Territory University
Planning Authority (1983-1985), the Museums
and Art Galleries Board of the Northern Territory
(1983-1989) and Heritage Commission of the
Northern Territory (Darwin) provided additional
funding to visit remote localities and the scattered
collections in Australian museums.
I am particularly grateful to Leonie Hooper for
line drawings and John Kennedy for many of the
SEMs. I also thank Bob Hardy (UQ), Charles
Webb (NTU), and Clive Wilkinson (AIMS) for
providing assistance with SEM photography.
For financial or logistic assistance with
fieldwork I acknowledge: George Elyakov,
Valodya Krasochin, Y. Yakovlev, USSR RV
`Akademik Opafin' (PIBOC); Alice Kay, Lester
Cannon (QM) and Queensland Fisheries Service;
Mick Ready (FV 'Hydronaue); Peter Murphy,
Martin Riddle, Shirley Sorokin, Rob McCauley
and other members of the NCI team (AIMS);
Danny Roberts (EPA); Scott Chidgey, Calwell
Connor and Associates; Patricia Byers (FV
`Skeleton'); Bill Rudman (AM); Ian Poiner and
Trevor Ward (CSIRO Fisheries, RV `Soela', RV
'Sprightly', FV 'Clipper Bird', RV 'Southern
Surveyor'); Martin Riddle and Lisa Miller
(EMU); Rob Capon (University of Melbourne);
Alan Butler (University of Adelaide); Clay Bryce
(WAM); Neville Coleman (Australian Marine
Photographic Index, Brisbane); Conservation
Commission of the Northern Territory, Darwin;
Darryl Grey, Dave Ramm and Anne Coleman
(NT Fisheries Darwin); Neil Smit (NT University
Darwin) Barry Russell and Helen Larson (NTM);
C.C. Lu (NMV); Cecile Debitus, George Bargibant, Jean-Louis Menou, Pierre Laboute
(ORSTOM Noumea); Pat and Lori Colin (NCI
CRRF Chuuk and Palau); and Ian and Pam Low
(FV 'Rachel').
I am grateful for competent field assistance and
laboratory technical assistance, during various
parts of this study, from Jodie Baxter, Steven
Cook, Lisa Hobbs, Alen Howard, Cathy
Johnston, John Kennedy, Daniel Loy Choy,
Anne-Marie Mussig, Paula Tomkins and Rex
Williams. I also thank Phil Alderslade (NTM) for
assistance in developing computer digitising
software, and Russell Hanley (NTM) for identifying commensal polychaetes.
I also thank many people for providing material
for examination, or for other information cited in
the text: Penny Barents (AM), Nicole Boury-Esnault (SME), Beatrice Burch (BPBM), Susan
Chambers (RSME), Frank Climo (NMNZ), Ruth
Desqueyroux-Faundez (MHNG and LMJG),
Jane Fromont (JCU), Manfred Grasshoff (SMF),
Jan Den Hartog (RMNH), Takomura Hoshino
(MMBS), Frank von Knorre (PMJ), Deiter
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Kuhlmann (ZMB), Vladimir Krasochin
(PIBOC), Romely Lockyer (Cootamundra
Shoals Survey team, UK), Susan Boyd (NMV),
Liz McCaffrey (UQ Brisbane), A.K. Mandal
(IM), Loisette Marsh (WAM), C. O'Riordan
(INM), Shane Parker (SAM), David Parry (NTU
Darwin), Urs Rahm (NMB), Martin Riddle
(EMU), Frank Rowe (AM), Klaus Riitzler and
Kathleen Smith (USNM), Shirley Stone
(BMNH), B.R. Stuckenberg (NM), Ole Tendal
(ZMC), Jean Vacelet (SME), Clare Valentine
(BMNH), Clive Wilkinson (AIMS), and
Wolfgang Zeidler (SAM).
DEFINITION OF CHARACTERS
MINERAL SKELETON. The form, composition
and division of the skeleton remains the most
important character for classification of
Demospongiae. Recent attempts at higher systematics of Demospongiae based on non-skeletal
characters (e.g., Simpson, 1968a; Bergquist &
Hartman, 1969; Bergquist, 1980a; Lee &
Gilchrist, 1985; Hooper et al., 1992) have had
only limited success because in many instances
they are unable to corroborate all skeletal and
non-skeletal evidence into a single systematics.
In some cases amongst Demospongiae (e.g.,
Verongida), non-skeletal evidence has been
decisive and to some extent well correlated with
other characters. In other cases (e.g., Axinellida)
that evidence has merely highlighted inadequacies in systematics based solely on skeletal
morphology (Simpson, 1968a; Bergquist &
Hartman, 1969; Bergquist, 1980a; Vacelet, 1985;
Hooper et al., 1992).
COMPOSITION OF THE SKELETON. All
microcionids are siliceous with discrete, free
spicules. So far no desma-bearing species or hypercalcified 'relict' species are known. Many
species undergo secondary acquisition, loss or
reduction of spicule mineralisation, particularly
when displaced by arenaceous particles (e.g.,
Holopsamma). C. (Wilsonella) is partly defined
by this feature, with various degrees of
arenaceous development among species; this
trend is widespread throughout the
Poecilosclerida.
ORGANISATION OFTHE SKELETON. Gross organic
and inorganic skeletal architecture, structural differentiation of the inorganic skeleton, and distribution of mineral components in that structure
are primary diagnostics (Levi, 1960a, 1973;
Bergquist, 1978a; Hartman, 1982). However,
when used alone skeletal architecture is not
necessarily a reliable indicator of relationships.
Hooper (1991, herein) noted that the so-called
typical, compressed, axinellid-like skeleton of
many Raspailiidae (Raspailia pinnatifida;
Hooper, 1991: Fig.2b) also occurred in C.
(Axociella) (Figs 7G, 119A). It is also probable
that skeletal structure is influenced to some extent
by environmental conditions, and there is some
evidence to suggest that flexible, compressed
axial skeletons are produced in response to high
energy environments (e.g., Palumbi, 1984).
Similarly, skeletal characters such as those found
in encrusting species have obviously evolved independently in many (otherwise unrelated) taxa.
Review of microcionid skeletal structures
showed that species which were similar in spicule
geometry had different skeletal architectures
(Hooper, 1988).
Architectural types amongst the Microcionidae
include: 1, hymedesmoid (with thin layer of basal
spongin lying on the substrate containing erect
megascleres (Fig. 7H); 2, microcionid (with a
compressed basal spongin, producing ascending
fibre nodes and plumose spicule columns) (Fig.
100F); 3, renieroid reticulate (consisting of a
rectangular basal reticulation of uni- or
paucispicular tracts fully enclosed within spongin
fibres or cemented at their nodes by loose collagen) (Fig. 7C); 4, isotropic reticulation (a disorientated, random uni-, pauci- or multispicular
reticulation in erect or massive forms, in which
there is no distinction between primary or secondary tracts (not figured; seen only in nominal
genus Qasimella); 5, isodictyal reticulation
(reticulation with triangular meshes formed by
uni- or paucispicular tracts of spicules, cemented
at their nodes by collagen or fully enclosed within
spongin fibres) (Fig. 131A); 6, regularly or irregularly reticulate (with large multispicular
tracts and/or fibres forming irregular oval or rectangular meshes (Fig. 7A-B); 7, plumo-reticulate
(producing ascending and consecutively diverging tracts and fibres, forming pauci- or multispicular primary lines, and interconnected by
transverse uni- or paucispicular tracts and fibres)
(Fig. 7D); 8, dendro-reticulate (similar to the
preceding, but where ascending tracts are sinuous
and more obviously diverging and branching than
the less conspicuous transverse elements) (Fig.
231C); 9, plumose (with ascending and diverging
primary lines that are not connected by transverse
elements) (Fig. 7E); 10, axially or basally compressed (having a skeleton clearly divided into a
compressed central or basal core of fibres and/or
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12
of types: 1, principal spicules
(Fig. 2A), robust, straight or
slightly curved primary styles,
subtylostyles, tylostyles or
SUBECTOSOME
quasi-monactinal forms
generally (but not invariably)
confined within fibres; 2,
auxiliary spicules (Fig. 2B),
CHOANOSOME
more slender, curved, sinuous
10 or straight styles, subtylostyles,
tylostyles or quasi-diactinal
forms most commonly located
outside fibres in the ectosomal
or subectosomal skeletons, or
dispersed throughout the
mesohyl; 3, accessory spicules
SPONGIN FIBRE
(Fig. 4), styles, acanthostyles
or modified quasi-diactinal
4568
forms generally echinating
fibres. Thus, most species have
FIG. 1. Idealised microcionid skeletal structure. 1, Echinating acanthostyles. principal spicules coring fibres
2, Reticulate fibre skeleton; 3, Isotropic extra-fibre skeleton. 4, Detrital (=choanosomal principals),
entrapping fibres. 5, Renieroid reticulate secondary fibre skeleton. 6,
some (e.g., Echinochalina)
`Microcionid' radial fibre skeleton. 7, `Spicate' spicule skeleton. 8,
Hymedesmoid spicule skeleton. 9, Plumose/dendritic fibre skeleton. 10, have coring spicules derived
Coring principal spicules. 11, Subectosomal auxiliary spicules. 12, from auxiliaries (=choanosomal
auxiliaries), and some (e.g.,
Ectosomal auxiliary spicules.
Clathria (Dendrocia)) lack
spicules, forming tight anastomosing meshes, principal spicules altogether.
and from which arise plumose or plumoreticulate
Ectosomal Skeleton. Within Microcionidae the
extra-axial (subectosomal) fibres and/or spicules)
ectosomal skeleton has been attributed greater
(Fig. 7G). Some species have combinations of
importance (Van Soest, 1984b) than over elethese skeletal structures with different structural
ments such as choanosomal architecture and
types found in different parts of the skeleton (e.g.,
growth form (e.g., Levi, 1960a; Berquist &
axis and periphery).
Fromont, 1988), microsclere diversity (e.g., de
STRUCTURAL REGIONS OF THE SKELETON. The Laubenfels, 1936a), or megascleres echinating
skeleton may be divided into three main struc- fibres and choanosomal spicules (e.g., Hallmann,
tures or regions (Fig. 1): 1, ectosomal (outer 1912, 1920). Within the family ectosomal
perforated epithelium comprised of single elon- development ranges from membraneous, without
gate flattened pinacocytes, including an ec- an ectosomal skeleton (Fig. 94F); membraneous,
tosomal skeleton, where present, arising from the with a specialised tangential, reticulate fibre
mesohyl directly below the exopinacoderm); 2, skeleton (Fig. 255C); with an ectosomal tangenchoanosomal or axial core (central or basal region tial reticulation of detritus (Fig. 92A); with subcontaining choanocyte chambers, fibres and ectosomal spicules erect, paratangential or
spicule tracts and the mesohyl ground substance, tangential to surface (Fig. 65F), or forming disin addition to the diverse and mobile cell types); crete bundles (Fig. 59E); with special ectosomal
3, subectosomal or cortical zone (strictly part of spicules erect or paratangential to surface, formthe choanosome since it lies within the bounds of ing a continuous palisade (Fig. 155G) or discrete
the external epithelial barrier, but is immediately bundles (Fig. 151F).
subdermal, and the delineation of this region from Subectosotnal Skeleton. The region between
the choanosome proper is based on the develop- choanosomal and ectosomal layers may or may
ment of an extra-fibre in some species structurally not be differentiated into a subectosomal (subderdifferentiated from the deeper choanosomal or mal or extra-axial) structure. This region may
occupy a small proportion of the peripheral
superficial ectosomal skeletons).
This system is used in conjunction with spicule mesohyl (e.g., Holopsamma; Fig. 269D), or it
nomenclature (Hallmann, 1912), based on origin may comprise the majority of sponge diameter
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FIG. 2. Major structural megascleres. A, Principal spicules (style, subtylostyle, acanthostyle subtylostyle, hastate
style). B, Auxiliary spicules (style, subtylostyle, rhabdose subtylostyle, spined subtylostyle, asymmetrical
styloid, tornostyle, quasistrongyle, mucronate styloid). C, Bases of principal spicules (style, subtylostyle, hastate
style, acanthose subtylostyle, vestigial basal spination). D, Bases of auxiliary spicules (pointed subtylostyle,
tylostyle, style, acanthose style, terminal spines, tornostyle).
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FIG. 3. Terminations of structural megascleres. A, Fusiform (pointed and rounded). B, Telescoped. C, Hastate
(pointed and rounded). D, Spined (granular, tuberculate, terminal sharp, subterminal sharp, mucronate, bifurcate).
(e.g., C. (Axociella); Fig. 118F). When present,
subectosomal extra-fibre development may be
completely disorganised or it may be well organised into radial non-plumose bundles of
spicules (Fig. 127B). The composition and
geometry of spicules within the subectosomal
skeleton varies from species lacking any form of
subectosomal spicules (e.g., C. (Microciona);
Fig. 102F), those with a single form of auxiliary
spicule throughout the skeleton (e.g., C.
(Dendrocia); Fig. 112D), those with two
categories of structural megascleres, one in the
ectosome and one in spongin fibres (e.g., C.
(Clathria); Fig. 73F), to those with three forms of
structural megascleres, all of which occur to some
extent in the subdermal region (e.g., C.
(Thalysias); Fig. 197G), and spicules may be
monactinal (for most microcionid species; e.g.,
Fig. 28A-B), or quasi-diactinal (Fig. 298A-B), or
truly diactinal (Fig. 307A-B) as seen in E.
(Protophlitaspongia).
Choanosomal Skeleton. In Microcionidae skeletal structures are diverse (listed above), and traditionally important in classification. In terms of
differentiated regions within the choanosomal
skeleton there are species with simple reticulate
fibres without any axial or extra-axial differentiation (Fig. 7C), those with marked axial and extraaxial differentiation of the choanosomal and
subectosomal skeletons, which are structurally
distinct from the peripheral skeleton (Fig. 7G),
and those in which the choanosome is simply
reduced to a basal layer of spongin lying on the
substrate (Fig. 7H). Similarly, choanosomal
skeletal tracts may be diversified into more than
one element, including those with no obvious
division of primary or secondary skeletal lines (Fig.
159F), those with more-or-less well differen-
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FIG. 4. Echinating and acanthose principal megascleres and spination. A, Microspined echinating acanthostyles
(typical recurved spines, typical erect, vestigial, poorly silicified, hastate smooth, entirely smooth). B, Myxillid-like echinating acanthostyles (aspinose apex, clavulate apex, entirely spined, bare neck). C, Modified
principal spicules (three forms of intermediate principal echinating spicules, acanthostrongyles). D, Spine
geometry (greatly recurved spines, slightly recurved spines, erect spines, granular spines, tuberculate spines).
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tiated primary (usually ascending) and secondary
(usually transverse) lines of skeletal fibres or
tracts (Fig. 48E), and those in which secondary
(choanosomal) connecting tracts may be absent,
leaving only radial primary lines (Fig. 227C).
DISTRIBUTION OF SPICULES WITHIN THE
SKELETON. Coring spicules. (Fig. 2) Spicules
coring spongin fibres or forming tracts within the
skeleton (bound by loose collagen) range from
principal megascleres (e.g., C. (Clathria); Figs
2A, 28A), auxiliary monacts (e.g., C. (Wilsonella); Figs 2B, 83A), auxiliary diacts or quasi-diacts
(e.g., E. (Echinochalina); Figs 2B, 280B), hastate
oxeas (e.g., E. (Protophlitaspongia); Figs 2B,
296A), secondarily incorporated echinating
spicules (e.g., C. (Thalysias)phorbasifonnis; Fig.
183G), acanthose principal styles, strongyles or
rhabdostyles ('plocamiform' species; e.g., A.
(Plocamia) ridleyi; Fig. 218C), coring spicules
partially or completely absent replaced by foreign
particles (detritus, algal filaments) (C. (Wilsonella); Fig. 91F; Holopsamma; Fig. 257D), primary
fibres cored by principal styles, whereas secondary (connecting) fibres contain auxiliary
megascleres (e.g., C. (Thalysias) mutabilis; not
figured), primary fibres contain auxiliary spicules
whereas secondary fibres mostly devoid of any
spicules (e.g., C. (Clathria) noarlungae sp. nov.;
Fig. 55F), axial fibres cored by auxiliary
megascleres whereas peripheral fibres heavily
arenaceous (e.g., C. (Clathria) echinonematissima; not figured).
Echinating spicules. (Fig. 4) The presence, absence, distribution, geometry and secondary
modification (including ornamentation) of
echinating spicules (accessory spicules in the terminology of Hallmann, 1912) have been used as
relatively important taxonomic characters by
some authors (e.g., de Laubenfels, 1936a),
whereas their importance in the supra-specific
classification of the microcionids has been
recently questioned (Simpson, 1968a; Van Soest,
1984b). The presence of echinating megascleres
in the two families Microcionidae and Raspailiidae indicate the retention of an ancestral character by both families, interpreted as a
synplesiomorphy (Hooper, 1991).
In Microcionidae echinating spicules may be
absent (e.g., C. (Axociella), Fig. 124F; C.
(lsociella), Fig. 134D; presumed to be a secondary loss), echinated sparsely by acanthostyles or
styles (e.g., C. (Clathria) nexus; Fig. 53E), heavily echinated by acanthostyles or smooth styles
(e.g., C. (77talysias)procera; Fig. 187G), echinated
by a second category of (acanthose) principal
spicule (e.g., A. (Antho) tuberosa; Fig. 214F), or
with both acanthose principal spicules and true
echinating spicules present on fibres (e.g., A.
(Plocamia) ridleyi; Fig. 218H).
Similarly, echinating spicules may be confined to
the exterior surface of fibres (most microcionids),
or clumped on basal spongin only and absent
from ascending skeletal lines (e.g., C. (Thalysias)
tingens sp. nov.; Fig. 201G), concentrated in tufts
at fibre nodes or fibre junctions (C. (Thalysias)
spinifera; Fig. 197G), concentrated on exterior
edges of fibres, or exclusively on primary fibres,
and/or in the peripheral skeleton only (e.g., C.
(Thalysias) abietina; Fig. 138F), or secondarily
incorporated into fibres (e.g., C. (Dendrocia)
myxilloides; Fig. 112D).
Extra-fibre spicules. (Fig. 2) The distribution of
structural megascleres within the choanosomal
skeleton is certainly a feature relied upon by
earlier authors (e.g., Carter, 1885 et seq.;
Hallmann, 1912), but its value to the supraspecific classification has been recently questioned (Van Soest, 1984b). In most species these
megascleres are confined within choanosomal
fibres, lying parallel to fibres (e.g., C. (Clathria)
raphana; Fig. 67D), or in the case of encrusting
species, embedded in basal spongin (e.g., C.
(Microciona) illawarrae sp. nov.; Fig. 100F). In
other species they may core choanosomal fibres
but also occur as plumose brushes on fibre endings (e.g., C. (Thalysias) spinifera; Fig. 197G).
Choanosomal spicules may also poke out of
fibres, much like echinating spicules (termed
`spicate'), forming plumose tracts near the surface (e.g., C. (Thalysias) major; Fig. 181A), or
they may be absent from the fibre core, standing
perpendicular to the nodes or fibre junctions (e.g.,
C. (Clathria) biclathrata; Fig. 30G), strewn in
loosely aggregated, halichondroid, extra-fibre tracts
(e.g., E. (Echinochalina)anomala; Fig. 277A), form
a renieroid structure, without a fibre component,
but merely bound at the nodes by loose collagen
(e.g., C. (Clathria) angulifera; Fig. 26F).
SPICULE GEOMETRY. Spicule geometry is an important, sometimes ultimate (e.g., de Laubenfels,
1936a), feature of existing sponge classifications,
based largely on the great diversity of
megascleres and microscleres throughout the
Porifera. This theoretical basis of sponge systematics using the mineral skeleton rests heavily
on the assumption that morphogenesis of
megascleres and microscleres is a function of the
genotype (e.g., Reid, 1963). Whilst there is no
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evidence to reject this hypothesis there is certainly some experimental data to show that spicule
geometry and morphogenesis is at least partly
influenced by environmental perturbations (e.g.,
Hartman, 1981; Jones, 1991), including examples from the Microcionidae (e.g., influence of
seasonality (Simpson, 1978) and geographical
distribution (Hooper et al., 1990) on spicule size
and geometry). But the extent to which these
phenotypic modifications occur within natural
populations has not yet been examined rigorously. In general, however, these features appear to
be relatively stable across wide geographical ranges as shown by studies on raspailiids (Hooper,
1991) and microcionids (Hooper & Levi, 1993a)
from east and west coasts of Australia and the
western Pacific. Hartman (1981) and Simpson
(1990) outline the various theories on the functional significance, process of silicification and
evolution of demosponge spicules.
STRUCTURAL MEGASCLERES. Spicule axes.
Microcionidae have exclusively monaxonic
spicule axes. Megascleres are usually monactinal, although some may have modified secondary axes (i.e., anisoxeote diactinal modifications
to styles), and a few appear to have true diactinal
forms (E. (Protophlitaspongia)). Unlike
Trikentrion and Cyamon in the allied Raspailiidae
(Hooper, 1991) there are no tetraxonic spicule
modifications in this family (triacti nal, tetractinal
or polyactinal forms). Furthermore, the
Microcionidae have a comparatively small range
of structural megasclere types in the skeleton,
whereas some raspailiids have many. Major types
of structural megascleres are illustrated in Figs
2-4. These range from hastate styles or tylostyles
(Fig. 87A), fusiform styles or tylostyles (Fig.
77A), asymmetrical styloid, rounded, quasi-diactinal or strongylote spicules (Fig. 280B), and
oxeote megascleres (Fig. 296A).
Spicule ornamentation. Spines on megascleres
are of dubious importance to supraspecific clas-
sification (e.g., Simpson, 1968a), although they
have been used frequently in the past to define
genera (e.g., de Laubenfels, 1936a). Microcionid
structural spicules frequently have basal
microspines (Fig. 2C-D), occasionally with
spines on shafts (Fig. 180B) or points of spicules
(Fig. 3D). Spicule ornamentation ranges from
entirely smooth (Fig. 28A), smooth shafts with
acanthose bases (Fig. 30A), vestigial spination on
the proximal portions of shafts only (Fig. 153A),
acanthose on both bases and points (Fig. 83A), or
entirely acanthose (Fig. 98A).
ECH1NATING MEGASCLERES. There is a diverse
range of echinating spicule geometries in
Microcionidae, although not as great as in
Raspailiidae. Major types (Fig. 4) include: evenly
spined (granular), claviform or stump-like acanthostyles; acanthose styles with aspinose bases;
acanthose styles with aspinose points; acanthose
styles with aspinose 'necks' (i.e., area proximal
to the basal swelling); acanthostrongyles; entirely
smooth styles identical in geometry to principal
megascleres; derived oxeotes; or entirely smooth
stylotes of different geometry than principal
spicules.
MICROSCLERES. The geometry, ornamentation
and modification of microscleres is an important
character for classification (Dendy, 1921), although it has probably been overemphasised by
some authors (de Laubenfels, 1936a) and its
primary importance has been questioned (Van
Soest, 1984b). Within Microcionidae there are
two forms of diactinal microscleres: meniscoid
forms (chelae) and toxas. Other poecilosclerid
microscleres (microxeas, raphides and meniscoid
forms such as true sigmas) are not present.
Microcionids show many modifications to both
chelae and toxas, the latter sometimes resembling
microxeas, and frequently microscleres are lost
altogether.
FIG. 5. Chelae geometry (A-H,J-N, Microcionidae; I,O-U, Other poecilosclerids). A, Palmate (C. australiensis).
B, Palmate, reduced alae (C. australiensis). C, Palmate sigmoid, vestigial alae (C. hesperia sp.nov.). D, Palmate,
arcuate-like alae with straight shaft (C. oxyphyla). E, Palmate, fused alae (C. curvichela). F, Palmate, contort
(C. abietina). G, Palmate, arcuate-like fusion (C. macropora). H, Palmate, central wing on shaft (C.
toxipraedita). I, Palmate, anisochelate (Mycale). J, Palmate, arcuate-like alae, fluted alae (C. macropora). K,
Arcuate-like, fusion of alae, curved shaft, alae practically fused together (C. grisea). L, Arcuate-like, unguiferous, detached alae (C. scabida). M, Anchorate-like, unguiferous, tooth-like alae (C. scabida). N,
Arcuate-like, unguiferous, tooth-like alae (C. myxilloides). 0, True arcuate (Ectyodotyx). P, Arcuate, unguiferous (Ectyodoryx). Q, Arcuate, unguiferous, tooth-like alae (Cretin). R, Palmate unguiferous (Crella). S,
Palmate, unguiferous, vestigial alae (Hamigera). T, Arcuate, unguiferous, tooth-like alae (Monanchora). U,
Anchorate, unguifcrous (Monanchora).
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FIG. 6. Toxa geometry. A, Oxhorn. B, Wing-shaped (intermediate oxhorn-accolada. C, U- and V-shaped. D,
Oxeote. E, Accolada. F, Sinuous and raphidiform.
CHELAE. These are autapomorphic for
Poecilosclerida, although Hajdu et al. (1994b)
suggested that the Isodictya might belong to
Haplosclerida (with chelae retained as an ancestral feature), although it is more likely that its
niphatid-like isodictyal skeleton is merely convergent on the haplosclerid condition. There are
many geometric modifications possible to chelae
(Fig. 5) which Hajdu et al. (1994a) proposed
could be accomodated into three main lines: pal-
mate, arcuate and anchotrate (the latter two also
termed bidentate- or tridentate-derived). These
authors hypothesised further that chelae
geometry might be an absolute indicator of
phylogeny for the Poecilosclerida based on the
assumption that they might be less susceptible to
phenotypic modifications than structural
spicules, following the arguments of Dendy
(1921, 1922). This interpretation is based on parsimony and to some extent is corroborated by
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
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other evidence (such as congruence of structural
features). There are, however, some anomolous
examples of chelae that fall between these three
categories (see Discussion).
Palmate. (Fig. 5A-B) This is the simplest form
with 'typical' morphology consisting of straight
shaft, front ala completely free and well developed,
and lateral alae more-or-less completely fused to
the shaft along its longest dimension. Most microcionids have unmodified 'typical' palmate
isochelae. Modifications to this 'typical' palmate
form include: partial reduction of alae (Fig. 5B),
nearly vestigial alae producing a sigmoid spicule
(Fig. 5C), partial fusion of alae along lateral margins producing spatulae (Fig. 5E), partial fusion
(Fig. or complete fusion along medial margins n
in which both the front alae meet and/or fuse
at the centre producing cleistochelae (Fig. 76G),
sculpturing on fluting on alae (Fig. 5J), contortion
of the shaft such that alae are at 90 0 to each other
at each end of the shaft (Fig. 5F), expansions of
the lateral alae fused with the shaft producing
wing-like process on the shaft (Fig. 5H), 'crocae'
or j-shaped sigmoid forms where the alae are
vestigial and asymmetrical, producing a simple
hook-like spicule (Fig. 17F), and deep curvature
of shaft and reduction of alae to tooth-like structures (termed unquiferous; Fig. 5R-T).
Arcuate. (Fig. 50) Here the lateral alae are more
fully developed than in palmate forms and become almost completely detached from the shaft,
and the shaft is usually prominently curved and
thickened. However, there is no clear transition
between the palmate and arcuate forms, whereby
an increase in curvature and thickening of the
shaft (Fig. 5D-E) and partial detachment of lateral
alae (Fig. 5J-K) extend along a continuum from
true palmate to true arcuate (compare Fig. 5D, G,
J-L, N-Q, T). Somewhere along this continuum
chelae are deemed to be arcuate (Fig. 50-P).
Anchorate. (Fig. 5U) Further along the continuum are anchorate chelae, in which all three
alae are fully formed, the lateral ones completely
detached from the shaft, and there are also lateral
ridges on the shaft. In this study I use the terms
'arcuate-like' or 'anchorate-like' for modified
chelae although it is equivocal whether these
spicules are truly arcuates or anchorates.
TOXAS. Toxas are found in only a few families of
poecilosclerids but also known from
Haplosclerida. There is also some evidence to
suggest that they may be particularly common in
young or larval tissue (e.g., Simpson, 1968b).
Eight major morphotypes are delineated here (Fig
6), although intermediates are also possible: 1,
Oxhorn toxas (wide central curve, reflexed arms
and greatly recurved points; usually thick) (Fig.
6A); 2, Wing-shaped toxas (sharply curved at
centre, with recurved arms and reflexed points;
usually thick) (Fig. 6B); 3, U-shaped toxas (with
wide central curvature but lacking reflexed arms)
(Fig. 6C); 4, V-shaped toxas (pinched hairpin-like
central curvature, straight arms running more-orless vertical, and slightly reflexed points; usually
thick) (Fig. 6C); 5, Oxeote toxas (virtually
straight shaft and points) (Fig. 6D); 6, Accolada
toxas (wide or slightly pinched central curvature,
strait arms running more-or-less horizontal, and
strait points; usually thin) (Fig. 6E); 7,
Raphidiform toxas (sharply angular central curvature, straight arms and straight points; very
thin, hair-like) (Fig. 6F); and 8, Sinuous toxas
(asymmetrical, sinuous, raphidiform; very thin,
hair-like) (Fig. 6F). The presence or absence of
microspines on toxas was at one time considered
to be an important supraspecific character (e.g.,
de Laubenfels, 1936a), but these have since been
found in many genera and may not be important
above the species level.
SIZE OF SPICULES. Variation of spicule size
has also been an important diagnostic criterion,
but this has been applied mainly at the species
level of classification. Numerous (possibly a
majority) of taxa have been erected solely on the
basis of megasclere and microsclere dimensions,
but only a few studies have investigated the statistical variability of spicule size or commented on
the effects of physico-chemical factors on that
variability (e.g., Hartman, 1958, 1981; Jones,
1984). There is some evidence to show that intraspecific variability can be significant for a given
taxon, and spicule size-ranges can span across
several closely related taxa which were otherwise
erected solely on that basis. Hooper et al. (1990)
demonstrated that two sibling species of Clathria
(Thalysias) could not be reliably distinguished by
their absolute spicule sizes, and only statistical
comparisons between these species were of any
value in this regard. Thus spicule dimensions
used as diagnostic characteristics are of most
significance at the species level of classification,
and consequently their application is generally
comparative rather than absolute.
ORGANIC SKELETON. The development of
the organic skeleton, the amount of spongin it
contains, its architecture and foreign inclusions
contained within it, are diagnostic features for the
Demospongiae in general. The organic skeleton
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FIG. 7. Examples of microcionid skeletal structures. A, Regularly reticulate (C.(C.) noarlungae sp.nov.). B,
Irregularly reticulate (C. (C.) multipes). C, Renieroid reticulate (A. (1.) chartacea). D, Plumoreticulate (C. (T.)
reinwardti). E, Plumose (C. (T.) procera). F, Arenaceous (C. (W.) tuberosa). G, Axially compressed, extraaxially radial (C. (A.) canaliculata). H, Hymedesmoid (C. (M.) tingens).
can be differentiated into two components: a fibre
system and a collagenous mesohyl.
Spongin fibres. These may be well developed,
present but relatively lightly developed, secondarily lost, with or without spicule tracts, or even
replaced partially or entirely by algal filaments
(e.g., Anomoclathria (Fig. 9A-B)). Simpson
(1984) elucidates two morphological types of
spongin fibres (= type A spongin of Bergquist,
1978a): those that are cored by siliceous spicules
and those that incorporate foreign particles.
Detrital-entrapping fibres are most prevalent in
the 'keratose' sponges (Dictyoceratida,
Dendroceratida, Verongida), Haplosclerida
(Arenosclera), and Poecilosclerida of the Desmacididae (Psammascus, Desmapsamnia), as
well as several species of Microcionidae (i.e., C.
(Wilsonella) and Holopsatntna). In these species
fibres may be totally void of true megascleres
(e.g., Holopsamma laminaefavosa), or they may
have a combination of both foreign particles and
indigenous megascleres (e.g., C. (Wilsonella)
tuberosa). The independent occurrence of detrital-entrapping fibres throughout the various orders of Demospongiae indicates that it is a
character of ecological importance rather than of
any phylogenetic significance.
Spiculose fibres are typical for microcionids,
and the form, size, orientation and origin of
megascleres coring the organic skeleton, and the
structural complexities of the fibres themselves
are important diagnostic characters for this family. In encrusting species that have a basal layer of
spongin anchoring themselves to the substrate
this spongin is continuous with the spiculated
fibres, and as such both actually lie outside the
living organism (Weissenfels, 1978).
Mesohyl. The development of collagen in the
mesohyl matrix, also termed 'interstitial' spongin, extra-fibre spongin, type B spongin
(Bergquist, 1978), or ground substance, and including collagenous fibrils, has not been given
much value as a systematic character for the
Microcionidae, whereas in the 'keratose' orders
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these features have more significance, and in
Aaptos (Hadromerida) it has been used to differentiate species through deposition patterns
(Kelly-Borges & Bergquist, 1995). However,
within the Microcionidae there is evidence to
show that the development of collagen
throughout the mesohyl varies intra-specifically,
especially between specimens in different
reproductive condition or as a consequence of
overwintering behaviour (e.g., Pandaros acanthifolium; Wiedenmayer, 1977; Van Soest,
1984b; Microciona prolifera; Simpson, 1963,
1968b; Knight & Fell, 1987). Simpson (1968a)
attempted to define species and genera of
Microcionidae on the basis of the organic
skeleton and cytological characteristics but to a
large extent his results did not corroborate with a
classification based on the mineral skeleton, and
in some cases evidence was directly conflicting.
GROWTH FORM. The use of external morphology
as an important or even crucial diagnostic characteristic has diminished since early systematics
(e.g., Lamarck, 1814). Bowerbank (1864), in
developing Grant's (1861) scheme for the
Porifera, de-emphasised sponge habit in his systematics although he recognised that growth form
was related to 'anatomical peculiarities'. That
external morphology is often closely linked to the
internal architecture and composition of the
skeleton has been well documented (e.g., Levi,
1973; Bergquist, 1978). Although there are some
groups which are immediately recognisable by
their growth form and skeletal architecture (e.g.,
the honeycomb reticulate structure of Holopsamma), other groups show a higher degree of intraspecific variability in their morphology (e.g.,
most Clathria), ranging from encrusting to massive forms. Moreover, there is now evidence to
suggest that gross morphology is highly plastic,
greatly influenced by prevailing environmental
conditions (temperature, depth, turbidity, currents, substrate etc.) (e.g., Hartman, 1958;
Simpson, 1968a; Fry, 1971; Palumbi, 1984). It is
not entirely clear to what extent abiotic factors
influence growth form, or the degree to which
genotype dictates possible shapes attainable by
particular species, but it is becoming more apparent that the sponge 'species' is not as immutable as previously suspected. Palumbi (1984)
proposed that sponges have evolved to be capable
of producing a quick and decisive response to
environmental adversities (unpredictable, high
energy environments), and those responses are
most readily seen as changes to both growth form
and skeletal structure (e.g., the degree to which
the skeletal becomes compressed). It is also not
clearly understood why some species seem to be
highly plastic (e.g., C. (Thalysias) lendenfeldi)
(Hooper et al., 1990), whereas others with comparable depth and geographical distributions are
much more conservative (e.g., C. (Thalysias)
abietina). Growth forms, as characters used in a
classification, can be defined as determinate (e.g.,
Holopsamma, C. (Microciona)) or indeterminate
(e.g., most other Clathria, Antho).
CONSISTENCY. Sponge texture is a highly subjective characteristic, which is difficult to quantify,
but one which may provide clues as to the composition of the skeleton, the amount of spongin
present, whether or not detritus is incorporated
into the sponge, and silicification of the skeleton
(Bergquist, 1978). A description of sponge consistency is usually an integral part of any species
description, but its application in systematics has
been mostly comparative rather than absolute.
More recently de Weerdt (1985) used consistency
as an objective feature in the systematics of
Haplosclerida. She noted that it was not only
useful in characterising particular species but that
in broad terms,texture was able to be used at the
family level of classification. For the
Microcionidae this character does not vary greatly, with most species being firm, compressible,
flexible (e.g., Clathria (Thalysias)) or soft, compressible, spongy (e.g., Holopsamma).
MACROSCOPIC FEATURES OF THE SURFACE. Surface sculpturing. Macroscopic features of the
sponge surface are important for some sponge
groups (e.g., Haplosclerida), and surface ornamentation, such as tangential webs of spicules
or fibres, perpendicular brushes of spicules, or
elevated oscules may be diagnostic for particular
genera. Within the Poecilosclerida however, including the Microcionidae, these features are less
consistent, and they are usually only reliable in
characterising particular species or occasionally
genera (Simpson, 1968a). Encrusting species frequently exhibit intricate drainage canals radiating
away from oscules, or highly hispid (furry) surfaces (e.g., C. (Thalysias) toxifera), whereas
more massive or digitate species may have surface papillae or conules (e.g., C. (Thalysias)
abietina), or a surface which is composed of
reticulate ectosomal fibres (e.g., Holopsamma
globosa).
Oscules. The distribution of oscules on the surface may vary considerably between related
species, ranging from being confined to distinct
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pore areas, such as sieve-plates of Echinochalina
tubulosa, or restricted to certain regions, such as
lateral sides of branches, the tops of digits, or the
exterior surface of vases, or scattered indiscriminantly over the surface. Oscules may also be conspicuous, discrete, with a membraneous lip, slightly
raised or flush with the surface (e.g., C. (Thalysias)
reinwardti), or terminal, raised on the apex of
surface papillae or stoloniferous tubes (e.g., C.
(Isociella) eccentrica), or scattered, conspicuous,
producing a porous reticulate surface (e.g.,
Holopsamma arborea), or minute and not easily
visible optically (e.g., C. (Thalysias) coppingeri).
COLOURATION. Sponge pigments are not generally diagnostic (Bergquist, 1978). Some species
may show high intra-specific variation in live
pigmentation, and this variability may be related
directly to microhabitat and depth distribution.
The nature of these pigments, their distribution
within the mesohyl and their specific cellular
association is still poorly known, but it is well
established that carotenoids are predominant
(Simpson, 1984). Isolation and identification of
these pigments is more difficult, as is the determination of whether they are produced or
modified by the host, or obtained directly from
symbiotic associations (e.g., Litchfield & LiaaenJensen, 1980). The major proportion of sponge
carotenoids are metabolised by the sponge (i.e.,
primary metabolites) and are intracellular
(Simpson, 1984), whereas it is suspected that
some sponges have a variable proportion (up to
20%) of pigments synthesised by symbiotic algae
(e.g., Litchfield & Liaaen-Jensen, 1980; LiaaenJensen et al., 1982). Litchfield & Liaaen-Jensen
(1980) studying C. (Microciona) prolifera suggested that the sponge could modify (aromatise)
a large proportion of algal carotenoids, and
Liaaen-Jensen et al. (1982) divide the classes of
carotenoids into a phytoplankton-type,
zooplankton-type, bacterial and/or fungal origin,
and sponge metabolised (oxidative) groups.
These authors found that phytoplankton derived
and sponge metabolised carotenoids comprised
the major proportion of carotenes in Demospongiae. The Poecilosclerida and Axinellida were
found to exhibit the highest capacity for
carotenoid accumulation and transformation, explaining their diverse and often brightly coloured
pigmentation, and furthermore they possessed a
similar carotenoid diversity.
Evidence suggests that carotenoid pigments
may be photoprotective, in which case it would
be expected that intertidal species contain a
higher proportion of these pigments than deeperwater species. But it is not clear why some sympatric species have consistent pigmentation (e.g
C. (Wilsonella) tuberosa), whereas in others pigmentation is highly variable even in specimens
growing side-by-side (e.g., C. (Thalysias)
abietina; Plate 4D). Colour consistency is not
generally used as a reliable diagnostic character,
but it is also true that only very few authors have
investigated the intraspecific colour variability of
any species. It is therefore advantageous to determine whether live colouration is stable and specific
to a species, or has very narrow limits in variation
(e.g., C. (Isociella) eccentrica). Alternatively,
pigmentation may be highly unstable, not specific
and without an accurately definable 'typical'
colouration (e.g., C. (Thalysias) abietina).
REPRODUCTIVE PRODUCTS AND REPRODUCTIVE
CYCLES. Reproductive products and modes of
reproduction, as diagnostic characters, have been
used predominantly at higher levels of classification (e.g., Bergquist, 1980a), whereas breeding
seasons and spawning cycles are most useful for
detecting sibling species (e.g., Fromont, 1989).
As far as known, within the Microcionidae larvae
are viviparous parenchymella with bare posterior
poles. The apparent form of sexuality varies from
gonochoristic to contemporaneous hermaphroditism (Fell, 1984, 1990; Simpson, 1984).
Breeding seasons and/or spawning cycles may be
continuous or periodical.
CYTOLOGY. Simpson (1984) provided a definitive treatment of sponge cell biology, including a
description of diverse cell types and their functional morphology. He suggested that descriptions of characters such as cells with inclusions
and the morphology of choanocyte chambers will
probably provide further information directly
relevant to demosponge systematics. For the
Microcionidae, Simpson (1968a) showed that
seemingly morphologically convergent genera
could be readily differentiated by the presence
and morphology of special cell types (gray cells),
and that their higher systematic relationships
could be defined in terms of cytological characters. However, there were many incongruities
between systematics based on skeletal characteristics and those indicated by cytological data.
Specific examples of these differences are discussed below in the synopsis of genera, but some
general comments are appropriate.
Taxonomic groupings indicated by Simpson's
microcionid cytological data suggested that many
skeletal characters used previously by authors
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had little importance in differentiating genera.
These included the presence or absence of palmate isochelae, the presence of acanthose versus
smooth echinating megascleres, quantity of spongin in the skeleton, plumose versus anastomosing
fibres, megascleres with basal spination or
smooth bases, the presence of surface conules and
distinct oscules, the production of upright
branches, and the presence or absence of an ectosomal skeleton. With the exception of the last
feature these conclusions are supported in the
contemporary classification of Microcionidae
(e.g., Van Soest, 1984b). However, other
cytological evidence presented by Simpson
(1968a) is more difficult to reconcile with
microcionid skeletal data. For example, encrusting species (i.e., the nominal genera Microciona
and Ophlitaspongia) were cytologically relatively homogeneous and distinct from ramose forms
(Clathria). The cytological characteristics of
these encrusting species were more similar to
renieroid microcionids (nominal genus Plocamilla) than they were to the ramose forms (nominal
genera Rhaphidophlus and Thalysias) which
otherwise had the most similar spicule and surface characteristics.
Simpson concluded that generic definitions
based primarily on spicule types did not lead to
natural classifications, and he proposed that these
definitions should include skeletal, cytological
and histological evidence. He suggested that the
numerous classification systems that were based
solely on various combinations of skeletal characters, such as those of Vosmaer (1933, 1935a-b),
de Laubenfels (1936a) and Levi (1960a), could
be defended with equal justification. Although
some cytological features have been incorporated into existing systematics (e.g., morphology and arrangement of choanocytes), much of
Simpson's (1968a) important work cannot be
used in classification based primarily on skeletal
characters.
SYSTEMATICS
Class Demospongiae Sollas, 1885
Order Poecilosclerida Topsent, 1928
Suborder Microcionina Hajdu, Van Soest &
Hooper, 1994
Poecilosclerida Topsent, 1928a: 64, 309.
REMARKS. This order is the largest and most
diverse of Demospongiae (Bergquist 1978). It is
characterised by a skeleton of both spicule and
spongin elements, usually well developed, sometimes vestigial, in which megascleres are monactinal, diactinal or both, and spongin development
varies from well developed horny fibres enclosing spicules to an interspicular collagen cement
(Bergquist, 1978; Hartman, 1982). Simpson
(1984) suggested that the order is characterised
by at least two distinctly localised types of
megascleres (with or without distinctive
geometry). Those megascleres are choanosomal
principal spicules embedded in spongin fibres,
and subectosomal auxiliary megascleres which
are free in the mesohyl or protrude from spongin
fibres in which they are embedded. This definition is consistent with the inclusion of
Raspailiidae in the Poecilosclerida as proposed
by Hooper (1991). Poecilosclerids usually have
an abundantly collagenous mesohyl matrix, and
microscleres may include chelae (apomorphic for
the order), although not all taxa have them.
Sexual reproduction is predominantly
viviparous, oviparous in two families, and in
those species incubating larvae they are
parenchymella with uniform flagellum size and
bare posterior poles.
The suborder Microcionina was established to
include four families of Poecilosclerida
(Microcionidae, Raspailiidae, Iophonidae and
Rhatxleremiidae), which have terminally spined
ectosomal monactinal megascleres (occasionally
modified to quasidiactinal forms), isochelae of
palmate origin, diverse forms of toxas, up to five
categories of megascleres and lacking sigmas.
The other suborders (Myxillina and Mycalina)
were also defined by their chelae morphology
(bidentate-derived and sigmancistra-derived
chelae, respectively), and absence of toxas and
presence of sigmas, respectively (Hajdu et al.,
1994), but assignment of particular genera to
these suborders is still contentious.
The number of families recognised in the order
varies according to different authors (e.g., Levi,
1973; Wiedenmayer, 1977; Bergquist, 1978;
Hartman, 1982; Van Soest, 1984b; Bergquist &
Fromont, 1988). Recently Hooper & Wiedenmayer (1994) included 16 families in the order:
12 with chelae microscleres, 3 without chelae,
and 1 of uncertain placement, whereas Hajdu et
al. (1994) recognise 17: Microcionina
(Microcionidae, Raspailiidae, lophonidae, Rhabderemiidae); Myxillina (Myxillidae, Crambidae,
Coelosphaeridae, Crellidae, Hymedesmiidae,
Anchinoidae, Phoriospongiidae, Tedaniidae);
and Mycalina (Mycalidae, Hamacanthidae,
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Desmacellidae, Cladorhizidae, Guitarridae).
Latrunculiidae, included in the order by Levi
(1973) and Van Soest (1984b) has also been
assigned to Hadromerida (Reid, 1968;
Bergquist, 1978; Hartman, 1982), but is now
considered to be polyphyletic (Kelly-Borges &
Vacelet, 1995) with Latrunculia having affinities
with Iophonidae and Diacamus, Sigmosceptrella, Negombata more closely related to the
Mycalidae.
Family Microcionidae Carter, 1875
Microcionina Carter, 1875. Microcionidae Hentschel,
1923; Wiedenmayer, 1977.
Clathriidae Lendenfeld, 1884a; Hentschel, 1923; Topsent, 1928a; Levi, 1960a; Simpson, 1968a;
Bergquist, 1978; Hartman, 1982; Van Soest, 1984b;
Bergquist & Fromont, 1988.
Ophlitaspongiidae de Laubenfels, 1936a; Thomas,
1968; Hoshino, 1981.
Growth form encrusting, lobate, arborescent or
flabellate; skeleton differentiated into
choanosomal (axial), subectosomal (extra-axial)
and ectosomal regions; axial skeleton formed by
unispicular or multispicular tracts of
choanosomal (principal) megascleres, typically
coring spongin fibres or sometimes simply bound
together by collagen; fibres echinated by (acantho-) styles (accessory spicules); skeletal structures include isodictyal, renieroid, reticulate,
plumo-reticulate, plumose or hymedesmoid, but
never radial; extra-axial skeleton formed by tracts
of subectosomal (auxiliary) spicules, usually dispersed outside of fibres, rarely well organised but
usually with some degree of difference between
axial and extra-axial regions; ectosomal skeleton
ranges from membraneous, or with protruding
subectosomal (auxiliary) spicules, or with a special category of ectosomal (auxiliary) spicules;
principal megascleres monactinal, predominantly smooth or partially spined only, occasionally
vestigial or absent completely, or sometimes
replaced by detritus in skeleton; auxiliary
megascleres usually monactinal, rarely quasidiactinal, smooth shaft and basal spines, more
slender than choanosomal spicules; echinating
styles or subtylostyles smooth, partially or completely spined; microscleres include toxas of
several morphologies (including raphidiform and
microxeotes), and isochelae primarily of palmate
origin (but occasionally with partial 'arcuate' and
`anchorate' modifications); larvae viviparous.
REMARKS. There has been disagreement as to
which of Microcionidae Carter and Clathriidae
Hentschel should be used. Wicdcnmayer (1977:
139) argued that Microcionidae was established
in 1875, whereas Clathriidae did not appear until
1884. He noted that under Article 40 of the International Code of Zoological Nomenclature
(Anonymous, 1984), it was irrelevant whether or
not Clathria Schmidt (1862) had priority over
Microciona Bowerbank (1862; apparently published 1863). Conversely, Van Soest (1984b: 89)
argued that the priority of Clathria over
Microciona did have bearing on the choice of the
family name. Although 'Clathriidae' is in current
usage by most contemporary workers, its
preferred use is in direct contravention with the
Code and to long term stability of the group and
Microcionidae is used here following Hooper &
Wiedenmayer (1994).
The definition given above restricts
Microcionidae to genera which possess
predominantly smooth monactinal ectosomal and
choanosomal spicules. It excludes certain
microcionid-like genera which have true tylotes
or strongylotes as their ectosomal spicules (e.g.,
Acamus, Megaciella). These taxa are now
referred to Iophonidae, as defined by their ectosomal features (Hajdu et al., 1994). However,
the definition barely distinguishes species with
modified or reduced quasidiactinal (styloid)
auxiliary megascleres (e.g., several
Echinoclathria, Holopsamma and Echinochalina
species), or quasimonactinal (amphistrongylote
or tornote-like) auxiliary megascleres (e.g., E.
(Protophlitaspongia)). These modified auxiliary
spicules are usually asymmetrical and are interpreted here as convergent upon true diactinal
spicules. These anomalous microcionids share
certain characteristics of both Microcionidae and
Desmacididae, and the importance of these characters at higher levels of systematics must therefore be questioned, or a certain level of
homoplasy must be accomodated in the
phylogeny of the order.
Similarly, the definition given above cannot
always clearly distinguish some Microcionidae
and Raspailiidae, but this is a problem of semantics rather than a biological one. As a general rule
most species of Raspailiidae have well compressed
axial skeletons, and well differentiated axial and
extra-axial skeletons. In contrast, most Microcionidae lack these features or they are only poorly developed and probably convergent, perhaps
related to growth form (e.g., Clathria (Axociella)). Nevertheless, there are examples in both
families where the boundaries between taxa blur,
such as the microcionid-like Raspailia (Clathriodendron) arbuscula (see Hooper 1991: Figs 19-
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20), and the raspailiid-like Clathria (Axociella)
canaliculata (Figs 118-119)). These families are
consistently differentiated by their ectosomal features and microscleres, which appear to be more
important characters than skeletal structure.
Hajdu et al. (1994) restricted Microcionina
(and hence Microcionidae) to taxa with only palmate isochelae, tacitly excluding several
microcionid-like genera specifically created for
species with bidentate-derived (arcuate or
anchorate) chelae. Theoretically this is a viable
system for the suprafamily classification of
Microcionidae but in practical terms it is not
always possible to distinguish between true
bidentate-derived chelae and palmate chelae with
'arcuate-' or 'anchorate-like' modifications. These
cases are discussed indivually below.
REVIEW. There are several problems in the
taxonomy of Microcionidae that need to be addressed in order to clearly recognise and define
valid genera and produce a phylogenetically valid
systematics for the family.
1) The family is large, containing about 540
described species and many other as yet undescribed species known from various collections. 79 nominal genera have been previously
included, of which 69 are currently recognised as
residing here although fewer than this number are
valid. Some of these genera have been merged in
others by previous authors (e.g., Levi, 1960a;
Simpson, 1968a; Van Soest, 1984b; Bergquist &
Fromont, 1988; Hooper, 1990a), but in some
cases these synonymies are now deemed wrong
and have produced further nomenclatural complexities. Several contemporary studies have attempted partial revisions of Microcionidae (Van
Soest, 1984; Bergquist & Fromont, 1988;
Hooper, 1990a), but these have mainly focused
on smaller regional faunas without consideration
of all the higher taxa. In the present work each of
these genera is redefined and illustrated from its
type species (i.e., strict definition).
2) The literature on Microcionidae is vast, scattered, mostly antiquated (pre-1900), descriptions
are far too brief for modern purposes and many
taxa have never been illustrated. The present
work deals primarily with museum material and
living populations of species, and decisions are
less reliant on the literature than previous studies.
3) There are many characters in sponges whose
expressions (character states) change subtly
within populations of supposedly single species
and across the whole range of species, usually
without clear boundaries between related taxa.
Some of these characters have been used as important diagnostic criteria in earlier works. This
study has examined large numbers of specimens
and species, and documents the range of intraspecific and inter-specific character states in an
effort to clearly define taxa and understand
relationships between them. Inclusion of nonskeletal evidence into the systematics can further
support or refute opinions based solely on skeletal
characters and gross morphology (to decide
whether one character is more important than
another, whether morphological characters are
homologous, and whether the observed high
levels of homoplasies within most Poriferan classifications are in fact real or acceptable). The
previous studies of Hooper et al. (1990) and
Hooper (1990a) are preliminary to this study.
4) There are nearly as many subjective interpretations between different authors, as to the
phylogenetic importance of one character over
another in the systematics, as there are taxa. This
has arisen partly as a consequence of over
reliance on definitions of type species (and hence
nominal genera) from the literature (especially
the work of de Laubenfels, 1936a), given that
many type species are poorly described, misdescribed or barely differentiated from their
cogeners. The present study uses a phylogenetic
framework to produce an objective and consistent taxonomy for the family. Two previous
studies (Van Soest, 1984b; Hooper, 1990a) partially resolved intrafamily relationships within
Microcionidae, both are preliminary to this work.
GENERIC NAMES INCLUDED IN
MICROCIONIDAE
Preoccupied generic names are shown in square
brackets. The synonomy lists provided in this
section refer to works in which the name is used
and in the case of genera considered valid do not
include the numerous synonyms. The diagnoses
provided in this list are based solely on the type
material of the type species unless otherwise
stated.
[Abila] Gray, 1867
(Fig. 8A-B)
Abila Gray, 1867: 539.
Not Abila Gray, 1867: 522.
TYPE SPECIES. Microciona laevis Bowerbank, 1866:
124 (by monotypy) (holotypc BMNH1877.5.21.1543).
Encrusting growth form. Surface hispid, even.
Choanosomal skeleton composed of short
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FIG. 8. Type species of microcionid genera. A-B, Abila (Microciona laevis Bowerbank, BMNH1877.5.21.424).
C-E, Allocia (Spanioplon chel(erum Hentschel, SMF1571). F-H, Anaata (A. spongigartina de Laubenfels,
USNM21428). I, Anomoclathria (Alcyonium opuntioides Lamarck, MNHNDT654).
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plumose spongin fibre nodes, cored by very long
smooth or basally spined choanosomal principal
styles, with only slightly swollen bases, and with
bases embedded in spongin fibres on substrate
and points protruding through ectosome.
Echinating acanthostyles erect on substrate. Subectosomal skeleton of a single category of very
long, curved, sinuous or straight subectosomal
auxiliary subtylostyle, with smooth or microspined
bases, forming irregular brushes on ectosome.
Special ectosomal spicules absent. Microscleres
thick wing-shaped toxas. Isochelae absent.
REMARKS. Gray (1867) used the generic name
Abila on two occasions, for two separate taxa.
The senior name (Gray, 1867: 522) refers to a
species originally described in Raspailia (viz. R.
freyerii Schmidt), which Strand (1928: 32) unnecessarily proposed a replacement name
Abilana Strand and which is a junior synonym of
Raspailia Nardo (Strand should have replaced the
junior name). The junior name (Gray, 1867: 539)
was used fora microcionid, M. laevis Bowerbank,
related to Microciona in skeletal structure, and
related to Thalyseurypon de Laubenfels in lacking isochelae and having an encrusting growth
form. De Laubenfels (1936a: 112) merged Abila
with Hymantho Burton (see below), and Van
Soest (1984b: 90) subsequently reduced Hymantho into synonymy with Clathria. Re-examination of the type species shows that the genus
belongs with Clathria (Microciona).
Allocia Hallmann, 1920
(Fig. 8C-E)
Allocia Hallmann, 1920: 768; Bergyuist & Fromont,
1988: 95.
TYPE SPECIES. Spanioplon chehferum Hentschel,
1 9 1 1: 362 (by original designation) (holotype
ZMB4440, paratype SMF1571).
Arborescent, foliose, planar growth form. Surface hispid, uneven. Choanosomal skeleton
reticulate, with multispicular ascending primary
fibres and paucispicular transverse connecting
fibres. Spongin fibres heavy, cored by styles and
tylotes (also scattered throughout mesohyl),
echinated by acanthostyles with spinose shafts
and bases, apinose 'necks', and large recurved
spines. Subectosomal region with ascending
primary fibres cored by choanosomal principal
styles, latter protruding through surface. Ectosomal skeleton with both tangential and
paratangential layers of auxiliary spicules, latter
protruding from peripheral fibres and also lying
tangential to surface. Megascleres completely
smooth principal styles of a single size category,
smooth subectosomal auxiliary stylote spicules,
with asymmetrical rounded and microspined
ends, and acanthostyles. Microscleres palmate
isochelae of two sizes. Toxas absent.
REMARKS. Hentschel (1911, 1912) expressed
doubts concerning the initial generic assignment
of S. cheliferum suggesting it showed certain
affinities with Ectodoryx Lundbeck. Hallmann
(1920) created Allocia to receive the species,
including it in Microcionidae because of its
'typical' microcionid microscleres, echinating
and coring megascleres. For similar reasons
Dendy (1922: 70) merged the type species with
Clathria. Based on its described ectosomal characteristics, supposedly consisting of true tylote
spicules, the genus would be assigned to
Iophonidae (see discussion for Acarnus), but reexamination of the type material found that these
spicules were asymmetrical, modified styles very
similar to those found in Clathria (Thalysias)
major.
Allocia is monotypic and belongs in Clathria
(Clathria). The type species is known from the
north and southwest coasts of Australia (Arafura
Sea and Perth region), Amirante (Indian Ocean),
and New Zealand (Bergquist & Fromont, 1988).
Anaata de Laubenfels, 1932
(Fig. 8F-H)
Not Aaata Semenov, 1906.
Aaata de Laubenfels, 1930: 27.
Anaata de Laubenfels, 1932: 89, 1936a:109.
TYPE SPECIES. Aaata spongigartina de Laubenfels,
1 93 0: 27 (by original designation) (holotype
USNM21428).
Encrusting growth form. Surface hispid, even.
Choanosomal skeleton hymedesmoid, with bases
of both echinating acanthostyles and
choanosomal principal subtylostyles embedded
in basal fibres. Ectosome with dense erect
palisade of subectosomal auxiliary subtylostyles
overlaying choanosomal megascleres, both
protruding through surface. Megascleres principal choanosomal styles with prominently acanthose bases and partially acanthose shafts,
echinating acanthostyles with aspinose points,
and auxiliary subtylostyles of a single size
category, being completely smooth. Microscleres
two size categories of palmate isochelae with
arcuate modifications (strong curvature, partially
detached lateral alae, slightly pointed unguiferous teeth). Toxas absent.
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FIG. 9. Type species of microcionid genera. A-B, Anomoclathria (Alcyonium opuntioides Lamarck,
MNHNDT654). C-E, Antherochalina (A. crassa Lendenfeld, BMNH1886.8.27.450). F-I, Anthoarcuata (A.
graceae Bakus, USNM36284).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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25
REMARKS. De Laubenfels (1932: 90) suggested
that Anaata was characterised in part by lacking
echinating acanthostyles, although he noted that
two size categories of choanosomal megascleres
may be present. However, in the holotype the
smaller category of spined spicule standing perpendicular to spongin fibres are true echinating
acanthostyles, as found in most other
microcionids. These smaller spicules are
predominantly spined, whereas larger
choanosomal spicules are only partially spined
and are principals.
Anaata was originally assigned to Hymedesmiidae Topsent, compared with genera such as
Leptosiopsis Topsent (1927: 13) and Leptosastra
Topsent (1904a: 194) (both of which have ectosomal tornotes with polytylote or anisotornote
modifications, and anisochelae). All three genera
have hymedesmoid architecture, which is persistent in Hymedesmiidae but also seen in many
encrusting Microcionidae. Van Soest (1984b)
mentioned that Anaata differed from other thinly
encrusting microcionids, such as Clathria
(Microciona) having arcuate isochelae replacing
(or in addition to) palmate isochelae. It is questionable, however, whether these chelae are truly
arcuate (as defined by Hajdu et al., 1994), given
that their lateral alae are fused to the spicule shaft
more than half way along their length. The
smaller chelae are more-or-less evenly curved,
slightly unguiferous (with reduced, slightly
pointed alae), and lateral alae are fused to the
shaft for about three-quarters their length. The
larger spicules have more pronounced central
curvature, they are more unguiferous (alae are
reduced, sharper), and lateral alae are fused to the
shaft for only about half their length. Anaata is
considered here to be a Clathria (Microciona)
with hymedesmoid architecture and chelae with
slight 'arcuate' modifications.
Anomoclathria Topsent, 1929
(Figs 81, 9A-B)
Anomoclaihria Topsent, 1929: 26.
TYPE SPECIES. Alcyonium opuntioides Lamarck,
1 8 1 5: 164 (by original designation) (lectotype
MNHNLBIMDT654).
Erect, arborescent or flabellate-digitate growth
form. Surface smooth, not hispid. Choanosomal
skeleton divisible into two components. Ascending primary skeleton plumose, with spongin
fibres cored by paucispicular tracts of
choanosomal principal styles and also incorporate algal filaments. Secondary basal skeleton
renieroid reticulate, with spongin fibres fibres
cored by large acanthostyles also grouped into
plumose bundles on surface and secondarily incorporated into ascending primary fibres.
Echinating spicules absent. Ectosome with sparse
tangential skeleton of subectosomal auxiliary
subtylostyles. Megascleres smooth choanosomal
principal styles, acanthostyles with large spines
in renieroid skeleton, with same morphology as
principal spicules, and smooth subectosomal
auxiliary subtylostyles. Microscleres palmate
isochelae, wing-shaped and accolada toxas.
REMARKS. This diagnosis on the lectotype differs slightly from Topsent's (1929, 1932)
redescriptions of the species, particularly in the
emphasis on the secondary renieroid skeleton
overlaying the ascending plumoreticulate
primary skeleton. This is a prominent diagnostic
feature of Antho. In A. (Antho) the secondary
renieroid skeleton is composed of acanthostyles,
as described above for A. opuntioides, whereas in
another species originally referred to
Anomoclathria, Spongia frondifera Lamarck,
1814: 445 (lectotype MNHNLB1MDT565), the
spicules forming the renieroid skeleton are acanthostrongyles (diagnostic for A. (Plocamia)). Topsent (1932: 103) considered that S. frondifera a
junior synonym of A. opuntioides, whereas
Lamarck's type material shows that their differences in a number of characters, including
spicule geometry, is sufficient to warrant species
level separation.
Anomoclathria was erected on the basis of
'styloprothese' (Topsent, 1929), whereby algal
filaments are incorporated into spongin fibres,
partially or completely replacing the coring
choanosomal spicules. But this symbiosis has
subsequently been shown to be relatively common in marine sponges (Scott et al., 1984), occurring in many families (e.g., Bowerbank, 1862a;
Carter, 1878; Lendenfeld, 1886b; Topsent, 1929,
1932, 1933; Weber-van Bosse, 1910,1921; Sciscioli, 1966; Scott et al., 1984; Price et al., 1984),
and is an ecological rather than phylogenetic
phenomenon. Although there is some evidence to
suggest that particular species of algae are
specific to particular sponge species, or at least
restricted to a narrow range of taxa (Price et al.,
1984)), it is unlikely that this symbiosis can be
used as a diagnostic character at the supraspecific level. Nevertheless, it is intriguing to
consider that a sponge can shed most or all of its
diagnostic characters (spicules), replacing them
with organic symbionts (e.g., see Clathria
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(Thalysias) abietina) or inorganic foreign particles (e.g., see Clathriopsamma, Wilsonella).
Antherochalina Lendenfeld, 1887
(Fig. 9C-E)
Antherochalina Lendenfeld, 1887b: 741, 786; Burton,
1934a: 558; de Laubenfels, 1936a:112.
TYPE SPECIES. Antherochalina crassa Lendenfeld,
1887b:787 (by subsequent designation; Burton,
1934a: 558) (holotype BMNH1886.8.27.450).
Erect, thin lamellate growth form. Surface
smooth, not hispid. Choanosomal skeleton with
differentiated axial (compressed, renieroid
reticulate) and extra-axial (loose plumose)
skeletons, although no regional differences in
spiculation. Spongin fibres heavy, cored by
choanosomal principal subtylostyles, echinated
by small acanthostyles. Ectosomal skeleton with
sparse tangential subectosomal auxiliary subtylostyles. Megascleres include robust, entirely
smooth choanosomal principal subtylostyles,
lightly spined acanthostyles, and auxiliary subtylostyles with basal spines. Microscleres palmate isochelae and wing-shaped toxas.
REMARKS. Burton (1934a) designated A. crassa as type species of Antherochalina, in
preference to Lendenfeld's (1887b) first-named
species, Veluspa polymorpha var. infundibuliformis Maclay which was unrecognisable. He noted
that A. crassa was a synonym of Clathria s.s..
Furthermore, of the eight species placed in the
genus by Lendenfeld, only the type species now
belongs here, whereas the other species are either
unrecognisable or have affinities with Raspailia
(Syringe/la), Phakellia, Ophlitaspongia, Cymbastela (A. concentrica; Hooper & Bergquist,
1992) or Ectyoplasia (A. frondosa; Hooper,
1991). Antherochalina also resembles some Desmacellidae (such as Sigmaxinella) and some
Axinellidae (such as Axinella) in skeletal structure, having a slightly compressed renieroid axial
and plumose extra-axial skeletons.
Antho Gray, 1867
(Fig. 10A-C)
Antho Gray, 1867: 524; Levi, 1960a: 57,76; Van Soest
& Stone, 1986: 42; Bergquist & Fromont, 1988: 96.
Plocamilla, in part, Burton, 1935a:402; Pulitzer-Finali,
1973:40 (not Plocamilla Topsent, 1928a: 63).
TYPE SPECIES. Myxilla involvens Schmidt, 1864: 37
(by monotypy) (schizotype BMNH1867.3.11.92).
Thinly encrusting (s.s.) or erect, arborescent,
lamellate or vasiform growth forms. Surface
rugose, hispid. Choanosomal skeleton renieroid
reticulate with acanthostyles-strongyles coring
spongin fibres, or simply united at nodes by variable quantities of spongin, producing triangular
and rectangular skeletal meshes. Junctions of
skeletal meshes with principal choanosomal
styles echinating fibre nodes, standing erect or at
oblique angles, in tufts or singly. True echinating
megascleres absent (i.e., undifferentiated from
choanosomal principal styles). Ectosome contains tangential or paratangential multispicular
brushes of subectosomal auxiliary styles protruding through surface. Megascleres acanthose principal styles/strongyles of renieroid basal
skeleton, smooth or acanthose principal
styles/subtylostyles of the choanosomal skeleton,
and smooth subectosomal auxiliary styles, often
with basal spines. Microscleres palmate
isochelae, wing-shaped and accolada toxas.
REMARKS. This diagnosis is based on the type
species and primarily on the type material, given
the existing confusion about the true identity of
A. involvens (cf. its alleged synonym A. inconstans; Ackers, Moss & Picton, 1992). Antho appears to be the earliest available name for a group
of myxillid-like plocamiform sponges (sensu de
Laubenfels, 1936a) which have microsclere
spiculation typical of other Microcionidae. These
taxa have an axial or basal skeleton composed of
mostly acanthose styles or strongyles, producing
a more-or-less regular renieroid reticulation. For
this reason Berquist & Fromont (1988) referred
Antho to the Myxillidae, but the genus has
monactinal rather than specialised tylote ectosomal spicules and does not fit the concept of
Myxillidae (Van Soest, 1984b).
The potential generic synonymy of Antho is
large. Burton (1930a: 501), de Laubenfels (1936a:77), Levi (1960a: 57) and Van Soest ( I 984b)
combined species of Diayoclathria Topsent in
Antho (see below), and that genus has largely
disappeared from current usage. Burton's (1959a:
252) merger of Myxichela de Laubenfels (1935:
331, 1936a: 85) (type species Lissodendoryx
tawiensis Wilson (1925: 432)) into Antho is not
upheld here, because it has a true myxillid ectosomal skeleton (di actinal (tylote) ectosomal
spicules).
Renieroid reticulate skeletal architecture is not
restricted to Myxillidae, known to occur in other
families of sponges (e.g., Chalinidae (Haplosclerida), Iophonidae (Acarnus), Raspailiidae
(Amphinomia Hooper, Plocamione Topsent,
Lithoplocamia Dendy) and Axinellidae (Pitalia
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
^
27
Gray)), although it is certainly most common in
the Myxillidae. The importance of a renieroid
skeleton is interpreted differently by different
authors, some giving it primary emphasis (e.g.,
Bergquist & Fromont, 1988) and others relegating it lesser importance (e.g., Van Soest, 1984b).
Several species-groups have been created to accomodate microcionid-like species, with spiculation typical of the family, supplemented by a
renieroid (myxillid-like) basal choanosomal
skeleton composed of acanthose styles or strongyles. Vosmaer (1935a: 653) called this group
Microciona prolifera tropus renieroides and de
Laubenfels (1936a) recognised it at the family
level (i.e., Plocamiidae sensu de Laubenfels (a
junior homonym of Plocamiidae Topsent,
1928a)). However, only three genera appear to be
sufficiently different to encompass all these
microcionids: 1) Plocamia Schmidt (including
Plocamilla Topsent, Dirrhopalum Ridley, and
Holoplocamia de Laubenfels), which has
predominantly (acantho)strongyles forming the
renieroid skeleton (less commonly styles), and
echinating acanthostyles overlap the main
skeleton; 2)Antho Gray (including Anthoarcuata
Bakus, Dictyoclathria Topsent, Isociona
Hallmann and Jia de Laubenfels)), which has
predominantly (acantho)styles forming the
renieroid skeleton (less commonly strongyles),
and echinating acanthostyles are absent; and 3)
Isopenectya Hallmann, which has an axially
compressed and extra-axially renieroid reticulate
skeleton composed of two forms of choanosomal
spicules inside spongin fibres, overlaid by a
second extra-axial plumose skeleton. Several
authors (Levi, 1960a; Simpson, 1968a; PulitzerFinali, 1973) were unable to reach a consensus of
whether or not Plocamilla and Antho were identical, since they only really differed by the
presence or absence of echinating spicules, and
the extent to which basal spicules of the renieroid
skeleton are styles or strongyles (i.e., intermediates occur). Most authors tentatively retain
these two genera; Simpson (1968a) and Van Soest
& Stone (1986) suggest edthat any decision on
these genera, which differ from Clathria in
having a renieroid skeletal architecture, would
require thorough re-examination of all type
species. This has now been done and the conclusion reached here is that differences between
all three species-groups (Antho, Plocamia and
Isopenecta) are not as great as the similarities (as
conferred by the possession of renieroid skeletal
structure), and these differences are emphasised
only at subgenus.
Anthoarcuata Bakus, 1966
(Fig. 9F-I)
Anthoarcuata Bakus, 1966: 431.
TYPE SPECIES. Anthoarcuata graceae Bakus,
1966:431 (by original designation) (holotype
USNM36284 (161848)).
Thickly encrusting, massive, cylindrical
growth form. Surface even, microscopically
hispid. Choanosomal skeleton with basal
renieroid reticulation composed of uni- or
paucispicular tracts (occasionally multispicular)
of acanthostyles joined at nodes by light collagen.
Peripheral ascending spicule tracts terminate in
brushes of smooth principal styles, originating in
choanosome and protruding through surface. Ectosomal skeleton has a dense, mostly tangential
crust of smooth subectosomal auxiliary styles (of
a single size category). Megascleres include
smooth ectosomal auxiliary styles-subtylostyles,
sometimes with mucronate bases and telescoped
points, and principal acanthostyles with even
spination forming basal skeleton. True echinating
megascleres absent. Microscleres palmate
isochelae with slight arcuate modifications (curvature, partial detachment of lateral alae from
shaft, slightly pointed teeth). Toxas absent.
REMARKS. The type species was originally
identified as Burtonancora lacunosa (Lambe,
1892) by de Laubenfels (1961: 195), but Bakus
(1966) noted that B. lacunosa (sensu de Laubenfels) was neither conspecific with Lambe's
species nor referable to Burtonanchora (which in
any case seems to be a synonym of Myxilla).
Anthoarcuata graceae has the same spicule
geometries and is structurally identical to Antho,
differing only in the supposed possession of arcuate rather than palmate isochelae. However, in
the type material chelae are not truly arcuate but
are predominantly palmate with some 'arcuate'
modifications including curvature and thickening
of the shaft, partial detachment of lateral alae
from the shaft (but for less than 20% of alar
length), and reduced, slightly pointed, slightly
unguiferous alae. They may be classed as palmate
on the basis that lateral alae are only partially
formed and are fused to the shaft for most of their
length.
Artemisina Vosmaer, 1885
(Fig. 10D-E)
Artemisina Vosmaer, 18856: 25; Ridley & Dendy,
1887: 112; Topsent, 1894: 12; Lundbeck, 1905:
110; Burton, 1930a: 501, 528-531; de Laubenfels,
^
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FIG. 10. Type species of microcionid genera. A-C, Antho (Myxilla involyens Schmidt, MNHNDCL636). D-E,
Artemisina (A. suberitoides, ZMAPOR443). F-G, Aulenella (A. foraminifera Burton & Rao, IM P I 167/1). H-I,
Axocielita (Microciona similis Stephens, RSME1921.143.1447).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
29
1936a: 117; Levi, 1960a: 61, 83; Ristau, 1978: 585;
Van Soest, 1984b: 122, 130; Bergquist & Fromont,
1988: 119.
Artenisina;Burton, 1934b: 54 [lapsus].
TYPE SPECIES. Artemisina suberitoides Vosmaer,
1885: 25 (by monotypy) (holotype ZMAPOR443);
junior synonym of Suberitesarciger Schmidt, 1870:47
(Burton, 1930a: 528) (schizotype BMNH1870.5.3.90).
Massive, subspherical growth form. Surface
skin-like, microscopically hispid, with few raised
oscules; texture distinctly stringy. Choanosomal
fibres indefinite or absent, overall architecture
plumo-reticulate, nearly halichondroid in places,
composed of multispicular ascending and
paucispicular transverse tracts of choanosomal
principal styles, bound together within spongin.
Echinating megascleres absent. Ectosomal
skeleton plumose, composed of single category
of subectosomal auxiliary styles forming discontinuous palisade of discrete brushes. Megascleres
smooth choanosomal principal subtylostyles and
smooth fusiform subectosomal auxiliary styles or
subtylostyles. Microscleres palmate isochelae,
and wing-shaped toxas with spinous extremities.
REMARKS. Defining Artemisina in phylogenetic terms is problematic. The taxon has no
real distinctive features, although it differs from
other Microcionidae in lacking a distinctive
choanosomal skeleton or definite spongin fibres
(also found in Qasimella Thomas), lacking
echinating spicules, and having a nearly radial
ectosomal skeleton reminiscent of some Ceratopsion (Raspailiidae; Hooper, 1991). These characteristics, two of which might be interpreted as
reductions or secondary losses and the third as a
convergence, are the only definable morphological apomorphies. Moreover, ectosomal structure
varies between several species, ranging from the
typical condition composed of erect brushes (e.g.,
A. arciger) to a tangential layer of spicules in
criss-cross fashion (e.g., A. melana Van Soest).
This variability is equivalent to (or analogues of)
the Mycale subgenera Carmia and Aegagropila,
respectively (e.g., Topsent, 1924b). At least one
species lacks a specialised ectosomal skeleton
completely (e.g., A. transiens Topsent).
Some species of Artemisina (e.g., A. foliata
(Bowerbank)) have honeycombed reticulate
growth forms, approaching the characteristic
Holopsamma morphology, but there is no consistency or pattern of gross morphologies amongst
Artemisina, and in any case it is unlikely that the
genus can be solely defined by its growth form.
De Laubenfels (1936a) and Ristau (1978b) suggested that the Artemisina was defined by the
absence of echinating acanthostyles and by the
presence of spinous extremities on toxas. Neither
character has much systematic value at the
generic level. In the present interpretation
echinating acanthostyles represent the retention
of an ancestral character, in which case their
presence or absence does not constitute a valid
reason to define a phylogenetic grouping, and in
any event they occur and disappear throughout
numerous microcionid and raspailiid taxa.
Similarly, toxas with spinous extremities are also
known to occur in many Microcionidae, including the type species of Clathria, C. cotnpressa
Schmidt, Microciona spinarchus Carter & Hope,
M. coccinea Bergquist, M. rubens Bergquist M.
spinatoxa Hoshino, Eutypon acanthotoxa
Stephens, Stylostichon toxiferum Topsent,
Labacea juncea Burton, Plocanzia ridleyi
Hentschel, and Ophlitaspongia thielei Burton.
They also occur in genera which have an ectosomal structure consistent with the Myxillidae
(e.g., Melonchela clathrata Koltun). Several species
of Artemisina have smooth toxas (e.g., A. melana),
and one (A. archegona Ristau) has oxeote toxas
similar to Paratenaciella Vacelet and Vasseur.
Thus, in the broad sense Artemisina contains a
heterogenous assemblage of species, which
prompted Burton (1930a) to divide the group into
three sections based on the number of megasclere
categories present. The simplest forms have only
one category of spicule (choanosomal principal
megascleres; e.g., A. transiens Topsent); the typical form has two categories of megascleres (larger
choanosomal principal styles and smaller subectosomal auxiliary styles; e.g., A. arciger); and the
third form has an incompletely differentiated
series of three megasclere types (two choanosomal spicules and one subectosomal spicule; A.
plumosa Hentschel). In all these forms species are
only really united in their lax choanosomal skeletal structure. The majority of species have been
described from Antarctic and Arctic regions.
[Aulena] sensu Lendenfeld, 1888
[Aulena]; Lendenfeld, 1888: 228, 1889a: 90; Topsent,
1894a: 19; de Laubenfels, 1936a: 16; Wiedcnmayer,
1989: 58.
Not Aulena Lendenfeld, 1885c: 309.
TYPE SPECIES. Aulena villosa Lendenfeld, 1885c:
309 (by subsequent designation; de Laubenfels, 1936a:
16) (syntypes AMZ130, G8901).
Bulbous, subspherical growth form, consisting
of fused digitate projections. Surface highly
papillose but not hispid. Choanosomal skeleton
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
regularly reticulate, with heavy spongin fibres
cored by sparsely scattered sand grains mostly at
nodes of fibres, and with a secondary fibre network between main skeleton. Ectosome lacks
sand cortex or any other mineral skeleton.
Megascleres and microscleres absent.
identical to Clathria (s.s.), lacking only toxa
microscleres. Vacelet et al. (1976: 75) correctly
synonymised Aulenella and Clathriopsamtna,
both of which now belong to C. (Wilsonella) (Van
Soest, 1984b; Wiedenmayer, 1989; Hooper &
Levi, 1993a).
REMARKS. Hallmann (1912:275) merged
Aulena and Echinoclathria Carter, whereas
Wiedenmayer (1989) noted that in the strict sense
(i.e., Lendenfeld, 1885c) it belongs to the Dictyoceratida (possibly related to Coscinoderma).
Conversely, most other species referred to Aulena
by Lendenfeld (1888) (A. laxa (Lendenfeld), A.
gigantea (Lendenfeld), A. crassa (Carter))
belong to Holopsamma.
Axocielita de Laubenfels, 1936
(Fig. 10H-I)
Aulenella Burton & Rao, 1932
(Fig. 10E-G)
Aulenella Burton & Rao, 1932:345.
TYPE SPECIES. Aulenella foraminifera Burton &
Rao, 1932: 345 (by original designation and
monotypy) (holotype IMP1167/1).
Flabello-digitate growth form, with subspherical, closely reticulate, honeycomb branching pattern. Surface uneven, not hispid. Choanosomal
skeleton irregularly reticulate, with spongin
fibres more-or-less fully cored by foreign particles and fewer choanosomal principal subtylostyles. Echinating acanthostyles abundant.
Ectosome with single size category of subectosomal auxiliary subtylostyles tangential to surface and also strewn randomly throughout
subectosome. Megascleres choanosomal principal
subtylostyles with spined and tuberculate bases,
echinating acanthostyles with evenly dispersed
large recurved spines, and subectosomal auxiliary
subtylostyles with or without spines on bases.
Microscleres palmate isochelae, toxas absent.
REMARKS. The 'honeycombed' reticulate
growth form of Aulenella is reminiscent of
Holopsamma and some E. (Echinochalina). The
incorporation of foreign particles into the
skeleton is also seen in species of Holopsamma,
Aulena of authors, Clathriopsamma,
Fisherispongia, Wilsonella and Anomoclathria.
It also occurs in other Poecilosclerida Raspailia
(Clathriodendron) (Raspailiidae), and many
species of Phoriospongiidae) and many Dictyoceratida, and is considered here to be a successful ecological adaptation independently
acquired by several groups. Thus,Aulenella is not
defined by any unique features, and furthermore
the holotype of A. foraminifera has spiculation
Axocielita de Laubenfels, 1936a: 118; Hechtel, 1965:
44; Wiedenmayer, 1977: 140.
TYPE SPECIES. Microciona similis Stephens, 1915:
441 (by original designation) (holotype RSME1921.143.1447).
Thickly encrusting growth form. Surface even,
and hispid. Choanosomal skeleton hymedesmoid, with spongin fibres forming basal layer on
substrate and ascending non-anastomosing fibre
nodes, each node containing plumose short unbranched tracts of choanosomal principal subtylostyles, standing perpendicular to substrate
with only bases embedded in spongin fibres.
Echinating subtylostyles erect on basal spongin
and also echinating erect plumose brushes of
choanosomal megascleres. Ectosomal skeleton
with single category of subectosomal auxiliary
subtylostyles forming paratangential tracts at surface and plumose brushes extending from ends of
choanosomal megascleres. Megascleres smooth
or minutely basally spined choanosomal principal subtylostyles, echinating subtylostyles with
only bases spined, subectosomal auxiliary subtylostyles mostly smooth, occasionally basally
spined, of a single size category. Microscleres
palmate isochelae and small wing-shaped toxas.
REMARKS. De Laubenfels (1936a) created
Axocielita for thinly encrusting sponges resembling Axociella (i.e., lacking spined echinating
megascleres). However, this is incorrect, based
on misconceptions of both the type species (i.e.,
the published characters of Microciona similis do
not agree with those seen in the holotype), and
Axociella by de Laubenfels' (1936a). In the
holotype of M. similis, diagnosed above, there is
only a single category of auxiliary spicule,
whereas Axociella s.s. has two distinct categories
of auxiliary spicules in the peripheral skeleton.
Thus, on the basis of its ectosomal features
Axociella is strictly a Thalysias, whereas
Axocielita has an unspecialised ectosomal
spiculation typical of Clathria. In having
plumose unbranched fibres forming a microcionid skeleton Hechtel (1965: 43) referred
Axocielita to Microciona. He supported this
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
31
decision on the basis that M. similis has both
spined and smooth styles, which he considered to
be diagnostic for Microciona (although in the
strict sense that feature is diagnostic for Anaata).
Axocielita is referred here to Clathria
(Microciona).
Axociella Hallmann, 1920
(Fig. 11A-B)
Axociella Hallmann, 1920:779; de Laubenfels, 1936a:
113; Wiedenmayer, 1977: 140; Bergquist &
Fromont, 1988: 116.
TYPE SPECIES. Esperiopsis cylindrica Ridley &
Dendy, 1886: 340 (by original designation) (holotype
BMNH1887.5.2.96).
Arborescent, dichotomously branched, stalked
growth form. Surface even, membraneous,
transparent, hispid. Choanosomal fibre skeleton
compressed at centre of stalk, with heavy spongin
fibres forming reticulate axis, with longitudinal
primary fibres cored by multispicular tracts and
secondary connecting fibres aspicular or
paucispicular tracts of choanosomal principal
styles. Echinating megascleres absent. Subectosomal extra-axial skeleton plumose or plumoreticulate, well-differentiated from axial core,
with light spongin fibres cored by multi- or
paucispicular larger subectosomal auxiliary
styles and few aspicular transverse fibres. Ectosomal skeleton composed of specialised
category of smaller ectosomal auxiliary styles
forming continuous palisade or individual
brushes of spicules. Megascleres entirely smooth
principal choanosomal styles-subtylostyles, and
two categories of entirely smooth auxiliary
styles-subtylostyles. Microscleres palmate
isochelae and oxhorn toxas.
REMARKS. Axociella has been misinterpreted
by all authors since it was first reviewed by de
Laubenfels (1936a). It is similar to Tenaciella
Hallmann in lacking echinating megascleres but
it has a distinctive, compressed reticulate axis and
plumose or plumo-reticulate subectosomal
(extra-axial) skeleton, reminiscent of the distinctive extra-axial skeletal architecture seen in
Raspailiidae. The structure of the ectosomal
skeleton in the type species of Axociella (i.e., the
possession of two categories of auxiliary
megascleres) is the same as seen in Thalysias, and
Van Soest (1984b) merged the two genera on this
basis. However, unlike typical species of
Thalysias or Clathria, those of Axociella have a
distinctive skeletal structure (compressed axis
and a radial extra-axial skeleton), which is more-
or-less homogeneous amongst the several known
species, and this structure is interpreted here as
indicative of supraspecific relationships. This interpretation is consistent with the treatment of
similar structures in Raspailiidae (Hooper, 1991).
In fact, Axociella could be justifiably included in
Raspailiidae (Hooper, 1991; Hooper et al., 1992),
apart from having isochelae and toxa microscleres
and lacking the unique raspailiid ectosomal
specialisation (large protruding auxiliary or principal spicules surrounded by bundles of smaller
auxiliary spicules). True examples of these ectosomal spicules are not seen in the Microcionidae,
although two species have analogues of this ectosomal condition: Esperiopsis canaliculata
Whitelegge, Ophlitaspongia thetidis Hallmann
— both belonging to Clathria (Axociella).
Axociella is convergent upon Raspailiidae in
skeletal structure, best developed in three
Australian species, Esperiopsis cylindrica, E.
canaliculata and Ophlitaspongia thetidis, all of
which also have a Thalysias ectosomal skeleton.
Another species from the NW Pacific,
Microciona lambei Koltun, has a mix of both
Raspailiidae and Microcionidae skeletons, being
convergent on Endectyon in structure (with a
markedly compressed axis, brushes of acanthostyles surrounding the bases of protruding
choanosomal principal styles) but it also has palmate isochelae and a tangential layer of auxiliary
styles lying tangential to the surface (i.e., the
Clathria condition). Axociella is recognised here
as a subgenus of Clathria based on its specialised
compressed skeletal structure.
Axosuberites Topsent, 1893
(Fig. 11C-D)
Axosuberites Topsent, 1893a: 179; de Laubenfels,
1936a: 118.
TYPE SPECIES. Axosuberitesfauroti Topsent, 1893a:
179 (by monotypy) (portion of holotype MNHNLBIMDT1859).
Flabellate, flattened digitate growth form. Surface hispid, conulose. Choanosome with compressed reticulate axis and plumose extra-axial
skeleton, with only light spongin fibres. Axial
fibres produce close-meshed reticulation of multispicular tracts cored by choanosomal principal
subtylostyles forming criss-cross reticulation,
tracts plumose near periphery. Echinating
megascleres absent. Subectosomal extra-axial
skeleton well differentiated from axial region,
with ascending plumose columns of larger subectosomal auxiliary subtylostyles arising from
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32zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 11. Type species of microcionid genera. A-B,Axociella(A. cylindrica Ridley & Dendy, BMNH1887.5.2.96).
C-D, Axosuberites (A. fauroti Topsent, MNHNDT1859). E-G, Bipocillopsis (B. nexus Koltun,
BMNH1963.7.29.56). H-I, Clathriella (C. primitiva Burton, BMNH1938.7.4.93).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
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peripheral choanosomal skeleton. Ectosome with
brushes of smaller auxiliary subtylostyles overlaying larger subectosomal spicules. Megascleres
entirely smooth choano somal principal subtylostyles-tylostyles, and two size classes of auxiliary
subtylostyles-tylostyles, both with smooth bases.
Microscleres absent.
REMARKS. Redescription of the type species
above is based on examination of a slide preparation in the MNHN collection and Topsent's
(1893a) description; the corresponding specimen
has not yet been seen. Topsent (1893a) initially
compared Axosube rites with Caulospongia Kent
and Pseudosuberites Topsent in Suberitidae
(Hadromefida), noting that it showed a certain
level of morphological similarity in choanosomal
fibres and geometry of tylostyle megascleres.
However, in skeletal structure and spiculation
Axosuberites appears to be a microcionid, most
similar to Axociella, both genera showing
similarities in their ectosomal specialisation,
axial and extra-axial differentiation, and absence
of echinating megascleres. Van Soest (1984b)
also suggested that the genus was probably an
Axociella without microscleres, and under his
scheme it was therefore referable to Thalysias,
but in both genera axial and extra-axial skeletal
structures are well developed and closely comparable, and it is suggested here that they both
should be included in the same subgenus (i.e.,
Clathria (Thalysias)).
Bipocillopsis Koltun, 1964
(Fig. 11E-G)
Bipocillopsis Koltun, 1964a: 79.
TYPE SPECIES. Bipocillopsis nexus Koltun, 1964a:
80 (by monotypy) (holotype ZIL10644, paratype
BMNH1963.7.29.56).
Erect, arborescent growth form with cylindrical
reticulated branches. Surface hispid, raised into
irregular sharp conules. Choanosomal skeleton
subrenieroid reticulate, with ascending spongin
fibres cored by multispicular plumose tracts and
interconnected by paucispicular transverse tracts
of choanosomal principal subtylostyles, and
echinated by acanthostyles. Subectosomal region
with heavy paratangential bundles of subectosomal auxiliary styles protruding through surface and also scattered between fibres. Ectosomal
skeleton without specialised spiculation but with
bundles of subectosomal auxiliary styles surrounding protruding choanosomal spicules, and
also lying paratangential to ectosome.
Megascleres choanosomal principal subtylostyles invariably with basal spines and sometimes
with spines on shaft, echinating acanthostyles of
similar morphology to principal megascleres but
shorter and more extensively spined, and single
category of entirely smooth subectosomal
auxiliary style. Microscleres isochelae, strongly
sigmoid, unguiferous with very reduced pointed
alae, possibly anchorate. Toxas absent.
REMARKS. Bipocillopsis resembles Damoseni
in lacking ectosomal specialisation and having
modified sigmoid isochelae, but differs in growth
form (arborescent versus encrusting) and subectosomal skeletal architecture (thickly paratangential versus tangential). The genus is
monotypic and may be included in Clathria
(Clathria) by its skeletal structure, whereas
Damoseni has hymedesmoid skeletal structure
typical of Clathria (Microciona). Chelae morphology has been described as tridentate sigmoid,
allegedly related to the arcuate form, but this is
very difficult to tell with certainty given that alae
are nearly vestigial. There is no doubt that these
chelae are more highly modified than most other
microcionids. They are very small, strongly unguiferous (sigmoid curved with small pointed
teeth), and lateral alae are completely free of the
shaft and undifferentiated from the front ala, suggesting they may be of anchorate origin. However, there is no lateral ridge on the shaft of chelae
to indicate an anchorate condition. The derivation
of these chelae is indeterminable. Under the
scheme of Hajdu et al. (1994) this genus could be
included in Myxillina in possessing tridentatederived isochelae, whereas all other features
(skeletal structure, principal and auxiliary
megascleres and ectosomal skeleton) indicate
relationships with the Microcionidae, in which it
is retained here tentatively. This decision is supported by the case of Damoseni, discussed below,
which have chelae of identical form to Bipocillopsis with the addition of oxhorn toxas (which
are not found in Myxillina).
Cionanchora de Laubenfels, 1936
(Fig. 12E-G)
Cionanchora de Laubenfels, 1936a: 108.
TYPE SPECIES. Hymeraphia tuberosocaphata Topsent, 1890b: 68 (by original designation) (fragment of
holotype MNHNLBIMDT939).
Encrusting growth form. Surface smooth, even,
microscopically hispid. Choanosomal skeleton
hymedesmoid. Spongin fibres reduced to basal
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34zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
layer lying on substrate, with choanosomal principal subtylostyles erect and forming unispicular
ascending columns protruding through ectosome,
and echinating acanthostyles standing parallel to
these. Ectosome tangential skeleton of subectosomal auxiliary subtylostyles, of a single
category, forming brushes surrounding protruding choanosomal principal spicules. Megascleres
choanosomal principal subtylostyles with tuberculate bases, echinating acanthostyles with large
spines evenly dispersed over entire spicule except
for bare point, and subectosomal auxiliary subtylostyles completely smooth. Microscleres
anchorate-like isochelae. Toxas absent.
REMARKS. The diagnosis is based on a slide of
the holotype and Topsent's (1890b) description;
the corresponding specimen has not been seen.
Cionanchora was erected for thinly encrusting
sponges with hymedesmoid skeletal construction, similar to Anaata, but with anchorate-like
instead of arcuate-like modifications to isochelae
(de Laubenfels, 1936a). Both those genera were
merged with Clathria (s.1.) by Van Soest (1984b)
on the basis that modified microscleres were a
homoplasy throughout the Poecilosclerida, and
are interspecific discriminators only. Anchoratelike modifications are also found in Folitispa
(both of which were included in Clathria by
Hooper, 1990a). Microciona dubia from
Christmas Island (Kirkpatrick, 1900a: 136), was
referred to Cionanchora by de Laubenfels
(1936a: 108) supposedly in having anchorate-like
chelae but these are of palmate origin. Both
species have skeletal architecture typical of
Microciona where they are referred.
Clathria Schmidt, 1862
(Fig. I2A-B)
Clathria Schmidt, 1862: 57; Ridley, 1884a: 443-449,
612-615; Ridley & Dendy, 1887: 31; Hentschel,
1911: 368; Hallmann, 1912: 205; Dendy, 1922: 64;
Dendy, 1924a: 352-354; Wilson, 1925: 439; Topsent, 1925: 645-658; Topsent, 1928a: 62,299; Burton & Rao, 1932: 334-337; Burton, 1932a: 319;
Burton, 1934a: 558; Koltun, 1959: 184; Levi,
1960a: 50,52,61; Melone, 1963: 1-8; Sara &
Melone, 1963: 362; Sara, 1964: 229; Simpson,
1968a: 102, 104-106; Van Soest, 1984b: 90;
Wiedenmayer, 1989: 56; Bergquist & Fromont,
1988: 106.
Clathria Schmidt, plus Rhaphidophlus Ehlers; Ridley
& Dendy, 1887: 146,151; Topsent, 1894a: 14-15,18.
Clatharia;Kumar, 1925: 221 [lapsus].
TYPE SPECIES. Clathria compressa Schmidt, 1862:
58 (by subsequent designation (Schmidt, 1864: 35))
(holotype LMJG15509).
Erect, arborescent, thinly lamellate, branching
growth form. Surface even, not hispid.
Choanosomal skeleton regularly reticulate, with
well developed spongin fibres forming regular or
irregular anastomoses of differentiated primary
and secondary spongin fibres. Fibres cored by
choanosomal principal subtylostyles in multispicular ascending tracts and uni- or bispicular
transverse connecting tracts, and echinated by
acanthostyles perpendicular to or at acute angles
to spongin fibres. Ectosomal skeleton with tangential layer of subectosomal auxiliary subtylostyles, of a single size category. Megascleres
basally spined choanosomal principal subtylostyles, entirely smooth subectosomal auxiliary
subtylostyles, and echinating acanthostyles with
even spination. Microscleres palmate isochelae
and forceps-shaped or accolada toxas with
spinose extremities.
REMARKS. Strictly defining Clathria is essential in assigning a vast number of microcionids
included in the genus by numerous authors. For
example, C. compressa has toxas with spinose
extremities, which therefore becomes a character
'typical' of Clathria, whereas earlier authors considered that this feature was diagnostic for Artemisina Vosmaer (de Laubenfels, 1936a).
Clathria-like (viz. Labacea de Laubenfels) and
Artemisina-like genera (viz. Ligrota de Laubenfels) also have spinous toxas, indicating that this
character is homoplasious and probably not important above the species level of classification.
In general, the original definition of Clathria
(s.s.) is upheld here, as re-examination of
Schmidt's syntype confirmed that all published
characters cited in the species description
(Schmidt, 1862: 58; Topsent, 1925: 647) are
present in type material.
Topsent (1925: 648) noted that C. compressa
has variable spicule dimensions, skeletal architecture, and live colouration. He correlated
this variability with the diverse growth forms
shown by the species: thinly encrusting examples
had a hymedesmoid skeletal architecture, thickly
encrusting forms had a plumose skeleton, and
erect ramose forms had anastomosing fibres (i.e.,
encompassing the nominal genera Leptoclathria,
Microciona and Clathria). He also found correlation between the size of megascleres and growth
form variability, although he could find no obvious trends. Topsent's observations are invaluable in deciding whether to maintain nominal
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
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35
encrusting genera and more massive sponges as
distinct genera, and whether to differentiate taxa
with hymedesmoid, plumose or plumo-reticulate
skeletal structure.
Clathria compressa is known only from the
north Atlantic and Mediterranean regions, but
judging by its extensive synonymy it appears to
be (or have been) moderately common within
those regions.
Clathriella Burton, 1935
(Fig. 11H-I)
Clathriella Burton, 1935c: 73; Koltun, 1959: 186.
TYPE SPECIES. Clathriella primitiva Burton, 1935c:
73 (by original designation) (holotype
BMNH1938.7.4.93).
Crumpled, irregular, massive growth form.
Surface porous, uneven, hispid, with meandering
ridges covered by thin transparent dermal
membrane. Choanosomal skeleton renieroid
reticulate, with spongin fibres forming regularly
triangular meshes cored by multispicular tracts of
smaller principal rhabdostyles (confined to
renieroid network) and larger principal styles (latter producing secondary plumose, subisodictyal
skeleton of bi- or paucispicular ascending tracts).
Echinating spicules absent. Ectosomal skeleton
radially arranged tracts of subectosomal auxiliary
styles, of a single size category. Extra-fibre
skeleton (apparently) has centrally curved (arcuate) oxeote megascleres scattered throughout
mesohyl. Megascleres two categories of
choanosomal principal styles, larger smooth with
rounded bases, smaller spined with rhabdose
bases, and smooth or basally spined subectosomal auxiliary styles-subtylostyles.
Microscleres absent.
REMARKS. Clathriella primitiva is obviously
closely related to Clathria chartacea Whitelegge
in its skeletal structure and spiculation, conforming to the definition of Isopenectya (see below).
Clathriella also shows similarities to Isociella in
having an isodictyal reticulate skeleton and in
lacking echinating spicules, although in Isociella
all megascleres are smooth and chelae and toxa
microscleres are present. The presence of smaller
spined rhabdostyles and an isodictyal component
of the choanosomal skeleton are reminiscent of
Rhabderemiidae, although the absence of thraustosigmata, thraustotoxa and other rhabderemiid
microscleres in C. primitiva suggest that these
similarities are convergent.
Burton (1935c) considered that Clathriella was
a primitive member of the Microcionidae, in
which styles and acanthostyles had not yet become differentiated or segregated into coring and
echinating megascleres. He suggested further
that the toxiform oxeas, recorded by both Burton
(1935c) and Koltun (1959), were derived from
acanthostyles, and that both forms were remnants
of a primitive condition. There is no empirical
evidence to support either argument, and the
present study takes the alternative point of view,
that species like Clathriella prinntiva are derived
or modified microcionids. Koltun (1959) also
suggested that arcuate oxeas of C. primitiva were
microxeas, and in that respect the genus should
be compared with Paratenaciella. However, in
Clathriella these spicules are supposedly large
(200x7 p.m), indicating that they are true
megascleres, whereas in Paratenaciella
microxeas are very small (40-75 x 0.7-311m). In
any case, re-examination of the holotype of C.
primitiva (above) and more recent material from
the Sakhalin Is, NW Pacific collected by PIBOC
(QMG300052), did not find any toxiform oxeas
although several examples of smaller auxiliary
styles were sinuous, and it may be these the
authors were referring to. The genus is referred
here into synonymy with Atitho (lsopenectya).
Clathriopsamma Lendenfeld, 1888
(Fig. 12C-D)
Clathriopsamma Lendenfeld, 1888: 227; Topsent,
1894a: 19; Hallmann, 1920: 771; de Laubenfels,
1936a: 98; Levi, 1973: 614; Vacelet et al., 1976: 75.
TYPE SPECIES. Clathriopsamma reticulata Lendenfeld, 1888: 227 (by subsequent designation; Hallmann,
1920: 771) (lectotype AMG9 1 35).
Erect, anastomosing, arborescent growth form.
Surface uneven, arenaceous, microscopically
hispid. Choanosomal skeleton irregularly reticulate. Spongin fibres without well developed
primary or secondary differentiation, cored by
choanosomal principal subtylostyles and abundant foreign debris. In subectosomal skeleton
principal subtylostyles also form plumose
brushes, protruding through ectosome. Fibres
heavily echinated by acanthostyles also associated with ectosomal spicule brushes. Ecto some with paratangential tracts of
subectosomal auxiliary subtylostyles, of a single
category, usually forming discrete brushes of
spicules at surface. Megascleres basally spined,
fusi form choanosomal principal subtylostyles,
smooth and basally spined subectosomal
^
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36zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 12. Type species of microcionid genera. A-B, Clathria (C. compressa Schmidt, LMJG15509). C-D,
Clathriopsamma (C. reticulata Lendenfeld, AMG9135). E-G, Cionanchora (Hymeraphia tuberosocapaata
Topsent, MNHNDT939). H-1, Colloclathria (C. ramosa Dendy, BMNH1921.11.7.64).
REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
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37
auxiliary subtylostyles, and echinating acanthostyles with large spines evenly dispersed.
Microscleres palmate isochelae of two size
categories, including contort forms, and sinuousraphidiform or accolada toxas.
REMARKS. In Clathriopsamma the ectosome is
structurally close to the Thalysias condition, but
there is only one category of auxiliary spicule
producing surface bundles. The genus is distinguished from most other microcionid genera in
having foreign particles incorporated into the
skeleton (Hallmann, 1920) (see remarks for
Aulenella). Detrital entrapment is also known to
occur in other Microcionidae (e.g., some Holopsamma), other Poecilosclerida (e.g., Phoriospongiidae), and it certainly also occurs commonly in
other sponge orders (Dysideidae, Thorectidae
and Ircinidae in the Dictyoceratida, and many
Haplosclerida). This evidence indicates that this
feature has arisen independently several times
within the Porifera, probably indicative of
ecological specialisation, and obviously arisen
independently in several groups. However,
species of Microcionidae that do incorporate sand
appear to be relatively homogeneous in most of
their other characters (i.e., there are no other
conflicting characters such as presence/absence
of ectosomal specialisation, or modifications to
chelae), and consequently this specialisation is
recognised here at the subgenus level. Vacelet et
al. (1976) synonymised Clathriopsamtna with
Aulenella, and Van Soest (1984b) merged
Clathriopsatnma with Clathria, whereas in this
work it is shown that the earliest available name
for these species is Wilsonella (see below).
Colloclathria Dendy, 1922
(Fig. 12H-I)
Colloclathria Dendy, 1922: 74.
TYPE SPECIES. Colloclaihria ramosa Dendy, 1922:
74 (by monotypy) (holotypeBMNH 1921.11.7.64).
Cylindrical, arborescent, digitate growth form.
Surface even, hispid. Choanosomal skeleton
reticulate, with slightly compressed axis and
slightly more cavernous extra-axial skeleton.
Spongin fibres cored by large principal
choanosomal styles, forming multispicular ascending tracts interconnected by multispicular
transverse tracts. Echinating acanthostyles distributed irregularly over fibres. Subectosomal
skeleton reduced to immediate outer edge of
skeleton, with plumose tracts of larger subectosomal auxiliary subtylostyles extending
through ectosome. Larger auxiliary megascleres
also scattered between fibres and sometimes
coring fibres. Ectosomal skeleton with smaller
ectosomal auxiliary subtylostyle forming discrete
brushes overlying subectosomal spicules.
Megascleres smooth principal choanosomal
styles, echinating acanthostyles with spined
bases and points (bare 'necks'), and two sizes of
auxiliary subtylostyles, usually with basal spines.
Microscleres palmate isochelae, cleistochelae
and accolada toxas.
REMARKS. Colloclathria has a specialised ectosomal identical to Thalysias with two
categories of auxiliary spicules forming surface
spicule bundles, and on that basis Van Soest
(1984b: 115) suggested the two genera should be
merged: it is included here in synonymy with
Clathria (Thalysias). The possession of cleistochelae in C. ramosa is not unique to Microcionidae also found in Plocamiopsis, Quizciona
and several species of Clathria (e.g., C.
(Clathria) toxipraedita).
Damoseni de Laubenfels, 1936
(Fig. 13A-C)
Damoseni de Laubenfels, 1936a: 110.
TYPE SPECIES. Hymeraphia michaelseni Hentschel,
1911:351 (by original designation) (fragment of
holotype SMF969T).
Encrusting growth form. Surface sparsely
hispid, even. Choanosomal skeleton hymedesmoid, with spongin fibres reduced to basal layer
of spongin lying on substrate, with bases of
choanosomal principal subtylostyles and acanthostyles embedded in spongin, standing perpendicular to substrate, spicules protruding through
ectosome. Ectosomal skeleton with paratangential, slightly plumose tracts of both larger and
smaller auxiliary subtylostyles. Megascleres
large principal choanosomal subtylostyles with
spined bases, echinating acanthostyles with
spined bases and shafts (bare 'necks'), and two
size classes of subectosomal auxiliary subtylostyles with either smooth or spined bases.
Microscleres strongly unguiferous, sigmoid
isochelae with vestigial alae, arcuate- or anchoratelike but of uncertain affinity, and large wing-shaped
(oxhorn derived) toxas.
REMARKS. Hentschel's (1911) description of
Hymeraphia michaelseni does not mention the
presence of toxa microscleres, nor that principal
spicules protrude a long distance through the
surface. The modified unguiferous isochelae (of
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possible anchorate or arcuate derivation) are
identical to those of Bipocillopsis, and the combination of toxas and unguiferous isochelae of
possible tridentate-derivation supports the inclusion of such taxa in Microcionidae contrary to
Hajdu et al.'s (1994) proposal. Damoseni is
similar to other encrusting genera with hymedesmoid architecture and it could be included in
Clathria (Microciona) on this basis. However, the
specialised ectosomal skeleton composed of
smaller ectosomal and larger subectosomal
auxiliary spicules indicates affinities to Clathria
(Thalysias).
Dendrocia Hallmann, 1920
(Fig. 13D-E)
Dendrocia Hallmann, 1920: 767; de Laubenfels,
1936a:109.
TYPE SPECIES. Clathria pyramida Lendenfeld,
1888: 222 (by original designation) (holotype
AMG9047).
Massive, lobate-digitate growth form. Surface
conulose, with subdermal sculpturing, oscules
slightly raised above surface with membraneous
lip. Choanosomal skeleton dendritic, slightly
plumo-reticulate near axis, without any obvious
division of primary or secondary fibres, but with
clear structural differences between choanosomal, subectosomal and ectosomal regions.
Spongin fibres heavy, meandering, cored by
auxiliary styles indistinguishable from those in
ectosomal skeleton, and heavily echinated by acanthostyles (sometimes also secondarily incorporated into fibres). Subectosomal skeleton
plumose or radial, with spicule tracts composed
of auxiliary styles diverging from ends of
peripheral choanosomal fibres and supporting
overlying ectosomal skeleton. Ectosomal skeleton
with continuous, heavy palisade of erect auxiliary
styles. Principal spicules absent, and megascleres
include only echinating acanthostyles with spined
bases and necks but smooth points, and single
category of structural spicule (auxiliary stylessubtylostyles), entirely smooth with hastate points
and sometimes secondarily pointed bases. Micro
scleres modified palmate isochelae with curved,
thickened shaft and reduced alae. Toxas absent.
REMARKS. This strict diagnosis based on the
type species should be widened to include the
presence of modifications to isochelae, ranging
from typical palmate forms (with straight shaft,
completely fused lateral alae, as in Clathria
(Dendrocia) abrolhensis sp. nov.), modified palmate isochelae (with greatly curved, thickened
shaft, partially detached lateral alae, as in C. (D.)
dura), to anchorate-like forms (in which lateral
alae are detached from shaft, shaft is curved,
thickened and has a lateral ridge, as in C. (D.)
myxilloides). One species also has oxhorn toxas
(C.(D.) scabida) supporting the hypothesis that
Dendrocia has affinities with Microcionidae.
Dendrocia, like Wilsonella, differs from other
microcionids in having auxiliary styles both
coring fibres and forming the ectosomal skeleton.
However, whereas Wilsonella has two categories
of auxiliary spicules and detritus is incorporated
into the skeleton, Dendrocia has only one
category of structural spicule throughout the
skeleton. Dendrocia also has a characteristic
dendritic or plumo-reticulate skeletal architecture, whereas Wilsonella is invariably reticulate.
In ectosomal skeletal structure (with a continuous
ectosomal palisade of spicules) Dendrocia
resembles the Thalysias condition, but it has only
one geometric form of auxiliary spicule producing the extra-fibre skeleton (i.e., subectosome
and ectosomal regions) more similar to Clathria.
For this reason Dendrocia is enigmatic, and
Hooper (1990a) maintained it as a separate taxon,
whereas earlier Van Soest (1984b) had indicated that it was probably a synonym of
Clathria. In this work it is referred to Clathria
(Dendrocia).
Two Australian species were initially included
in Dendrocia by Hallmann (1920), the type
species and C. alata Dendy (holotype NMV
G2280). Both are undoubtedly synonyms. They
apparently differ only in their spicule dimensions
(styles: 175-230x2-p,m versus 240-250x4-p,m;
acanthostyles: 63-95x3-1p.m versus 79-154x81p.m; arcuate isochelae: 23-2p.m versus 20-2p,m,
respectively). In contrast, D. antyaja Burton &
Rao, from the Indian Ocean, should be removed
from Dendrocia because it has regularly reticulate choanosomal architecture, lacks an ectosomal
skeleton, and has palmate isochelae indicating
affinities to Clathria. Several other Australian
species previously referred to Clathria s.s. should
also be referred to Dendrocia on the basis of their
skeletal architecture and spiculation.
Dictyociona Topsent, 1913
(Fig. 13F-G)
Dictyociona Topsent, 1913a: 579, 618; de Laubenfels,
1936a: 110; Levi, 1960a: 60; Bergquist & Fromont,
1988: 104.
TYPE SPECIES. Microciona discreta Thiele, 1905:
447 (by monotypy) (holotype ZMB3302).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
39
Thickly encrusting, lobate growth form. Surface composed of interconnected micropapillae.
Choanosomal skeleton plumose, becoming
plumo-reticulate in thicker sections. Spongin
fibres divided into primary and secondary elements, heavier in axis, lighter in subectosomal
region where dermal spicules implanted at nodes
of peripheral fibres. Fibres cored by multispicular
tracts of choanosomal principal subtylostyles,
and heavily echinated by acanthostyles projecting from fibres at all angles. Ectosomal skeleton
composed of paratangential tracts of subectosomal auxiliary subtylostyles, of a single size
category, arising from peripheral fibres and piercing surface. Megascleres choanosomal principal
subtylostyles with acanthose bases and acanthose
shafts near basal end, echinating acanthostyles
with spined bases and points (bare 'necks'), and
subectosomal auxiliary subtylostyles with
microspined bases. Microscleres palmate
isochelae and thin toxas intermediate between
wing-shaped and oxhorn forms.
REMARKS. Dictyociona resembles Microciona
in its plumose skeletal structure but has partially
acanthose choanosomal megascleres similar to
those found in Anaata and Antho. These spicules
are also present in several Raspailiidae (e.g.,
Hymeraphia, Eurypon), and they are considered
here to be of minor diagnostic importance above
the species level following Simpson (1968a). Levi
(1960a: 60) merged Dictyociona with Clathria, although the species could as easily be placed in
Microciona. It is included here within Clathria
(Clathria). Apart from the type species, other taxa
referred to Dictyociona at one time or another
include: Microciona clathrata Whitelegge, M.
heterotoxa Hentschel, M. pyramidalis Brondsted,
Hymedestnia oxneri Topsent, Eurypon asodes de
Laubenfels, E. acanthotoxa Stephens, E. ditoxa
Stephens, E. microcizela Stephens, E. tenuissima
Stephens, Clathria terra novae Dendy, D. contorta Bergquist & Fromont and D. atoxa Bergquist
& Fromont.
Dictyoclathria Topsent, 1920
(Fig. 13H-I)
Dictyoclathria Topsent, 1920b: 18; Burton, 1930a:
501, 533, 1933: 50; de Laubenfels, 1936a: 77; Levi,
1960a: 80.
Dyctioclathria [lapsus]; Ferrer Hernândez, 1921: 172.
TYPE SPECIES. Clathria morisca Schmidt, 1868: 9
(by original designation) (schizotype BMNH1868.3.2.21); junior synonym of Antho involvens
(Schmidt, 1864) (Levi, 1960a: 57) (schizotype
BMNH1867.3.11.92).
Arborescent growth form. Surface hispid, uneven. Choanosomal skeleton renieroid reticulate
composed of spined acanthostyles forming triangular or rectangular meshes, enclosed within
fibres (axis) or bound together at nodes by collagen (near periphery). Echinating megascleres
absent. Subectosomal (extra-axial) skeleton
plumose, with smooth principal styles standing
perpendicular to fibre nodes, individually or in
bundles, protruding through surface. Ectosomal
skeleton with dense plumose or paratangential
bundles of subectosomal auxiliary subtylostyles,
sometimes surrounding protruding principal
styles. Megascleres shorter lightly spined acanthostyles (renieroid skeleton) and longer completely smooth principal styles (extra-axial
skeleton), and smaller subectosomal auxiliary
subtylostyles with spined bases. Microscleres
palmate isochelae and toxas intermediate between wing-shaped and oxhorn forms.
REMARKS. Dictyoclathria is an objective
synonym of Antho, since the type species of both
genera are conspecific (Levi, 1960a). The type
specimen of Dictyoclathria is remarkable in having
a nearly raspailiid ectosomal condition with
bundles of ectosomal spicules appearing to surround the protruding extra-axial spicules, but this
is not as perfectly developed as in many Raspailia.
Dirrhopalum Ridley, in Ridley & Duncan,
1881
Plocamia Schmidt, 1870: 62.
Dirrhopalum Ridley in Ridley & Duncan, 1881: 477.
See Plocamia.
Echinochalina Thiele, 1903
(Fig. 14A-B)
Echinochalina Thiele, 1903a:961; Hallmann, 1912:
288; Topsent, 1928a: 61; Burton, 1934a: 562; de
Laubenfels, 1936a:118; Thomas, 1977: 115,
TYPE SPECIES. Ophlitaspongia australiensis Ridley,
1884a: 442 (by subsequent designation (Hallmann,
1912: 288)) (holotype BMNH1881.10.21.299).
Massive-digitate, reticulate growth form. Surface with low ridges and interconnected lamellae,
producing angular cells with thin dermal
membrane stretched between. Choanosomal
skeleton irregularly reticulate, with heavy
primary spongin fibres cored by paucispicular
tracts of auxiliary spicules (tornotes), and heavy
secondary spongin fibres cored by uni- or
^
MEMOIRS OF THE QUEENSLAND MUSEUM
40zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
-
.01k1...
FIG. 13. Type species of microcionid genera. A-C, Damoseni (D. michaelseni Hentschel, SMF969T). D-E,
Dendrocia (Clathria pyramida Lendenfeld, AMG9047). F-G, Dictyociona (Microciona discreta Thiele,
ZMB3302). H-I, Diayoclathria (Clathria morisca Schmidt, MNHNDT unreg.).
^
41
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
bispicular tracts of auxiliary spicules. Fibres
lightly echinated by principal subtylostyles. Ectosomal skeleton with undulating fibres and loose
paratangential tracts of auxiliary spicules (also
distributed throughout mesohyl). Megascleres include quasidiactinal auxiliary tornotes with aymmetrical or symmetrical ends (coring fibres and
on ectosome), and completely smooth principal
subtylostyles-styles (echinating fibres), sometimes modified to oxeotes (asymmetrical with
two pointed ends). Microscleres wing-shaped
toxas. Isochelae absent.
REMARKS. In the type species coring spicules
are tornotes, whereas in other species of
Echinochalina they range from true styles to
oxeotes. Re-examination of the holotype of 0.
australiensis, and Thiele's (1903a) specimen
from Ternate, Indonesia (SMF1855) also found
that echinating principal styles can sometimes be
modified to oxeote spicules, providing support
for the otherwise tenuous placement of Protophlitaspongia with this group of microcionids.
Hallmann (1912: 288) noted that Echinochalina
differs from the allied Echinoclathria of authors
(= Holopsamma as defined here) in having fibres
cored by auxiliary spicules (tylotes or strongyles), which may be modified to quasi-monactinal forms (tylostrongyles, tornostrongyles), and
echinated by smooth styles or subtylostyles. By
comparison, Echinoclathria in the strict sense (or
Ophlitaspongia of authors) has principal
choanosomal styles (or modified monactinal
megascleres) which both core and echinate fibres,
as well as a second category protruding through
the surface; Holopsamma has only a single
category of coring and echinating principal
spicule. As such, Van Soest (1984b: 129) suggested that Echinochalina possibly did not
belong with the Microcionidae, and might be
more appropriately placed elsewhere within the
Poecilosclerida (e.g., Phoriospongiididae). However, examination of a suite of species included
here in Holopsamma found that this group also
undergoes a reduction in coring spicules (e.g., H.
pluritoxa (Pulitzer-Finali)), whilst retaining other
characteristics common to the genus, and it is
possible that Echinochalina, sensu Hallmann, undergoes similar reduction.
Probably of greater importance in determining
the appropriate placement of Echinochalina is its
lack of any special ectosomal skeleton, and in this
respect it is similar to Dendrocia (both with only
one form of structural (auxiliary) spicule). Whereas
Echinoclathria (s.s.) possesses monactinal auxiliary
spicules in the peripheral skeleton (sometimes
reduced to quasi-diactinal forms), Echinochalina
has spicules which are closer to true diactinals
(although sometimes modified to quasi-monactinal forms). Both genera lack definite ectosomal
specialisation. Some Echinochalina appear to have
affinities with certain species of Echinodictyum
(Raspailiidae) and with the Desmacididae, and
both Thiele (1903a) and Topsent (1904a) have
already noted this resemblance, suggesting that
they differ mainly in the geometry, ornamentation
and derivation of the echinating megascleres. By
comparison, Hallmann (1912) considered that
these differences, and the presence of long subectosomal styles in many Echinodictyum species,
are sufficient to maintain the two genera as distinct taxa (see Hooper, 1991d).
Although Echinochalina usually has smooth
echinating megascleres, linking it to
Echinoclathria and Holopsamma, there are two
species which were previously referred to
Echinodictyum (E. ridleyi Dendy and E. spongiosum Dendy), which have acanthose echinating
megascleres but otherwise conform to
Echinochalina in their spicule geometries and
skeletal architectures. Hooper (1991) transferred
these species to Echinochalina.
Tablis de Laubenfels is an obvious synonym of
Echinochalina. The two genera differ only by the
absence of microscleres and the presence of a
reticulate architecture in Tab/is.
Echinoclathria Carter, 1884
(Fig. 14C-E)
Echinoclathria Carter, 1884: 204 [nomen nudum];
Carter, 1885E 355; Ridley & Dendy, 1887: 159;
Topsent, 1894a: 18; Thiele, 1903a: 962; Hallmann,
1912: 275-276; Dendy, 1922: 71 [in part]; Topsent,
I928a: 61; de Laubenfels, 1936a: 119; Wiedenmayer, 1977: 143, 1989: 58. (not Uriz, 1988: 89].
TYPE SPECIES. Echinoclathria tenuis Carter, 1885E
355 (by subsequent designation; Burton, 1934a: 562)
(holotype BMNH1886.12.15.147); junior synonym of
Spongia leporina Lamarck, 1814: 444 (Topsent, 1932:
101) (holotype MNHNLBIMDT567).
Thinly flabellate, flattened palmate, stalked
growth form. Surface membraneous, microscopically hispid. Choanosomal skeleton renieroid
reticulate, slightly compressed with well
developed spongin fibres in axis, more openly
reticulate, less compressed and with lighter
spongin towards periphery. Axial fibres cored
by pauci- or multispicular tracts of smaller principal styles producing rounded or irregularly
shaped meshes, and echinated by same spicules.
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MEMOIRS OF THE QUEENSLAND MUSEUM
42zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Subectosomal (extra-axial) skeleton radial, unior paucispicular, with larger principal spicules
erect on terminal fibres and usually protruding
through surface. Ectosomal specialisation absent,
with bundles of subectosomal auxiliary subtylostyles embedded perpendicularly and forming
paratangential brushes surrounding larger principal spicules. Megascleres include smaller,
robust, entirely smooth principal subtylostyles
(coring and echinating fibres), larger principal
subtylostyles of similar geometry (projecting from
peripheral fibres and protruding through surface),
and smooth subectosomal auxiliary subtylostyles, straight or flexuous. Microscleres absent.
REMARKS. There is substantial confusion concerning the precise definition of Echinoclathria,
and its relationship with other nominal genera
such as Holopsamma, Halme, Aulena and Ophlitaspongia. Consequently the above diagnosis is
strict, pertaining only to the type species, and a
detailed explanation is justified below.
Most authors follow Hallmann's (1912: 275)
interpretation of Echinoclathria in which the
genus is essentially characterised by 'a
honeycomb mass of anastomosing flattened
trabeculae', a reticulate skeleton of heavy spongin fibres cored and echinated by smooth monactinal principal megascleres of the same geometry,
and with monactinal subectosomal auxiliary
styles distributed throughout the mesohyl (and in
some species also forming a radial subectosomal
skeleton). Hallmann also noted that in some
species he assigned to Echinoclathria there are
both chelae and toxa microscleres, with quasimonactinal auxiliary megascleres, or they may
have their coring megascleres replaced partially
or completely by detritus (e.g., Holopsamma
laminaefavosa). Hallmann suggested further that
Echinoclathria and Ophlitaspongia essentially
differed only in growth form, a view perpetuated
by Wiedenmayer (1989). This interpretation is
emended here.
Most species included in Echinoclathria prior
to the present study do have the characteristics
outlined by Hallmann (1912), and most are relatively homogeneous and easily recognisable in
the field by their characteristic 'honeycomb
reticulate' growth form. It is therefore unfortunate that Burton (1934a) subsequently designated E. tenuis (a junior synonym of Spongia
leporina) as the type species of Echinoclathria,
because this species has a flabellate growth form
(very dissimilar to 'honeycombed reticulate'
species), skeletal architecture consisting of a dif-
ferentiated axis, extra-axis, a renieroid skeleton,
and a second category of principal spicules protruding through the ectosome. Echinoclathria leporina
is typical of most Ophlitaspongia (of authors)
(e.g., 0. axinelloides Dendy).
Confusing the generic boundaries even further,
Spongia leporina closely resembles Antho
(Isopenectya) in growth form and gross skeletal
construction, but differentiated by their skeletal
structures, spicule ornamentation and localisation of particular spicules to certain regions of the
skeleton. Nevertheless, it could be argued that
Isopenectya could be included in Echinoclathria
as equally as in Antho. The former option is
tentatively rejected here based on the unequivocal possession of spined (versus smooth) styles
composing the renieroid skeleton, and possession
of a secondary, longitudinal, subisodictyal secondary skeleton in Isopenectya. Similarities in
renieroid skeletal construction may link both
these genera into a clade based on secondary
reduction or loss. The difficulty in positively assigning Isopenectya is discussed further below.
Wiedenmayer (1977: 144) suggested that
Echinoclathria should be restricted to IndoPacific species, although there were similarities
in skeletal architecture with Pandaros from the
West Indies (which lacks microscleres). He noted
further that Echinoclathria had mostly smooth
choanosomal spicules, except for occasional vestigial spines on the bases of some auxiliary
spicules, whereas Pandaros had acanthose
echinating spicules. Simpson (1968a) has already
demonstrated that the loss of spination on
echinating spicules is common amongst
microcionids and not correlated with any
cytological differences (i.e., of low taxonomic
value). Wiedenmayer (1977, 1989) concluded
that although two genera intergraded they could
not be consistently differentiated, and Pandaros
is not included in this group.
Thus, on the basis of evidence presented by
Hallmann (1912: 275), Burton (1959a: 246) and
Wiedenmayer (1989: 58), and re-examination of
all nominal species belonging to these groups, it
is clear that we are dealing with two distinct,
homogenous groups. One, agreeing with the
definition of Spongia leporina above, includes
the genera Echinoclathria (in the strict sense
only, and not of authors), and Ophlitaspongia (of
authors, and not in the strict sense).
Echinoclathria is the earliest available name for
this group, and its characteristics should be taken
as those traditionally associated with the concept
of Ophlitaspongia (of authors). This genus is
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
43
most closely related to A ntho in its renieroid main
skeletal structure, differing in having smooth
choanosomal spicules and a single skeletal stucture (i.e., Antho has 2 skeletal structures: a
renieroid skeleton composed of spined spicules
and a plumose or subisodictyal skeleton composed of smooth spicules).
The second group contains honeycombed
reticulate species traditionally associated with
Echinoclathria (of authors, not in the strict
sense), together with Holopsamma, HaIme, Plectispa and Aulena (of authors, not in the strict
sense). The earliest available name for this group
is Holopsamma Carter (18850. There is no doubt
that this honeycombed reticulate group of
microcionids warrants inclusion in a separate
taxon, although its level of divergence is arguable
(cf. Wiedenmayer, 1977, 1989; Hooper, 1991). Its
peculiar growth form is consistent for all 12
Australian species (all from southern Australia
(Gondwanan) faunas), and 4 non-Australian species
(2 from South America (Gondwanan) and 2 from
the Indo-west Pacific (Tethyan) faunas). This
growth form is correlated with a reticulate skeletal architecture, forming a homogeneous group
which is recognised here at the generic level.
A honeycombed reticulate growth form is also
known for Acamasina de Laubenfels (1936a: 117)
in Mycalidae (Van Soest, 1984b) and Panda ros
(see below), both known only from the West
Indies.
Echinonema Carter, 1875
(Fig. 14F-G)
Echinonema Carter, 1875: 194 [nomen nudum];
Carter, 1881a: 378; Ridley, 1884a: 615; Topsent,
I 894a: 19; Dendy, 1896: 32; Whitelegge, 1901:
80; Topsent, 1928a: 61, 1932: 89, 98; de Laubenfels, 1936a: 112; Levi, 1960a: 56.
TYPE SPECIES. Echinonema typicum Carter, 1881a:
377 (by typonymy) (lectotype BMNH1877.5.21.149);
junior synonym of Spongia cactifonnis Lamarck,
1814: 440 (lectotype MNHNLBIMDT580).
Arborescent, shrubby, lamellate growth form.
Surface even, hispid, subectosomal striations.
Choanosomal skeleton reticulate, with open rectangular or elongate open meshes although slightly compressed in axis. Primary spongin fibres
ascending, heavy, cored by multispicular tracts of
choanosomal principal styles, interconnected by
pauci- or aspicular secondary spongin fibres.
Fibres echinated by acanthostyles heaviest in
peripheral skeleton. Subectosomal skeleton
plumose, with brushes of larger subectosomal
auxiliary subtylostyles erect on peripheral
choanosomal fibres. Ectosomal skeleton dense,
with smaller ectosomal auxiliary subtylostyles
forming a dense palisade on surface. Megascleres
include smooth choanosomal principal styles,
larger subectosomal auxiliary subtylostyles with
smooth or microspined bases, smaller ectosomal
auxiliary subtylostyles with smooth or
microspined bases, and short thick echinating
acanthostyles with spined base and point but bare
neck. Microscleres palmate isochelae of two
sizes, including contort forms, and thin accolada
and asymmetrical toxas.
REMARKS. It is confirmed here that Carter's
(1881a) Echinonema typicum is identical to
Lamarck's (1814) Spongia cactifonnis, and consequently the name cactifonnis has seniority over
the better known junior synonym Clathria typica,
widely used in the literature. Lendenfeld (1888),
Whitelegge (1901) and Hallmann (1912) created
many new subspecific names (as varieties) for
this species, and Hooper & Wiedenmayer (1994)
assigned Lamarck's (1814) specimen to Clathria
(Thalysias) cactifonnis cactiformis, and Carter's
(1881) specimen to C. (T) cactiformis typica.
Examination of type material of all these subspecies (var. typica (SMF1589); var. porrecta
(SMF1653); var. brevispinus (AMZ931); var.
favosus (AMZ944); var. geminus (AMZ928); var.
obesus (AMZ937); var. proximus (AMZ930);
var. stelligera (AME648); and three unnamed
varieties of Hallmann (1912) (AMZ1158;
AMZ1430, G9135, Z938; and AMZ41), showed
that they were conspecific with the nominotypical variety (based on spicule size, spicule
geometry and skeletal structure), whereas growth
forms and surface features varied substantially
between each taxon. The recognition of these
subspecies, a preoccupation of many earlier
authors, does at least demonstrate a high degree
of external morphological variability for the
species, but is not of particular nomenclatural
importance because there are no other morphological characters that correlate with these
differences in external morphology. It may be
eventually determined from biochemical or
genetic data, that this polymorphism is indicative
of sibling species relationships, but no studies of
this sort have yet been undertaken.
De Laubenfels (1936a: 112) suggested that
Echinonema was identical to Thalysias (sensu de
Laubenfels), whereas Van Soest (1984b) and
others placed both genera into synonymy with
Rhaphidophlus (see below). The genus is in-
^
MEMOIRS OF THE QUEENSLAND MUSEUM
44zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 14. Type species of microcionid genera. A-B, Echinochalina (Ophlitaspongia australiensis Ridley,
BMNHI881.10.21.299). C-E, Echinoclathria (E. tenuis Carter, BMNH1886.12.15.147). F-G, Echinonema (E.
typicum Carter, BMNH1877.5.21.149), H-I, HaIme (Holopsamma laminaefavosa Carter,
BMNHI886.12.15.312).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
45
eluded here in Clathria (Thalysias) on the basis
of its ectosomal specialisation.
Fisherispongia de Laubenfels, 1936
(Fig. 15A-C)
Fisherispongia de Laubenfels, 1936b: 460.
TYPE SPECIES. Fisherispongiaferrea de Laubenfels,
1936b: 460 (by original designation) (holotype
USNM22239).
Encrusting growth form. Surface tuberculate,
arenaceous, hispid. Choanosomal skeleton
hymedesmoid, with basal layer of spongin fibre
incorporating detritus and bases of larger
choanosomal principal subtylostyles and smaller
echinating styles, standing perpendicular to substrate, in groups or individually, ascending to but
not protruding through ectosome. Ectosomal
skeleton plumose, with subectosomal auxiliary
polytylostyles, of a single category, arising from
ends of choanosomal megascleres in multispicular bundles protruding through surface.
Megascleres large choanosomal principal subtylostyles with smooth bases, smaller smooth
echinating styles with smooth or microspined
bases, and polytylote auxiliary tylostyles with
smooth or microspined bases. Microscleres palmate isochelae, including contorted forms, and
thick wing-shaped toxas.
REMARKS. De Laubenfels (1936b) distinguished Fisherispongia from other microcionids
by the polytylote bases on their subectosomal
auxiliary styles. In all other respects, however, the
type species resembles other encrusting species
with hymedesmoid architecture (e.g., Leptoclathria). Polytylote ectosomal megascleres
are known in several other species of
Microcionidae (e.g., Clathria aceratoobtusa,
Paratenaciella microxea), as well as in other
Poecilosclerida (e.g., Camptisocale Topsent and
Phellodenna Ridley & Dendy; Coelosphaeridae). The incorporation of detritus into the
choanosome and fibres is well known for several
microcionids and other sponges (see remarks for
Aulenella) and on this basis the species is included in Clathria (Wilsonella). Re-examination
of the holotype found a marked contrast in size
between the smaller and larger (so-called principal) spicules, with no intermediate sizes, and
these smaller spicules are interpreted here as
being smooth echinating styles. By its toxa morphology Fisherispongia ferrea (from the Atlantic
coast of Panama) is very similar to Clathria
aceratoobtusa (from the Indo-west Pacific).
Folitispa de Laubenfels, 1936
(Fig. 15D-F)
Folitispa de Laubenfels, 1936a: 119.
TYPE SPECIES. Hymedesmia laevissima Dendy,
1922:81 (by original designation) (holotype
BMNH1921.11.7.69).
Thickly encrusting growth form. Surface even,
slightly hispid. Choanosomal skeleton hymedesmoid, with spongin fibres lying on substrate and
bases of choanosomal principal subtylostyles embedded, standing perpendicular to substrate individually or forming short multispicular
plumose columns protruding through surface.
Echinating megascleres absent. Subectosomal
skeleton irregularly plumose, with loosely aggregated bundles of subectosomal auxiliary subtylostyles erect on surface or lying tangential to
it. Megascleres include smooth choanosomal
principal subtylostyles, and smooth subectosomal auxiliary subtylostyles with mucronate
or telescoped points. Microscleres palmate
isochelae with anchorate-like modifications (curvature, partially detached alae, continuous ridge
on shaft). Toxas absent.
REMARKS. The type species of Foliti.spa differs
from other encrusting microcionids with
hymedesmoid skeletal architecture (e.g., Leptoclathria) in lacking echinating acanthostyles
(cf. Axocielita) and having chelae with
anchorate-like modifications instead of typical
palmate isochelae (cf. Cionanchora). These
chelae are strongly curved, with lateral alae partially detached from the shaft and a continuous
lateral ridge running the length of the shaft. However, these lateral alae are not fully formed (being
about two-thirds the size of the front ala), nor are
they completely detached from the shaft (attached for approximately 50% of their length),
and consequently they cannot be considered true
anchorate chelae but perhaps palmate isochelae
with substantial anchorate modifications. The
genus is included here in Clathria (Microciona)
based on its skeletal structure.
[Halme] Lendenfeld, 1885
(Fig. 14H-I)
Ha/me Lendenfeld, 1885c: 285, 1889a: 446; de
Laubenfels, 1936a: 17; Bergquist, 1980b: 454;
Wiedenmayer, 1989: 58 (preoccupied).
Not Ha/me Pascoe, 1869.
TYPE SPECIES. Holopsamma laminaefavosa Carter,
1885b: 212 (by subsequent designation; de Laubenfels,
1936a: 17) (holotype BMNH1886.12.15.312).
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46zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 15. Type species of microcionid genera. A-C, Fisherispongia (F. ferrea de Laubenfels, USNM22239). D-F,
Folitispa (Hymedesmia laevissima Dendy, BMNH1921.11.7.69). G-I, Holoplocamia (H. penneyi de Laubenfels, USNM22460).
REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
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Massive, globular, lobate-digitate
honeycombed reticulate growth form. Surface
composed of small branches (lacunae') interconnected to form regular network. Choanosomal
skeleton irregularly reticulate, with heavy spongin fibres fully cored by both sand particles and
other detritus, and with fewer choanosomal principal subtylostyles both coring and echinating
fibres. Subectosomal skeleton with peripheral
fibres cored and echinated by principal
megascleres, slightly heavier, more plumose at
periphery than at core, and with subectosomal
auxiliary strongyles forming irregular paratangential tracts near surface. Ectosome with external fibre reticulation reinforced by sand.
Megascleres vary from common to relatively
scarse (or spicules reportedly absent entirely in
some specimens), including short entirely smooth
choanosomal principal subtylostyles, and
smooth sinuous or straight subectosomal
auxiliary strongyles or quasidiactinal styles.
Microscleres absent.
REMARKS. Ha/me (sensu Lendenfeld, 1889b)
is virtually identical to Aulena (of authors, e.g.,
Lendenfeld, 1888, but not Lendenfeld, 1885c), in
skeletal construction, growth form and the
presence of detritus within the fibre skeleton. In
contrast, Hahne (of Lendenfeld, 1885c) differs
from Aulena (of authors) by the virtual absence
(or inconsistent presence) of proper, heavily
mineralised spicules. However, examination of
relevant type material, recent material from
southern and eastern Australian waters (see
below) and the literature (e.g., Wiedenmayer,
1989) shows that H. laminaefavosa is relatively
polymorphic in its growth form, surface structure, spongin fibre construction, amount of
detritus incorporated into the skeleton and the
number and presence or absence of megascleres.
Despite this variability, the species is clearly a
synonym of Holopsamma, closely related to other
honeycombed reticulate species. Of the
numerous species referred to Hahne by Lendenfeld (1885c, 1888) many are Dictyoceratids
(Bergquist, 1980b), whereas the type species is
undoubtedly a microcionid. Unfortunately the
name Halme Carter, 1885b is preoccupied by
Hahne Pascoe, 1869 (Wiedenmayer, 1989), and
Holopsamma Carter is the senior-most available
name for this group of honeycomb reticulate
microcionid sponges.
Heteroclathria Topsent, 1904
(Fig. 16A-B)
Heteroclathria Topsent, 1904b: 93; Burton, 1935a:
403.
TYPE SPECIES. Heteroclathria hallez; Topsent,
1904b: 94 (by original designation and monotypy)
(schizotype MNHNLBIMDT1884).
Erect, digitate growth form. Surface even,
microscopically hispid. Choanosomal skeleton
regularly renieroid reticulate, with heavy spongin
fibres well differentiated into primary ascending
multispicular fibres, cored by choanosomal principal tylostyles, and secondary transverse uni- or
bispicular fibres cored by amphistrongyles. Subectosomal skeleton plumose or paratangential,
with subectosomal auxiliary subtylostyles forming poorly developed brushes arising from
peripheral primary fibres, and also forming
clusters around margins of oscules. Echinating
acanthostyles absent, but choanosomal principal
styles sometimes echinate primary fibres.
Megascleres choanosomal principal tylostyles
with microspined bases, large strongyles (dumbell spicules) of renieroid skeleton with spined
bases, and subectosomal auxiliary subtylostyles
with microspined bases. Microscleres palmate
isochelae and thin wing-shaped toxas.
REMARKS. Heteroclathria is unusual to the
plocamiform group of sponges (de Laubenfels,
1936a), such as Antho and Plocamilla, in having
a differentiated primary and secondary fibre network cored by monactinal and diactinal
megascleres, respectively. The type species is
only known from the holotype which unfortunately lacks collection data. Burton (1935a:
403) referred two other species to the genus,
Plocamia karykinos de Laubenfels (1927: 262)
and P. manaarensis (sensu Lambe, 1895: 124;
holotype USNM6331; not Carter, 1880a: 34),
which he renamed H. lambei Burton, and which
Bakus (1966: 512) also renamed Plocamilla zhnmeri. De Laubenfels (1936a: 78) suggested that
Heteroclathria was a synonym of Plocamia, and
this is confirmed here from re-examination of
both H. lambei and H. hallezi. Heteroclathria is
referred here to Antho (Plocamia) in having
(acantho)-strongyles in the renieroid skeleton,
although true echinating spicules are absent.
Another species which shows some similarities
to Heteroclathria in skeletal structure is Stylotella
corn uta Topent (1897b: 464), from the Andaman
Sea off Malaysia, for which Burton & Rao (1932:
343) created Acanthostylotella. That species
lacks the characteristic `dumbell spicules' but has
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a renieroid skeleton of primary ascending multispicular fibres interconnected by unispicular
tracts of smooth styles. It lacks an ectosomal
skeleton and lacks microscleres, and it possibly
best placed in Iophonidae.
to Plocamione Topsent (Raspailiidae; Hooper,
1991), whereas the poecilosclerids (including
Holoplocamia) are considered here to belong to
Antho (Plocamia) (the latter a senior name for
Plocamilla).
Holoplocamia de Laubenfels, 1936
(Fig. 15G-I)
Holopsamma Carter, 1885
(Fig. 16C-D)
Holoplocamia de Laubenfels, 1936a: 75; Levi, 1960a:
80; Little, 1963: 47.
TYPE SPECIES. Holoplocamia penneyi de Laubenfels, 1936a: 75 (by original designation) (holotype
USNM22460).
Thinly encrusting growth form. Surface rugose,
microscopically hispid. Choanosomal skeleton
hymedesmoid, with basal layer of spongin fibre,
principal choanosomal styles embedded in fibre
nodes, standing perpendicular to and protruding
through surface, and with basal mass of acanthostrongyles forming an irregular renieroid
secondary reticulation of spicules around principal spicules, interconnected by sparse collagen
at nodes. Smaller acanthostyles also present
echinating fibre nodes. Mesohyl incorporates
large quantities of detritus and auxiliary spicules.
Ectosomal skeleton with tangential or paratangential tracts of subectosomal auxiliary subtylostyles. Megascleres principal choanosomal
styles-subtylostyles with either smooth or
microspined bases, acanthostrongyles or acanthostyles of basal skeleton more heavily spined at
ends than middle, echinating acanthostyles evenly spined, and subectosomal auxiliary subtylostyles with microspined bases. Microscleres
palmate isochelae, including contorted forms,
and wing-shaped toxas.
REMARKS. Holoplocamia was erected for
sponges similar to Plocamia Schmidt, but having
spiny rather than smooth principal spicules. Levi
(1960a) suggested that the genus was a synonym
of Plocamilla, whereas Topsent (1928a) and Little (1963) argued that Plocatnilla was different
from both Plocamia and Holoplocamia in lacking
any differentiation between primary and secondary skeletal tracts. This opinion is not upheld
here. De Laubenfels (1936a: 75) referred several
`plocamiform' species to Holoplocamia, including the type species of Plocamilla, and it is now
generally accepted that Holoplocamia and
Plocamilla are synonymous (Bakus, 1966;
Simpson, 1968a; Levi & Levi, 1983a; PulitzerFinali, 1983; Van Soest, 1984b). Most of the
`plocamiform' species discussed by de Laubenfels (1936a) were subsequently found to belong
Holopsamma Carter, 1885c: 211.
TYPE SPECIES. Holopsamma crassa Carter, 1885c:
211 (by subsequent designation, de Laubenfels, 1936a:
98) (lectotype BMNH 1886.12.15.313; Hooper &
Wiedenmayer, 1994), a senior synonym of Ha/me
globosa Lendenfeld, 1885c: 303 (lectotype
BMNH1886.8.27.71) (cf. Wiedenmayer, 1989: 63).
Subspherical, digitate, regularly 'honeycomb'
reticulate growth form. Surface arenaceous,
porous, with tympanic membrane-like ectosomal
crust stretched across adjacent subdermal
cavities. Choanosomal skeleton reticulate, with
well developed spongin fibres not well differentiated into primary or secondary elements, although many ascending fibres have core of small
quantities of detritus (mostly spicule fragments),
whereas other fibres clear of detritus completely.
Coring and echinating spicules absent from
choanosomal skeleton. Ectosomal skeleton
heavily arenaceous, with crust of sand and scattered reticulate (or plumose or paratangential in
places) bundles of subectosomal auxiliary strongyles lying tangential on surface crust. Mesohyl
matrix relatively heavy between fibres .
Megascleres only smooth subectosomal auxiliary
strongyles. Microscleres absent.
REMARKS. De Laubenfels (1936a: 97) noted
that Holopsamma differs from other 'sandy
sponges' (i.e., the polyphyletic 'family
Psammascidae' de Laubenfels) in lacking
microscleres and having both monactinal and
diactinal megascleres. It is unfortunate that he
designated H. crassa as the type species because
in some of the 5 'valid' syntypes the monactinal
(principal) styles may be lost completely, and the
diactinal (auxiliary) strongyles are vestigial,
leaving only heavy spongin fibres (the major ones
with a core of detritus), and a heavy ectosomal
sand cortex.
The status and affinities of this genus are still
confused, despite the comprehensive redescription and discussion of the type species by
Wiedenmayer (1989). This confusion is due to the
fact that no-one had previously nominated a lectotype amongst the 31 syntypes of the type
species, which are composite and represent at
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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least 6 different species: only 5 of these actually
conformed to Carter's (1885c) original description of Holopsamma crassa.
Furthermore, the lectotype (BMNH1886.12.15.313) designated by Hooper &
Wiedenmayer (1994) and figured by Wiedenmayer (1989: p1.6, fig.7) is identical to Ha/me
globosa Lendenfeld, and there is some conjecture
as to which name is the most valid. Wiedenmayer
(1989: 63) chose to use the name H. globosa over
H. crassa, although he admitted that crassa was
more senior (apparently by only several months),
and therefore under the rules of the ICZN it must
take precedence. His arguments in choosing
globosa over crassa were that type material of
globosa was firmly established whereas the type
series of crassa was an unresolvable mess, but
this is inrelevant with the subsequent designation
of a lectotype for H. crassa by Hooper & Wiedenmayer (1994).
Wiedenmayer (1989: 63) provided many further details concerning these species synonymies
and the affinities of 'sandy sponges' belonging to
the Microcionidae. But more important than the
nomenclatural problems associated with the type
species (and the genera Ha/me and Holopsamma), there are some biological questions unanswered by Wiedenmayer's (1989) work.
Holopsamma crassa is very similar to Holopsamma laminaefavosa, the type species of Ha/me
Lendenfeld, 1885 (not Ha/me Pascoe, 1869), and
it is possible that in fact the two species are
synonymous (given that they are both allegedly
very polymorphic). A comparison of type
material shows that the only substantial differences are that in H. crassa spongin fibres contain
virtually no sand and principal spicules have been
lost, whereas in H. laminaefavosa primary fibres
are virtually fully cored with sand, and both principal and auxiliary spicules are retained. In dealing with preserved material (i.e., without
accompanying field characters), these definitions
should be adhered to strictly. Conversely, following Wiedenmayer's (1989) definition, diagnostic
characters in each species overlap substantially
providing reasonable cause to synonymise the
species (and genera Holopsamma and Ha/me).
In the present work Holopsamma is the senior
name for the group of honeycomb reticulate
sponges traditionally known as Echinoclathria
(of authors).
Hymantho Burton, 1930
(Fig. 16F-G)
Hymantho Burton, I930a: 503.
TYPE SPECIES. Hymantho normani Burton, 1930a:
503 (by original designation) (holotypeBMNH
1910.1.1.791).
Thinly encrusting growth form. Surface even,
hispid. Choanosomal skeleton hymedesmoid,
with basal layer of spongin on substrate and bases
of choanosomal principal subtylostyles and
echinating acanthostyles embedded and standing
perpendicular to substrate. Ectosomal skeleton
with paratangential bundles of subectosomal
auxiliary subtylostyles of single size category.
Mesohyl matrix with some debris incorporated.
Megascleres choanosomal principal subtylostyles with acanthose bases, echinating acanthostyles with spined bases and shafts but aspinose
points, and smooth auxiliary subtylostyles or
polytylostyles. Microscleres palmate isochelae
and thick forceps-shaped or v-shaped toxas.
REMARKS. Hymantho normani was originally described with only toxa microscleres but reexamination of the holotype discovered that large
palmate isochelae (18-22m), with large alae, are
also present. Hymantho was erected by Burton
(1930a), being similar to Leptoclathria Topsent
in its hymedesmoid skeletal architecture, but supposedly lacking chelae microscleres (Levi,
1960a: 60). In this latter respect the genus was
also be compared with Pseudanchinoe and QuercicIona, which Van Soest (1984b) considered to
be junior synonyms of Clathria (s.1.), but in any
case the discovery of isochelae in the type species
negates the concept of the genus. Alander (1942)
and Van Soest & Stone (1986) also suggested that
the secondary loss of isochelae and the presence
of a leptoclathriid skeleton have little generic
value in the Microcionidae. The genus is referred
into synonym with Clathria (Microciona).
Isociella Hallmann, 1920
(Fig. 16H-I)
Isociella Hallmann, 1920: 784; de Laubenfels, 1936a:
152; Bergquist & Tizard, 1967: 187; Bergquist &
Fromont, 1988: 114.
TYPE SPECIES. Phakellia flabellata, in part (sensu
Ridley & Dendy, 1886: 478) (by monotypy) (not P.
flabellata Carter, 1885f: 363); =Phakellia jacksoniana
Dendy (replacement name; Dendy, 1897: 236)
(holotype BMNH1887.5.2.9), both junior synonyms of
Clathria macropora, in part, Lendenfeld, 1888: 221
(holotype AMZ466) (this work).
Erect, stipitate, flabelliform growth forms. Surface hispid, relatively smooth, with ridges and
lamellae. Choanosomal skeleton relatively
homogeneous, renieroid, composed of primary,
multispicular, plumose, ascending tracts of
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FIG. 16. Type species of microcionid genera. A-B, Heteroclathria (H. hallezi Topsent, MNHNDT1884). C-D,
Holopsamma (H. crassa Carter,BMNH1886.12.15.313). E, Ha/me (H. globosa Lendenfeld, synonym of H.
crassa, BMNH1886.8.27.71). F-G, Hymantho (H. normani Burton, BMNHI910.1.1.791). H-I, Isociella
(Phakelliaflabellata sensu Ridley & Dendy, BMNH1887.5.2.9).
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
choanosomal principal styles, interconnected by
secondary, uni- or paucispicular, transverse tracts
of same spicules, together forming regular
renieroid or sub-renieroid (triangular) meshes;
principal spicules bonded together at nodes by
collagen or enclosed in relatively poorly
developed spongin fibres. Echinating acanthostyles absent. Subectosomal auxiliary styles of a
single category, arranged tangentially, paratangentially or in plumose brushes on surface, with
some principal styles also protruding through
peripheral skeleton singly or in sparse plumose
brushes. Megascleres robust smooth
choanosomal principal styles, and small smooth
or basally spined subectosomal auxiliary stylessubtylostyles. Microscleres palmate isochelae
with 'fluted' alae, no toxas (in type species).
REMARKS. Isociella contained 3 species prior
to the present study (Clathria macropora Lendenfeld, Ophlitaspongia eccentrica Burton
(1934a: 560), and !. incrustans Bergquist (1961a:
42)), although Hallmann (1920: 784) suggested
that there were some other species included in his
concept of Ophlitaspongia (=Echinoclathria as
defined in this study) which could also be referred
here. The definition above is widened below to
include species with toxas and different forms of
palmate isochelae.
Dendy (1897) renamed the type species jacksoniana because Phakellia flabellata Ridley &
Dendy (1886) was preoccupied by Phakellia
flabellata Carter (1885f) from Port Phillip Bay,
but Hallmann (1920) considered that the 2
species belonged to different genera, and the replacement name flabellata was unnecessary.
However, P.flabellata Ridley & Dendy is a junior
homonym of P. flabellata Carter, and Dendy's
(1897) replacement name P jacksoniana is a
valid emendment. Neither species belongs to
Phakellia. In any case C. macropora Lendenfeld
is identical to, and the senior available name for,
P jacksoniana.
Isociella is distinguished from other
microcionids by its relatively homogeneous,
wide-meshed, sub-renieroid reticulation of a
single category of smooth choanosomal styles,
lacking echinating spicules, and without any differentiation between axial and extra-axial regions
(although choanosomal spicules may diverge
slightly towards periphery, sometimes becoming
plumose on surface). It differs from the two other
groups of microcionids that have renieroid skeletal structure. Antho (including Isociona,
Plocamia, Plocamilla, Plocamiopsis, and
Isopenectya), has two categories of choanosomal
megascleres, one acanthose forming a basal
renieroid skeleton, and one smooth forming ascending plumose extra-fibre tracts of the true
choanosomal skeleton. Some Echinoclathria
species (as defined in the present study, including
Ophlitaspongia in the sense of most authors),
have a renieroid main skeleton of smaller, smooth
choanosomal styles and the same spicules
echinating fibres, a radial peripheral skeleton in
which larger, smooth choanosomal styles are embedded in the subectosomal skeleton and
protrude a long way through the surface, and a
very well developed spongin fibre reticulation
seen in most species, often with some axial compression (reflecting a digitate or flabellate growth
form).
The most abundant Australian species, Isociella eccentrica, has a choanosomal skeleton
reminiscent of Callyspongia (Haplosclerida) and
an open, reticulate ectosomal skeleton, and all 3
known Australian species have closest affinities
with certain Clathria species (e.g., C. (Clathria)
conectens, C. (Thalysias) hirsuta). Thus, the
definition of the type species is expanded below
to include forms which are predominantly semiencrusting, branching, with rugose reticulate surface sculpturing, well developed spongin fibres
(as opposed to only having collagen binding principal spicules together); poorly differentiated
primary and secondary skeletal tracts, and to
allow for the presence of toxa microscleres.
Isociella is included here as a subgenus of
Clathria.
Isociona Hallmann, 1920
(Fig. 17A-B)
Isocionu Hallmann, 1920: 768; de Laubenfels, 1936a:
TYPE SPECIES. Lissodendoryx tuberosa Hentschel,
1911: 326 (by monotypy) (holotype ZMB4417).
Thickly encrusting, bulbous growth form. Sur-
face irregularly microconulose, hispid.
Choanosomal skeleton with differentiated
primary and secondary skeletons. Secondary
skeleton renieroid or subrenieroid reticulate, with
acanthose styles forming uni-, pauci-, or less frequently multispicular tracts, bound together by
very light fibres or collagen at spicule nodes.
Primary skeleton plumose, with choanosomal
principal styles forming larger primary ascending
tracts irregularly connected by smaller secondary
transverse tracts. Subectosomal skeleton
plumose, with principal styles protruding through
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ectosome overlayed by erect bundles of subectosomal auxiliary subtylostyles. Echinating
megascleres absent. Megascleres large smooth
choanosomal principal styles, basally spined or
entirely lightly spined styles-subtylostyles of
renieroid skeleton, and smooth auxiliary subectosomal subtylostyles. Microscleres palmate
isochelae and thick wing-shaped toxas.
REMARKS. Isociona tuberosa and Isociella eccentrica appear very similar from published
descriptions whereas examination of type
material shows that they are different. Isociella
eccentrica has a wide meshed renieroid reticulate
skeleton composed of smooth principal spicules,
whereas I. tube rosa has a close-meshed renieroid
secondary skeleton of spined monactinal
spicules, overlayed by a plumose primary skeleton
of smooth principal spicules. Van Soest (1984b)
merged lsociona with Antho, although this relationship is not straightforward given that Antho is
usually reserved for forms with diactinal
megascleres (in the renieroid basal skeleton).
Nevertheless, the two taxa have similar skeletal
architecture and Isociona is maintained as a
synonym of Antho herein.
Isopenectya Hallmann, 1920
(Fig. 17C-D)
Isopenectya Hallmann, 1920: 789; de Laubenfels,
1936a: 125.
TYPE SPECIES. Clathria (?) chartacea Whitelegge,
1907: 497 (by monotypy) (holotype AMZ436).
Thinly flabellate-lamellate growth form. Surface smooth, even. Choanosomal skeleton
renieroid reticulate, with differentiated axial and
extra-axial regions of skeleton. Axial skeleton
with compressed spongin fibres running through
centre of lamellae, cored by smooth choandsomal
principal styles (marginally smaller than those in
surface bundles) forming subisodictyal tracts,
overlain by renieroid skeleton of small spined
styles. Extra-axial skeleton with more openmeshed, regularly reticulate spongin fibres, cored
by uni- or bispicular renieroid tracts of small
acanthose styles, and plumose, subisodictyal
tracts of longer smooth styles standing perpendicular to axis, both fully enclosed in spongin
fibres. Echinating megascleres absent. Subectosomal skeleton plumose, with bundles of
smooth choanosomal principal styles protruding
through surface and tangential or paratangential
tracts of subectosomal auxiliary styles. Ectosome
lacks specialised spiculation. Megascleres lightly
acanthose styles-subtylostyles forming renieroid
skeleton, short and long smooth choanosomal
principal styles, and subectosomal auxiliary
styles with spined bases. Microscleres absent.
REMARKS. Isopenectya is similar to other
renieroid `plocamiform' genera (sensu de
Laubenfels, 1936a), Antho, Plocamilla,
Plocamiopsis, Isociona, Labacea, Panda ros,
lsociella and Echinoclathria, in having a primarily renieroid reticulate skeleton. Unlike these
other genera, however, Isopenectya has 2 forms
of choanosomal megascleres enclosed within
spongin fibres. The smaller acanthose styles
forming the renieroid structure, whereas smooth
styles form the subisodictyal, mostly longitudinal
skeleton. These latter spicules also produce the
extra-axial plumose tracts that ascend to the surface, and larger, smooth choanosomal styles
produce brushes on the ectosome. Although the
fibre skeleton is compressed in the axis the
renieroid skeleton is barely different between
axial and extra-axial regions.
Isopenectya chartacea is remarkable in that it
closely resembles the type species of
Echinoclathria (E. leporina) in growth form,
gross skeletal architecture and fibre characteristics. Whereas E. leporina has one category of
relatively homogeneous smooth principal styles
throughout the choanosomal skeleton, I. chartacea has 2 differentiated structures within the
choanosome (renieroid and plumose or subisodictyal skeletons) and 2 geometrically different categories of megascleres forming these
skeletal structures. Skeletal architecture is also
more regularly renieroid and renieroid
fibres/tracts are relatively homogeneous than in
most Echinoclathria species, but this is a matter
of degree. Echinoclathria has a radial skeleton of
larger, smooth principal styles embedded in
peripheral fibres, poking through the surface, and
it could be argued that this is a vestigial subisodictyal skeleton like that in Isopenectya. But this
similarity is inferred and any relationship is
equivocal, with emphasis placed here on the possession of spined spicules of the renieroid
skeleton and clearly differentiated renieroid and
subisodictyal skeletal structures in deciding on
affinities of Isopenectya. Nevertheless, these 2
species are certainly remarkably similar and
potentially may be confused.
Van Soest (1984b) suggested that Isopenectya
may be valid, but `plocamiform' microcionid
type shows that it is a close relative of this group,
of Antho in particular, differing from it only in
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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having a unique subisodictyal skeleton of smooth
spicules overlaying the renieroid basal skeleton.
It is recognised here as a subgenus of Antho.
TYPE SPECIES. Clathria juncea sensu Burton,
1931a: 343 (by original designation) (type fragment
BMNH1926.2.19.2).
Erect digitate, arborescent growth form. Sur-
Jia de Laubenfels, 1930
face even, minutely hispid. Choanosomal
skeleton reticulate, with well differentiated
primary ascending and secondary transverse
skeletal tracts. Primary skeleton with well
developed thick spongin fibres, ascending to surface, cored by multispicular tracts of
choanosomal principal styles, interconnected by
thin transverse secondary fibres cored by paucior unispicular tracts of same spicules. Echinating
acanthostyles moderately common in axial
skeleton, very heavy in peripheral skeleton, forming plumose brushes at surface. Subectosomal
skeleton tangential with subectosomal auxiliary
subtylostyles lying on surface. Ectosome without
specialised spiculation, but with many foreign
spicule fragments embedded in outer layer of
skeleton. Mesohyl with abundant detritus and
auxiliary spicules dispersed between fibres.
Megascleres include both smooth and completely
lightly spined fusiform choanosomal principal
styles, evenly spined echinating acanthostyles,
and smooth auxiliary subectosomal subtylostyles. Microscleres large palmate isochelae and
accolada toxas with microspined points.
(Fig. 17E-F)
Jia de Laubenfels, 1930: 28, 1932: 97.
TYPE SPECIES. Jia jia de Laubenfels, 1930: 28 (by
original designation) (holotype USNM21510).
Encrusting growth form. Surface uneven,
hispid. Choanosomal skeleton confused renieroid
reticulate, composed of smaller smooth or acanthose styles forming basal more-or-less rectangular network, overlayed by larger smooth
principal styles standing erect, forming ascending bundles or single spicules projecting through
surface. Subectosomal auxiliary subtylostyles
paratangential to surface. Ectosome without special spicules. Megascleres larger smooth
choanosomal principal styles, smaller smooth or
acanthose styles-subtylostyles of basal skeleton,
and subectosomal auxiliary subtylostyles with
basal spines. Microscleres palmate isochelae,
modified J-shaped chelae resembling sigmas (=
`crocae'), and wing-shaped toxas with spinous
extremities.
REMARKS. De Laubenfels (1932) description
of the type is incomplete. The essential specific
characteristics include the modified (J-shaped)
isochelae (crocae of Van Soest & Stone, 1986),
and a renieroid albiet ill-defined reticulation.
Those characters are contrasted with the Antholike nature of closely related A. (Jia) brattegardi
Van Soest & Stone, which has acanthose monactinal and diactinal spicules in the main skeleton,
whereas Jia (s.s.) has predominantly smooth
spicules. Despite these differences, Van Soest &
Stone (1986) justifiably merged Jia and Antho
because many other Antho-like species without
crocae also have predominantly smooth styles
and poorly defined skeletal construction (e.g., A.
dichotoma (Esper)). Jia is referred to Antho
(Ant ho).
Crocae are not unique to Jia; other taxa with
similarly modified chelae are Dendoryx luciensis
Topsent (Myxillidae) and Zygherpe hyaloderma
de Laubenfels (Hamacanthidae) (de Laubenfels,
1932; Bakus, 1966; Van Soest & Stone, 1986.
Labacea de Laubenfels, 1936
(Fig. 17G-H)
Labacea de Laubenfels, 1936a: 125.
REMARKS. Clathria juncea is attributed to Burton (1931a), and a lectotype designated from the
syntypes (BMNH1933.7.4.4-7). However, Burton and de Laubenfels (1936a) suggested that
Clathria juncea sensu Burton may be
synonymous with Alcyonium junceutn Lamarck,
1816. Topsent (1933: 26) merged Lamarck's
species as a variety of Anomoclathria opuntioides, but noted that the type was missing from
the MNHN. It is still missing, and must now be
presumed destroyed. If Lamarck's species is
identical with A. opuntioides then C. juncea sensu
Burton is quite different (see remarks for
Anomoclathria).
Burton's (1931a) description, and de
Laubenfels' (1936a) interpretation of the type
species are both erroneous. Burton's described
material exhibits several characters not noted by
either of these authors, so that Labacea is incorrectly diagnosed. The skeletal architecture of C.
juncea is reticulate, not renieroid as supposed by
de Laubenfels, and echinating acanthostyles (a
prominent feature of the peripheral skeleton in
particular) and palmate isochelae were overlooked. I assign it to Clathria (Clathria).
De Laubenfels (1936a) assigned Clathria
oxeifera Ferrer Hernandez to this genus; it sup-
^
MEMOIRS OF THE QUEENSLAND MUSEUM
54zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 17. Type species of microcionid genera. A-B, lsociona (Lissodendoryx tube rosa Hentshel, ZM B4417). C-D,
Isopenectya (Clathria chartacea Whitelegge, AMZ436). E-F, Jia (J. jia de Laubenfels, USNM21510). G-H,
Labacea (Clathria juncea sensu Burton, BMNH1926.2.19.2). I, Leptoclathria (L haplotoxa Topsent,
MNHNDT1101).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
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55
posedly differed from the type species in having
echinating acanthostyles (as distinct from acanthose varieties of choanosomal principal
megascleres (Levi, 1960a: 84)) but as noted
above these spicules are typical for the genus, and
de Laubenfels' observation is redundant.
Leptoclathria Topsent, 1928
(Figs 171, I8A)
Leptoclathria Topsent, 1928a: 298.
TYPE SPECIES. Leptoclathria haplotoxa Topsent,
1928: 298 (by monotypy) (holotype MNHNLBIMDT1101).
Encrusting growth form. Surface irregular,
hispid. Choanosomal skeleton hymedesmoid,
with spongin fibres reduced to basal layer of
spongin lying on substrate, with bases of
choanosomal principal subtylostyles and abundant smaller acanthostyles standing perpendicular to substrate. Subectosomal skeleton with
light tangential tracts of subectosomal auxiliary
subtylostyles occurring in bundles or individually, forming irregular dermal brushes surrounding
protruding choanosomal megascleres. Ectosomal
skeleton without specialised spiculation.
Megascleres include entirely acanthose
choanosomal principal subtylostyles, evenly
spined echinating acanthostyles, and subectosomal auxiliary subtylostyles with spined bases
and telescoped or mucronate points, sometimes
quasidiactinal. Microscleres palmate isochelae
and toxas intermediate between wing-shaped and
v-shaped.
REMARKS. The type species is simply a thinly
encrusting Microciona-like species with a
hymedesmoid skeletal architecture, but which
differs from Microciona (s.s.) in having entirely
acanthose principal styles (similar to Dictyociona,Hymeraphia and Anaata). Levi (1960a)
proposed that Leptoclathria should be merged
with Microciona, and this is certainly true for the
type species (referred here to C. (Microciona)),
but there are some species which have a clearly
differentiated ectosomal and subectosomal
auxiliary spicules (e.g., L. lambda Levi), and
these species are more appropriately included in
C. (Thalysias).
Topsent (1928a) overlooked the spined bases
and telescoped (or mucronate) points on subectosomal auxiliary subtylostyles which are
prominent in the holotype. Furthermore, and of
greater taxonomic significance, there are at least
a small proportion of auxiliary megascleres with
tylote (quasidiactinal) geometry, including basal
microspination, which is reminiscent of the
specialised diactinal ectosomal megascleres
characteristic of Myxillidae and Iophonidae.
Only a few other microcionids have this feature,
C. (Clathria) chelifera, C. (Wilsonella) australiensis and C. (Thalysias) major (see also Hooper
et al., 1990). These quasi-diactinal spicules are
analogous to (but not homologous with) ectosomal tylotes found in Myxillidae for example
(see remarks for Acamus below).
Ligrota de Laubenfels, 1936
(Fig. 18B-C)
Ligrota de Laubenfels, 1936a: 125.
TYPE SPECIES. Clathria lobata Vosmaer, 1880: 151
(by original designation) (holotype RMNH276).
Arborescent growth form with flattened
branches. Surface even, microscopically hispid.
Choanosomal skeleton regularly reticulate, with
poorly developed spongin fibres forming differentiated primary and secondary tracts. Primary
fibres ascend to surface, cored by plumose multispicular tracts of choanosomal principal styles;
secondary fibres transverse, paucispicular, cored
by same spicules. Subectosomal skeleton
plumose, with bundles of choanosomal principal
megascleres extending into ectosome and
protruding through surface. Echinating acanthostyles most abundant in periphery, below surface
spicule brushes. Some detritus incorporated into
mesohyl, scattered between spicule tracts. Ectosome radially arranged, erect brushes of subectosomal auxiliary subtylostyles of a single size
class. Megascleres robust choanosomal principal
styles-subtylostyles with smooth bases, echinating acanthostyles with large spines and aspinose
points, and subectosomal auxiliary subtylostyles
with basal spines. Microscleres palmate
isochelae and wing-shaped toxas with spined
points.
REMARKS. This definition from the holotype
differs from descriptions provided by Vosmaer
(1880), Ridley & Dendy (1887), and Levi (1963),
but corresponds with Stephens's (1915) concept
of the species. In particular, the species has a
regularly reticulate skeleton with radial architecture, but it lacks a well differentiated axial and
extra-axial region as suggested by these authors.
Ligrota has spiculation virtually identical to that
of Clathria (s.s.), including spines on the points
of toxas. It is not a Thalysias, as supposed by
Ridley & Dendy (1887), because it has only a
MEMOIRS OF THE QUEENSLAND MUSEUM
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
single, undifferentiated category of auxiliary subtylostyle and is referred here to Clathria
(Clathria). Similarly, de Laubenfels' (1936a)
diagnosis of Ligrota is incorrect. Clathria lobata
of Stephens (1915) does not have diactinal
megascleres or sigmas. Those attributes were
described by Vosmaer (1880) for the type
specimen, but were discounted by Ridley &
Dendy (1887) as being probable contaminants;
nor were they observed in type material.
Lissoplocamia Brondsted, 1924
(Fig. 18D-E)
Lissoplocamia Brondsted, 1924: 470.
TYPE SPECIES. Lissoplocamia prima Brondsted,
1924: 470 (by original designation) (holotype in ZMC,
not seen; MNHNLBIMDCL637 from South Africa).
Digitate, arborescent, flattened or cylindrical
branches. Surface prominently hispid, velvetty.
Choanosomal skeleton renieroid reticulate, with
differentiated axial and extra-axial regions. Axial
skeleton slightly compressed, with well
developed spongin fibres cored by uni- or
paucispicular tracts of tylotes forming renieroid
reticulation, overlaid by plumose (or
plumoreticulate) extra-axial skeleton composed
of diverging single or multiple choanosomal principal styles, echinating (protruding from) spongin
fibres and ascending to surface. Spongin fibres in
extra-axial skeleton lighter, more-widely spaced.
Subectosomal skeleton plumose, with bundles of
subectosomal auxiliary styles perched on ends of
principal megascleres. Ectosome without special
category of megascleres, but isochelae microscleres predominant in peripheral skeleton.
Megascleres tylotes ('dumbell-shaped' spicules)
of renieroid skeleton with swollen microspined
bases, choanosomal principal styles-subtylostyles, with smooth or faintly microspined bases,
and smooth subectosomal auxiliary styles. Microscleres palmate isochelae and wing-shaped toxas.
REMARKS. This definition is based on a recent
redescription of the holotype from New Zealand
(Bergquist & Fromont, 1988: 122) and Levi's
(1963) specimen from South Africa. Levi's
material is more complete, with an intact ectosomal skeleton and containing palmate
isochelae, whereas Brondsted's (1924) holotype
is poorly preserved, lacking an ectosomal
skeleton in which the isochelae are predominantly found (Bergquist & Fromont, 1988).
Lissoplocamia belongs to de Laubenfels'
(1936a) `plocamiform' group of sponges in
having diactinal, quasi-diactinal or monactinal
`dumbell-shaped' spicules forming a renieroid
basal skeleton (it should be noted that this character is different from (analogous to) similar
'sausage-shaped' diactinal spicules found in the
ectosomal skeletons of some genera in families
such as Coelosphaeridae and Petrosiidae). Some
of these `plocamiform' genera belong to
Raspailiidae (Hooper (1991: 1319)), whereas
others are closely related to Microcionidae and
are referred to Antho (Plocamia). Bergquist &
Fromont (1988: 122) mentioned several of these
genera (Lissoplocamia, Holoplocamia, Plocamilla, Heteroclathria, and Plocamia), and to
this group should also be added Antho and Jia.
Axoplocamia, included by them with the
microcionids, was shown to be a raspailiid
(Hooper, 1991). Bergquist & Fromont (1988)
included only 2 New Zealand species in
Plocamia, Dirrhopalum novizelanicum Ridley
and L. prima, both of which are referred here to
Antho. Lissoplocamia differs from most other
`plocamiform' microcionids, such as Antho
(Antho) in having predominantly smooth 'clumbell-shaped' spicules, similar to the type
species of Plocamia, P gymnazusa, but this character is considered insignificant at the generic
level given its large variability within the family.
Litaspongia de Laubenfels, 1954
(Fig. 18F-G)
Litaspongia de Laubenfels, 1954: 162.
TYPE SPECIES. Ophlitaspongia arbuscula Row,
191 1: 347 (by original designation) (holotype
BMNH1912.2.1.63).
Convoluted arborescent growth form. Surface
irregularly conulose, arenaceous, minutely
hispid. Choanosomal skeleton regularly reticulate, with heavy spongin fibres cored by uni- or
paucispicular tracts of choanosomal principal
subtylostyles (sometimes fibres aspiculose).
Echinating acanthostyles sparse. Subectosomal
skeleton plumose, with dense bundles of principal styles (identical to coring spicules) diverging from ends of peripheral fibres and forming
discrete brushes on surface. Ectosomal skeleton
with sparse subectosomal auxiliary subtylostyles
tangential to surface and also dispersed
throughout mesohyl. Megascleres thin smooth
choanosomal principal subtylostyles, smooth
quasi-diactinal subectosomal auxiliary subtylostyles (usually resembling asymmetrical strongyles),
and evenly spilled echinating acanthostyles. MICIDscleres wing-shaped toxas. Isochelae absent.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
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57
REMARKS. De Laubenfels (1954) stated that
Litaspongia was established for sponges like
Echinoclathria (as defined here) in having
monactinal megascleres, toxa microscleres and
arborescent growth form. The holotype also has
echinating acanthostyles, previously overlooked
by Row (1911). The resemblance between 0.
arbuscula and Echinoclathria is here considered
superficial, based on the fact that principal
spicules in 0. arbuscula are thin, attaining only
the thickness typical of auxiliary spicules found
in most other Clathria species; auxiliary spicules
are quasidiactinal in 0. arbuscula, resembling
diactinal ectosomal spicules in some species of
Echinoclathria (e.g., E. chalinoides) and Holopsamma (e.g., H. ramosa); and spongin fibres are
regularly reticulate, well developed, and tend to
dominate skeletal structure over spicule components. But unlike Holopsamma and
Echinoclathria which have undifferentiated
coring and echinating spicules, Litaspongia has
different principal spicules coring fibres from
those echinating fibres. I consider it a reduced
Clathria (Clathria).
De Laubenfels (1954: 162) synonymised 0.
arbuscula and 0. horrida Row (1911: 349), and
re-examination of both holotypes (the latter
BMNH1912.2.1.65) supports this decision. He
added Echinoclathria nodosa (which he merged
with E. subhispida) to Litaspongia, but both are
species of Echinoclathria. Pulitzer-Finali (1982:
105) referred 0. arbuscula and 0. horrida to
Kerasemna (Desmacellidae) (Hooper, 1984b),
but neither have sigma microscleres and this
placement was unjustified.
Marleyia Burton, 1931
(Fig. 18H-I)
Marleyia Burton, 1931a: 346; de Laubenfels, 1936a:
109.
TYPE SPECIES. Marleyia irregularis Burton, 1931a:
346 (by original designation) (holotype NM 1279).
Digitate growth form, flattened branches. Sur-
face uneven, porous, with specialised reticulate
external fibrous skeleton. Choanosomal skeleton
reticulate, with subisodictyal reticulation of well
developed primary and secondary spongin fibres.
Primary fibres ascending, cored by multispicular
tracts of both choanosomal principal subtylostyles and subectosomal auxiliary subtylostyles.
Fibres heavily echinated by short acanthostyles.
Secondary fibres transverse, aspicular, but of
similar diameter and density of echinating
spicules as primary fibres. Abundant detritus
scattered through mesohyl, but not incorporated
into fibres. Ectosome with tangential reticulation
of spongin fibres, more closely reticulate and
slightly thinner than choanosomal fibres, lightly
cored by both subectosomal and choanosomal
subtylostyles and echinated by abundant acanthostyles. Megascleres smooth robust
choanosomal principal subtylostyles, smooth
straight or sinuous subectosomal auxiliary subtylostyles, and robust echinating acanthostyles
with bare neck. Microscleres absent.
REMARKS. The holotype is dry and lacks a well
preserved ectosomal skeleton). Burton suggested
that Marleyia differs from all other microcionid
genera in having a special ectosomal skeleton
formed by a tangential reticulation of spongin
fibres, which are thinner and more closely compacted than choanosomal fibres. This feature was
not as remarkable as we are led to believe from
the original description (although the holotype is
not well preserved), nor is it unique in the
Microcionidae (also seen in Echinochalina
(Protophlitaspongia) labouti Hooper & Levi) or
other families (e.g., Callyspongiidae
(Haplosclerida)).
In gross morphology and fibre characteristics
Marleyia is similar to several Holopsamma
species, but it has different spicules coring and
echinating spicules (whereas Holopsamma has
only one category of principal spicule performing
these functions). Burton (1931a) made further
casual comparisons between Marleyia and certain Dictyoceratida, based on fibre characteristics
and a greatly reduced skeleton. De Laubenfels
(1936a: 109) suggested that Marleyia may have
affinities with Acantheurypon, representing a
more mature form of that genus, but he noted that
Marleyia had a remarkable external resemblance
to `keratose' sponges. On the basis of its unusual
ectosomal fibre characteristics Van Soest (1984b)
suggested that Marleyia might be a valid genus
of Microcionidae, but re-examination of type
material indicates that it clearly belongs to
Clathria (Clathria). Marleyia is monotypic, and
known only from the Durban region, Natal coast,
South Africa.
Microciona Bowerbank, 1862
(Fig. 19F-G)
Microciona Bowerbank,1862b: 1109; Topsent, 1894a:
18; Dendy, 1922:60; Topsent, 1928a: 62; Vosmaer,
1935a: 604 (in part); de Laubenfels, 1936a: 447;
Levi, 1956b: 399; Koltun, 1959: 181; Burton,
1959a: 225; Levi, 1960a: 51; Sara, 1963: 210, 1964:
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FIG. 18. Type species of microcionid genera. A, Leptoclathria (L haplotoxa Topsent, MNHNDT1101). B-C,
Ligrota (Clathria lobata Vosmaer, RMNH276). D-E, Lissoplocamia (L. prima Brondsted, MNHNDCL637).
F-G, Litaspongia (Ophlitaspongia arbuscula Row, BMNH1912.2.1.63). H-I, Marleyia (M. irregularis Burton,
NM1279).
59
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
230; Bergquist, 1965: 168; Hechtel, 1965: 41;
Simpson, 1968a: 93, 102; Levi, 1973: 613; Wiedenmayer, 1977: 140; Bergquist, 1978: 172; Bergquist
& Fromont, 1988: 100.
TYPE SPECIES. Microciona atrasanguinea Bowerbank, 1862b: 1109 (by subsequent designation
(Bowerbank, 1864: 188)) (holotype BMNH
1930.7.3.225).
Encrusting growth form. Surface hispid, uneven. Choanosomal skeleton hymedesmoid, with
spongin fibres reduced to basal layer lying on
substrate, bearing erect, non-anastomosing, rarely branching, scattered fibre nodes perpendicular
to substrate ('microcionid' fibres), each cored by
plumose ascending columns of choanosomal
principal subtylostyles, wholly or partly embedded in fibres, with points of spicules usually
projecting through ectosome. Echinating acanthostyles also erect on fibre nodes. Subectomal
skeleton with tangential layer of subectosomal
auxiliary subtylostyles, singly or in bundles on
surface. Ectosomal skeleton without specialised
spiculation, but choanosomal and subectosomal
spicules protude through surface. Megascleres
choanosomal principal subtylostyles with smooth
or microspined bases, evenly spined echinating
acanthostyles, and smooth or basally spined subectosomal auxiliary subtylostyles. Microscleres
palmate isochelae and wing-shaped toxas.
REMARKS. In many publications, Clathria
Schmidt and Microciona Bowerbank have the
date 1862. In merging the two genera, Wiedenmayer (1977, 1989) and Van Soest (1984b) note
that the former has priority, because Bowerbank's
work was not officially published until 1863.
This ddefinition of Microciona is compiled
from description of the type species (Bowerbank,
1862b, 1864, 1866) and histological preparations
of the holotype and paratypes (BMNH1910.1.1.68, 1930.7.3.226). Microciona (s.s.) differs
from Clathria (s.s.) in its encrusting growth form,
a reduced hymedesmoid skeleton with erect fibre
nodes cored by plumose tracts of principal and
echinating spicules standing erect on the substrate (= `microcionid' architecture of Levi,
1960a), and smooth toxas. The critical difference
between these genera, therefore, is the possession
of the plumose, non-anastomosing fibre nodes,
whereas cofamilial encrusting genera have
hymedesmoid skeletal construction (Leptoclathria, Anaata, Cionanchora, Hymantho) or
a basal renieroid reticulation (e.g., some Antho,
Plocamilla). By comparison, erect or massive
Microcionidae commonly have reticulate or
plumo-reticulate skeletons (Clathria, Holopsam-
ma), renieroid or subisodictyal skeletons (e.g.,
Isociella, Isopenectya, Pandaros), or virtually
halichondroid skeletal architecture (Artemisina).
Many authors maintain the distinction between
Microciona and erect non-plumose genera, such
as Clathria (e.g., Levi, 1969; Wiedenmayer,
1977; Pulitzer-Finali, 1983; Uriz, 1984a-b;
Wintermann-Kilian & Kilian, 1984; Boury-Esnault & Lopes, 1985; Bergquist & Fromont,
1988). Cytological evidence to support this argument (Simpson, 1968a) is difficult to reconcile
completely with other morphological features.
Moreover, only a few species were examined in
this context, and the cytological characters themselves are obscure, not having been tested subsequently in other taxa. Conversely, no evidence
was found from biochemistry (protein
electrophoresis, free amino acid or carotenoid
protein profiles) to support the differentiation of
encrusting (hymedesmoid or microcionid)
species from erect (reticulate, non-plumose)
species (Hooper et al., 1992), but once again only
few taxa were examined in this context. Thus, I
have little clear evidence from non-skeletal sources on the boundary between these genera.
Overlying this classification based on skeletal
structure many authors subdivide Microcionalike species groups (at the generic level) according to modifications in microsclere geometry,
ornamentation on megascleres, or further reductions in skeletal structure (e.g., Anaata, Axociella
(of authors), Axocielita, Dictyociona, Leptoclathria, Pseudanchinoe, Thalyseurypon; e.g.,
Little, 1963; Koltun, 1976; Alcolado, 1980;
Hoshino, 1981; Lee & Gilchrist, 1985; Sim &
Bakus, 1986). Undoubtedly, recognising all these
genera is a matter of pragmatism, as this scheme
offers a convenient and relatively quick means to
manage a large and diverse assemblage of
microcionids. Many authors have questioned the
validity of these genera, debating generic distinction between Microciona and Clathria(Alander,
1942; Hechtel, 1965; Sara & Melone, 1966; Van
Soest, 1984b; Wiedenmayer, 1989; Hooper,
1990a). Some empirical evidence supports
counterarguments, that these character states intergrade between these genera (ontogenetically
and/or phenotypically), but no consensus has yet
been reached.
Van Soest (1984b) and Hooper (1990a) questioned separation of encrusting genera with
hymedesmoid or `microcionid' skeletons (e.g.,
Microciona) from erect growth forms with reticulate or plumoreticul ate skeletons (e.g., Clathria),
because they are clearly linked to some extent by
^
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FIG. 19. Type species of microcionid genera. A-B, Megaciella (Amphilectus pilosus Ridley & Dendy,
BMNH1887.5.2.125). C-E, Melonchela (M. clathrata Koltun, BMNH1963.7.29.7). F-G, Microciona (M.
atrasan guinea Bowerbank, BMNH1930.7.3.438). H-I, Naviculina (N. cliftoni Gray, BMNH1877.5.21.270).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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61
the ontogeny of the sponge individual (Simpson,
1968a). They argued from a phylogenetic basis
that the recognition of this character as being a
primary one confers exceptionally high levels of
homoplasy within the classification, cutting
across lines of apparent evolutionary decent
based on a combination of other characters (such
as the origin and disposition of structural
megascleres within the skeleton, spicule
geometry). For example, there are many thinly
encrusting species with hymedesmoid architecture and plumose spongin fibre nodes referable to
Microciona (e.g., M. maunaloa de Laubenfels, M.
microchela Hechtel), but also having two differentiated categories of auxiliary spicules (i.e., a
specialised ectosomal skeleton), which is characteristic of Thalysias species. Which genus do
these species then belong to, Microciona or
Thalysias, or do we recognise a third genus because they have both characters ? This third alternative was adopted by Bergquist & Fromont
(1988), largely following de Laubenfels' (1936a),
who recognised Axociella for microcionids with
Microciona-like fibre skeletons and Thalysiaslike ectosomal skeletons (although, like de
Laubenfels (1936a), they misinterpreted Axociella which is neither encrusting nor hymedesmoid).
This is a 'convenient classification' for managing
species, but it is also responsible for most of the
70 or so microcionid genera that exist today (i.e.,
approximately 1 genus per 6 species).
Few authors agree on the level of taxonomic
divergence, or the phylogenetic interpretation of
character polarity: is encrusting growth form and
hymedesmoid architecture a primitive or derived
strategy? Is a `microcionid' architecture
(hymedesmoid with ascending plumose fibre
nodes) a subsequent developmental stage of
hymedesmoid architecture (spongin fibres flat
against the substrate)? It is anticipated that
evidence from biochemistry or genetic sources
will eventually contribute towards the resolution
of these conflicts. A resolution is provided here
based on phylogenetic interpretation of morphology. We follow Levi's (1960a) arguments in considering three alternatives.
1) One generic taxon may be used to encompass
all the skeletal types from leptoclathriid
(hymedesmoid), `microcionid', to reticulate. This
option was used by Vosmaer (1933, 1935a-b),
who suggested that varying grades of skeletal
construction (from hymedesmoid, plumose to
renieroid reticulate) could be seen within populations of particular species and was apparently
largely related to ontogenetic development. Van
Soest (1984b), Hooper & Levi (1993a) and others
used this ontogenetic argument to merge
Microciona and Clathria (as well as Thalysias
and Clathria), and Hooper et al. (1992) presented
some empirical data to show that there was no
homogeneity in biochemical profiles to support
the retention of distinct taxa for encrusting versus
erect species.
2) Two separate taxa can be recognised for
species with hymedesmoid plumose
(Microciona) and reticulate (Clathria) architectures. This was the argument accepted by Levi
(1960a), Bergquist & Fromont (1988), and
others, on the basis that these skeletal structures
were consistent for populations of particular
species, and therefore represented fixed genetic
differences. Simpson (1968a) suggested further
that this option had some empirical support from
cytological evidence, although (unfortunately)
the cytological characters themselves are at the
moment not particularly useful taxonomic characters. A more pragmatic argument for the retention of the name Microciona is that it is in current
widespread use by sponge biochemists,
ecologists and experimental biologists, and
retaining this name provides some sort of
nomenclatural stability consistent with the previous literature. This argument, unfortunately,
has a pragmatic rather than biological basis.
3) The name Clathria could be used for an adult
terminal phase of skeletal architecture, related
directly to the ontogeny of the sponge, whereas
Microciona could be used for the juvenile phase
of the same species. Dictyociona is an example
of a Clathria with an intermediate Microcionalike skeleton, and Pseudanchinoe is an example
of a Microciona verging on a reticulate Clathrialike skeleton. This argument presupposes that
plumose skeletons are always precursors of, and
juvenile to, reticulate skeletons. But there are
several thinly encrusting species which do have
reticulate skeletons (e.g., Sara & Melone, 1966),
even though most plumose species are also persistently encrusting.
In recognising M icrociona at the subgeneric
level, Van Soest (1984b) implicitly also recognised a phylogenetic basis for the `microcionid'
skeletal specialisation, even though there were no
other corroboratory characters, such as unique
spicule geometries. The phylogenetic interpretation of this skeletal specialisation taken in the
present study is that Microciona is a persistently
encrusting sponge which consistently has nonanastomosing basal spongin fibres cored by nonanastomosing plumose spicule tracts or single
62zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
spicules, also lacking any ectosomal specialisation; it is an incompletely differentiated sister
taxon of Clathria and recognised here at the
subgenus level (Clathria (Microciona)). The Leptoclathria (flat, hymedesmoid) and Microciona
(ascending `microcionid' fibre nodes) conditions
are not differentiated, these being interpreted as
being more likely to be related to the thickness of
encrusting growth forms than anything else.
Echinoclathria). Wiedenmayer (1989) provided
further explanation of these relationships, although his nomenclatural decisions are not entirely correct.
Seriatula was erected for Spongia seriata Grant
(Gray, 1867) which is conspecific with 0. papilla
Bowerbank (e.g., Simpson, 1968a), and therefore
Seriatula becomes an objective synonym of Ophlitaspongia.
Ophlitaspongia Bowerbank, 1866
(Fig. 20A-B)
Pandaros Duchassaing & Michelotti, 1864
(Fig. 20C-E)
Ophlitaspongia Bowerbank, 1866: 14, 378; Wieden-
Pandaros Duchassaing & Michelotti, 1864: 88;
mayer, 1989: 59; Bergquist & Fromont, 1988: 113.
Not Ophlitaspongia; Dendy, 1896: 36; Hallmann,
1912: 253; Levi, 1960a: 58; Wiedenmayer, 1977:
140.
Seriatula Gray, 1867: 515; de Laubenfels, 1936a: 122.
TYPE SPECIES. Ophlitaspongia papilla Bowerbank,
1866: 378 (by original designation) (holotype
BMNH1910.1.1.395); = Spongia seriata Grant, 1826:
116 (Simpson, 1968a:37) (holotype BMNH1847.
9.7.14).
Encrusting bulbous growth form. Surface
microconulose, microscopically hispid.
Choanosomal skeleton isodictyal reticulate, with
compressed layer of spongin fibre lying on substrate and regularly reticulate spongin fibres arising from base, producing regular isodictyal fibre
network divided into primary ascending and
secondary transverse fibre elements. Primary
fibres cored by plumose tracts of choanosomal
principal styles, which also protrude from fibres
at acute angles resembling quasi-echinating spicules.
Secondary transverse fibres uncored or with unispicular tracts of usually smaller choanosomal
principal styles. Subectosomal skeleton paratangential, with scattered subectosomal auxiliary styles
lying on or near surface and also dispersed
throughout mesohyl. Ectosome without specialised
spiculation, but plumose tracts of choanosomal
principal megascleres protrude through surface.
Megascleres large and small, entirely smooth
choanosomal principal styles-subtylostyles, and
thin smooth subectosomal auxiliary styles.
Microscleres u-shaped toxas. Chelae absent.
REMARKS. This definition is based on type
material and Simpson's (1968a) description of
live populations. Ophlitaspongia papilla, was
shown by Simpson (1968a: 95) to be a synonym
of the type species of Microciona (M. atrasanguinea). A broader concept of Ophlitaspongia
(Wiedenmayer, 1989), (not 0. seriata (Grant)), is
identical to Echinoclathria (s.s.) (see remarks for
Schmidt, 1870: 59; de Laubenfels, 1936a: 123;
Wiedenmayer, 1977: 143; Van Soest, 1984b: 127.
TYPE SPECIES. Pandaros acanthifolium Duchassaing & Michelotti, 1864: 90 (by subsequent designation
of de Laubenfels, 1936a: 123)) (lectotype TMPOR57).
Bushy arborescent growth form. Surface highly
conulose, with flattened or lobate lamellae.
Choanosomal skeleton reticulate, with welldeveloped flattened spongin fibres (trabeculae)
cored by choanosomal principal subtylostyles
lying in all directions within fibres (from isodictyal reticulate to echinating) and with sparse
acanthostyles echinating or also incorporated into
fibres. Subectosomal skeleton radial, reduced to
single long subectosomal auxiliary subtylostyles
protruding through surface and also scattered
throughout mesohyl. Ectosome without special
spicules. Megascleres smooth choanosomal principal subtylostyles-tylostyles, often with slightly
rhabdose bases and terminal or subterminal basal
swellings, long curved or straight subectosomal
auxiliary subtylostyles, and lightly acanthose or
rarely smooth styles `echinating' fibres.
Microscleres absent.
REMARKS. This definition is based on the type
and a fragment of the type MNHNLBIMDNBE
1309, specimen BMNH1884.7.11.2 and description of live populations by Van Soest (1984b).
Important features are: 1) the prominently flattened fibres cored by smooth slightly rhabdose
principal subtylostyles-tylostyles (more reminiscent of Rhabderemiidae than of Microcionidae);
2) the sparse, lightly spined styles which more
closely resemble a second category of principal
spicules than than they do echinating (accessory)
spicules typical of other Microcionidae; furthermore, these spicules are only rarely seen echinating fibres, but more commonly they are
incorporated into them together with the principal
megascleres; and 3) the long subectosomal
auxiliary subtylostyles protruding through the
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
63
surface (more similar to a reduced Raspailiidae,
such as Echinodictyum or Ceratopsion, than to
typical Microcionidae).
Pandaros could be legitimately included in
either Raspailiidae or Microcionidae. Van Soest
(1984b) noted that only the rare echinating acanthostyles in P acanthifolium gives any cause to
link it to the Microcionidae at all. He speculated
that it might be necessary to erect a separate
family for the species, or even remove it from the
Poecilosclerida altogether, as it also shows affinities with axinellids such as Ptilocaulis. There
are no microscleres to give any further clues as to
its affinities. Pandaros is maintained as a separate
genus and tentatively included in Microcionidae.
Of 12 species referred to Pandaros, only the type
clearly belongs here. Raspailia kasumiensis
Tanita (MMBS SIS-052) was assigned to Pandaros (Hooper, 1990a).
Wiedenmayer (1977) merged Thalyseurypon
with Pandaros, because he considered that its
type species had architecture closely comparable
to P acanthifolium (Hechtel, 1965), but this is not
upheld here. The only features these genera have
in common is lacking microscleres. Wiedenmayer (1977) also speculated that the genus had
a close relationship with Echinoclathria (=
Holopsamma as defined here), based on alleged
similarities in skeletal architecture, and he suggested that the two genera probably intergrade in
habit and spiculation, but these suggested affinities are not evident in relevant specimens.
Paradoryx Hallmann, 1920
(Fig. 20E-G)
Paradoryx Hamann, 1920: 767; de Laubenfels,
1936a: 109.
TYPE SPECIES. Clathria dura Whitelegge, 1901: 83
(by original designation) (holotype AMG3046).
Arborescent, flabellate growth form with compressed branches. Surface even, smooth.
Choanosomal skeleton reticulate, with heavy
spongin fibres forming irregular anastomoses in
axis becoming more regular and rectangular
towards periphery. Spongin fibres differentiated
into primary ascending and secondary connecting
components, cored by auxiliary styles and heavily echinated by acanthostyles. Subectosomal
skeleton plumose, with bundles of auxiliary
styles forming plumose brushes on peripheral
fibres. Ectosome fibrous, without specialised
skeleton, and peripheral fibres form more-or-less
reticulate structure through which choanosomal
spicules protrude. Megascleres entirely smooth,
hastate or quasidiactinal auxiliary styles or subtylostyles, and echinating acanthostyles with
large and even spination. Microscleres palmate
isochelae resembling arcuate forms with lateral
alae fused to shaft for about 1/2 alae length, and
shaft greatly curved, thickened. Toxas absent.
REMARKS. Ectosomal fibres of Paradoryx are
unspecialised, identical to choanosomal fibres in
size and form, unlike the Callyspongia-like ectosome of Marleyia. Hallmann (1920) erected
Paradoryx for Wilsonella species (with auxiliary
megascleres coring fibres) which had arcuate
chelae instead of palmate isochelae. Loss or replacement of principal megascleres varies considerably between otherwise allied microcionid
taxa and this feature is considered to be of lesser
significance at the generic level than assumed by
Hallmann (Hooper, 1990a; Hooper et al., 1990).
Similarly, isochelae described by Hallmann
(1920) as arcuate in the type species are modified
(curved, thickened) palmate forms.
Apart from the type species, Hallmann (1920)
included: Dictyocylindrus pinifonnis Carter and
W oxyphila (both of which belong to Clathria
(Clathria)), and Clathria elegantula Ridley &
Dendy and Wilsonella curvichela Hallmann
(which are referred here to Clathria
(Dendrocia)).
Paratenaciella Vacelet & Vasseur, 1971
(Fig. 20H-0
Paratenaciella Vacelet & Vasseur, 1971: 103.
TYPE SPECIES. Paratenaciella microxea Vacelet &
Vasseur, 1971: 103 (by original designation) (holotype
MNHNLBIMD.W27).
Encrusting growth form. Surface uneven,
hispid. Choanosomal skeleton hymedesmoid,
with spongin fibres reduced to basal layer on
substrate and with ascending fibre nodes enclosing bases of choanosomal principal subtylostyles
standing perpendicular to substrate. In thicker
sections up to 3-4 spicules form ascending
plumose tracts, protruding through surface.
Echinating megascleres absent. Subectosomal
skeleton plumose or paratangential composed of
bundles of subectosomal auxiliary subtylostyles
often forming brushes around protruding principal spicules and protruding through ectosome.
Megascleres entirely smooth choanosomal principal subtylostyles, and subectosomal auxiliary
subtylostyles with microspined bases and
polytylote shafts. Microscleres palmate
^
MEMOIRS OF THE QUEENSLAND MUSEUM
64zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 20. Type species of microcionid genera. A-B, Ophlitaspongia (0. papilla Bowerbank,
BMNH1910.1.1.395). C-E, Pandaros (P. acanthifolium Duchassaing & Michelotti, BMNH1884.7.17.2). F-G,
Paradoryx (Clathria dura Whitelegge, AMG3046). H-I, Paratenaciella (P. microxea Vacelet & Vasseur,
MNHNDJV27).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
65
isochelae, including contort forms, and oxeote
toxas.
REMARKS. Paratenaciella has a Microcionalike choanosomal fibre skeleton composed of
non-anastomosing, plumose fibre nodes, but
lacks acanthostyles (similar to Axociella,
Axocielita, Tenaciella), has polytylote modifications to auxiliary spicules (similar to Fisherispongia ferrea de Laubenfels and Microciona
aceratoobtusa Carter), and has modified (oxeote)
toxas, like those in Artemisina archegona Ristau
(although type material shows that they do
resemble true microxeas). This latter feature appears to be the only unique character for the type
species, and as far as is known it is also unique
amongst the Microcionidae. Paratenaciella
shows some similarities to Esperiopsis
canaliculata Whitelegge (which is referred here
to Axociella) in its spiculation, although P.
microxea has a microcionid architecture,
microxeote microscleres and undifferentiated ectosomal-subectosomal megascleres. The genus is
monotypic and is synonymised with Clathria
(Microciona).
Plectispa Lendenfeld, 1888
(Fig. 21C-D)
Plectispa Lendenfeld, 1888: 225; Topsent, 1894a: 19;
de Laubenfels, 1954: 164.
TYPE SPECIES. Plectispa macropora, in part, Lendenfeld, 1888: 226 (by subsequent designation of
Hallmann, 1912: 204) (holotype AMG9159).
Lobodigitate growth form. Surface 'honeycomb'
reticulate. Choanosomal skeleton reticulate, with
well developed spongin fibres forming more-orless regular oval or eliptical meshes, sparsely
cored and abundantly echinated by principal subtylostyles, and fewer auxiliary styles-tornotes
also in fibres. Echinating megascleres less
predominant in periphery than in axis. Some
detritus also incorporated into choanosomal
fibres. Subectosomal skeleton plumose, with
brushes of principal subtylostyles from ascending
fibres of peripheral skeleton protruding through
surface, surrounded by paratangential loose
bundles or individual auxiliary spicules.
Megascleres smooth principal subtylostyles, and
long curved, setaceous or sinuous auxiliary styles
or quasi-diactinal tornotes, both with blackened
axial canals. Microscleres palmate isochelae, including contort forms. Toxas absent.
REMARKS. This genus is poorly defined due to
the dubious status of all of Lendenfeld's syntypes,
inaccuracies in Lendenfeld's (1888) original
description, and misinterpretation of the species
(e.g., Clathria macropora; Whitelegge (1901:
91) is different from C. macropora Lendenfeld
(1888)). Hallmann (1912: 165) attempted to sort
out these problems but was only partially successful. He rejected Whitelegge's proposed merger of
P. macropora with Echinonema levis Lendenfeld
(1888), but instead he synonymised the latter with
C. macropora (sensu Whitelegge), as Crella incrustans Carter, var. levis Lendenfeld. However,
Hallmann's (1912) remarks and assumptions
concerning the genus are not entirely correct (see
remarks below for Clathria macropora and
Holopsamma macropora), and there is no
evidence that he actually re-examined relevant
type material. His nomenclatural decisions are
erroneous and not supported here.
There are 4 type specimens and one other
specimen in the AM and BMNH which bear the
name macropora'. 1) AMZ959 (with the label
'dry, cotype') from Port Jackson, NSW, is the
so-called 'type' of Clathria macropora (sensu
Whitelegge), and is a species of Crella. It is
probably a syntype of Lendenfeld's (1888)
Echinonema levis, but that assumption is difficult
to corroborate due to the loss of Lendenfeld's
'key-list' from the Australian Museum (E. Pope
& E Rowe, pers.comm.). 2) AMZ466 (with label
stating 'spirit preserved, ?type') from Port
Stephens, NSW is the lectotype of Lendenfeld's
(1888: 221) Clathria macropora, later referred to
Wilsonella by Hallmann (1912: 240). It is neither
a Wilsonella, nor closely related to such species
as W australiensis Carter, but is a Clathria lacking echinating megascleres and having an axially
compressed skeleton (e.g., Clathria (Axociella)).
3) AMG9159 (with the label stating 'dry, type'),
probably from Port Jackson, NSW (i.e., there is a
discrepancy between the published locality of
Torres Strait, Qld. and the museum specimen
label and register), is the holotype of Plectispa
macropora. 4) BMNH1925.11.1.555 (with label
stating Plectispa macropora, dry, Dendy coll.')
from Manly Beach, NSW, is identical to Clathria
multipes Hallmann. 5) Another specimen
(BMNH1957.8.30.2, with label stating Plectispa
macropora, dry') belongs to Clathria lendenfeldi.
Consequently, only a single syntype is valid becoming the holotype of P. macropora, and the
diagnosis of Plectispa is based on this specimen.
Hallmann (1912:205) referred the type species
(and genus Plectispa) to Wilsonella, but then
returned it to Clathria (Hallmann, 1920: 768).
The species has a 'honeycombed reticulate'
MEMOIRS OF THE QUEENSLAND MUSEUM
66zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
growth form and smooth echinating spicules (not
acanthose as supposed by Lendenfeld (1888) and
Hamann (1912)). It also has long, setaceous,
quasi-diactinal auxiliary megascleres, which also
core fibres, scattered throughout the mesohyl and
associated with spicule brushes on the surface.
These characters indicate that Plectispa belongs
to Holopsamtna, and the species shows some
resemblance to H. ramosa (Hallmann, 1912) and
H. laminaefavosa Carter, 1885b (sensu Wiedenmayer, 1989).
Plocamia Schmidt, 1870
(Fig. 21A-B)
Plocamia Schmidt, 1870: 62.
Dirrhopalum Ridley in Ridley & Duncan, 1881: 477.
spicules are much better developed in Plocamilla.
Plocamia is the earliest available name for a
group of Antho-like ('plocamiform') microcionids having basal tylotes and echinating
spicules (see remarks for Holoplocamia, Lissoplocamia).
Plocamilla Topsent, 1928
(Fig. 21E-F)
Topsent, 1928a: 63; Levi, 1960a: 80; Levi,
1960b: 760; Pulitzer-Finali, 1973: 40; Simpson,
1968a: 95; Van Soest, 1984b: 26, 29, 125; (not
Burton, 1935a: 402).
Plocam illa
TYPE SPECIES. I sodiciy a coriacea Bowerbank,
1874: 136 (by original designation of Topsent, 1928a:
63) (holotype BMNH1910.1.1.251).
Plocam ia gym nazusa Schmidt, 1870:
TYPE SPECIES. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
DIAGNOSIS. Encrusting growth form. Surface
62 (by subsequent designation (Burton, 1935a: 401))
uneven, porous, microscopically hispid.
(holotype possibly LMJG, schizotype
Choanosomal skeleton renieroid, with regular
MNHNLBIMDCL1105L).
DIAGNOSIS. Encrusting growth form. Surface
even, microscopically hispid. Choanosomal
skeleton renieroid, with tylotes forming basal
reticulation, interconnected at nodes by light
spongin fibres, and with plumose columns of
choanosomal principal styles and echinating
styles embedded in basal renieroid skeleton, individually or in bundles, largest protruding
through surface. Subectosomal skeleton with
paratangential and erect tracts of subectosomal
auxiliary styles. Megascleres robust slightly
curved choanosomal principal styles with
microspined or smooth bases, robust tylotes with
swollen microspined bases, long subectosomal
auxiliary styles with smooth or microspined
bases, smaller echinating styles with microspined
bases. Microscleres palmate isochelae and wingshaped toxas.
basal reticulation of acanthose strongyles in unior paucispicular tracts, overlaid by plumose
brushes of larger choanosomal principal styles
and echinating acanthostyles projecting perpendicularly from nodes of renieroid spicule
skeleton. Ectosomal skeleton contains plumose
or paratangential brushes of subectosomal
auxiliary styles and projecting echinating acanthostyles from peripheral nodes of choanosomal
renieroid skeleton. Megascleres acanthose strongyles or tylotes with spines mostly on basal ends,
larger choanosomal principal styles-subtylostyles with spined bases, subectosomal auxiliary
styles-subtylostyles with smooth or microspined
bases, and small echinating acanthostyles with
few spines concentrated mainly on basal end.
Microscleres palmate isochelae, smaller wingshaped toxas, and larger accolada toxas with
spined points.
REMARKS. Ridley (in Ridley & Duncan, 1881)
suggested that Plocamia Schmidt was preoccupied by Plocamium Lamouroux, 1828, a
seaweed, and consequently proposed the replacement name Dirrhopalum Ridley. However, under
Article 56b of the ICZN (Anonymous, 1985) this
is an unjustified emendation and Plocamia
stands. Bergquist & Fromont (1988) also used
Plocamia over Dirrhopalum.
This definition is based on slides of the
holotype and published descriptions (Schmidt,
1870; Ridley in Ridley & Duncan, 1881). The
type species is essentially the same as Plocamilla
in structure and spicule geometry, but spines on
both the basal tylote spicules and echinating
REMARKS. This definition is based on the
holotype and descriptions of the type species
(Topsent, 1928a; Levi, 1960a). Plocamilla has
spiculation similar to Plocamia Schmidt, but it
has an encrusting growth form and therefore its
skeleton is not clearly differentiated into primary
or secondary lines (Topsent, 1928a: 63; Levi,
1960a: 80). This skeletal development is probably related to growth form and not an important
generic character. The genus was not formally
diagnosed until Burton (1935a: 402) differentiated a number of `plocamiform' genera, all of
which had acanthostrongyles or acanthostyles
forming the basal renieroid skeleton. Burton emphasised the diagnostic value of choanosomal
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
67
acanthostrongyles ('dumbell spicules') which
united those genera, but he differentiated them by
their skeletal architecture and the presence or
absence of various spicule types.
Holoplocamia is a synonym of Plocamilla
(Levi, 1960a: 80), although Little (1963: 47) argued to the contrary. Levi (1960a) also noted that
the skeleton of P coriacea was very close to
Antho and Dictyoclathria (both of which are objective synonyms), lacking special echinating
spicules and having acanthostyles instead of
acanthostrongyles in the basal renieroid skeleton.
Levi (1960a) and Pulitzer-Finali (1973: 40) were
cautious in interpreting whether Plocamilla was
distinct from Antho, because the spined acanthostyles and smooth principal styles could not be
consistently differentiated in all taxa. For example, P. elegans (Ridley & Dendy) does not
show any clear separation between the smaller
category of smooth principal styles and the more
sparsely spined acanthostyles (Pulitzer-Finali,
1973). On that basis, Pulitzer-Finali suggested
that future studies may show that P. elegans
should be referred to Antho, whereas P coriacea
has clearly differentiated coring and echinating
megascleres, and should remain in Plocatnilla.
Levi (1960a), Simpson (1968a), Pulitzer-Finali
(1973) and Van Soest (1984b) maintained
Plocamilla and Antho as separate genera, the
former having echinating acanthostyles and
predominantly (acantho)strongyles in the
renieroid skeleton. However, all these authors
admitted that the two genera may be too similar
to maintain sufficient generic separation. Van
Soest & Stone (1986) noted that the genus should
probably be merged with Antho, together with
other plocamiform genera containing renieroid
skeleton of acanthose megascleres, and this suggestion is supported here (i.e., Antho (Plocamia)).
Simpson (1968a: 95) found that although
Plocamilla and Microciona were easily differentiated by their skeletal construction and spicule
geometries, they were remarkably similar in their
cytological characteristics. These findings contradict the more obvious similarities between
microcionid genera, based on spicule geometry
and skeleton construction. Microciona, Clathria
and Thalysias are a relatively homogenous group
in skeletal construction, compared to any inferred
relationship between Plocamilla and Microciona
for example. Probably Simpson's (1968a) cytological characters have not been incorporated into
the classification, nor have the the implications of
his findings to phylogeny of demosponges been
widely discussed.
Plocamiopsis Topsent, 1904
(Figs 21G-H, 22A)zyxwvutsrqponmlkjihgfe
Plocam iopsis
Topsent, 1904a: 155; Burton, 1935a:
402.
TYPE SPECIES. Plocam iopsis signora Topsent,
1904a: 155 (by monotypy) (holotype MNHNLBIMDT947).
DIAGNOSIS. Encrusting growth form. Surface
even, hispid. Choanosomal skeleton irregularly
renieroid, with basal reticulation of acanthostrongyles in uni- or bispicular tracts lying on substrate. Choanosomal principal subtylostyles,
smaller acanthostyles and spicules intermediate
to both embedded individually in basal skeleton,
standing perpendicular to substrate. Subectosomal skeleton plumose, with ascending
brushes of subectosomal auxiliary subtylostyles
surrounding principal megascleres. Ectosome
without specialised spiculation, but with both
tangential and plumose tracts of subectosomal
auxiliary spicules protruding through surface.
Megascleres choanosomal principal subtylostyles with spines on bases and proximal region of
shaft, smaller acanthostyles with aspinose points,
and subectosomal auxiliary subtylostyles with
spined bases. Microscleres cleistochelae and
smooth wing-shaped toxas.
REMARKS. Plocamiopsis is a member of de
Laubenfels' (1936a) plocamiform group of sponges having a basal renieroid skeleton of acanthostrongyles. It differs from others in this group
(Dirrhopalum, Heteroclathria, Holoplocamia,
Lissoplocamia, P locamilla) having cleistochelae, which are also seen in Colloclathria and
Quizciona. Cleistochelae are hypersilicified palmate isochelae with the front alae fused (Fig.
22A) but probably have little phylogenetic significance given the complete transitional series
from palmate to cleistochelate (Fig. 76G).
Plocamiopsis is referred to Antho (Plocamia).
Protophlitaspongia Burton, 1934
(Fig. 22B-C)
Prot ophlit aspongia
Burton, 1934a: 562.
TYPE SPECIES. Siphonochalina bispiculat a Dendy,
1895: 246 (by original designation) (lectotype
NMVG2319).
Lamellar, tubulo-digitate growth form, with
osculum on apex of each tube. Surface uneven,
microscopically hispid. Choanosomal skeleton
subisodictyal reticulate, with more-or-less
regularly anastomosing heavy spongin fibres
^
MEMOIRS OF THE QUEENSLAND MUSEUM
68zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 21. Type species of microcionid genera. A-B, Plocamia (P. gymnazusa Schmidt, MNHNDCL1105L). C-D,
Plectispa (P. macropora sensu Lendenfeld, AMG9159). E-F, Plocamilla (Isodictya coriacea Bowerbank,
BMNH1910.1.1.251). G-H, Plocamiopsis (P. signata Topsent, MNHNDT947).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU
^
69
forming primary ascending and secondary
transverse lines. Primary fibres cored by multispicular tracts of longer auxiliary oxeas; secondary fibres usually aspiculose occasionally with
only single coring auxiliary oxeas. Fibres sparsely echinated by smaller, smooth hastate oxeas
(presumed to be homologous with principal
spicules). Ectosome lacks specialised spiculation, but has plumose erect brushes of oxeas from
peripheral skeleton protruding through surface.
Megascleres modified diactinal, including longer
smooth hastate auxiliary oxeas, and smaller smooth
hastate principal oxeas. Microscleres absent.
REMARKS. The genus contains 7 previously
named species: P bispiculata (Dendy, 1895: 246)
from Port Phillip Bay, Victoria, P. oxeata Burton
(1934a: 562) from the Papuan Pass, northern
Australia, P. ada de Laubenfels (1954: 96) from
Ponape, P aga de Laubenfels (1954: 97) from the
central west Pacific, P antillana Pulitzer-Finali
(1986: 138) from the West Indies, Echinochalina
(P.) laboutei Hooper & Levi (1993a: 1277) and
E. (P.) bargibanti Hooper & Levi (1993a: 1280)
both from New Caledonia. These species
resemble Haplosclerida to some extent; P. anti!lanais a haplosclerid, (Niphatidae). Similarly, de
Laubenfels' (1954), P ada and P aga, are placed
in the Phoriospongiidae leaving 4 known species
and 4 new Australasian species described below.
The genus has a characteristic three-dimensional ectosomal architecture reminiscent of
Hemigellius and Anzphimedon (Haplosclerida:
Niphatidae), with only oxeas as megascleres.
However, smooth hastate oxeas echinating fibres,
with a similar geometry but smaller size than
coring spicules, gives some clues as to probable
phylogenetic relationships (i.e., Microcionidae,
some lophonidae, and Raspailiidae). This pattern
of spiculation is seen in Echinochalina, and to a
lesser extent Echinoclathria as defined here. It is
particularly well developed in E. axinelloides,
and for this reason Burton (1934a: 562) tentatively referred Protophlitaspongia to the Microcionidae. Conversely, de Laubenfels (1936a: 54,
1954: 96) suggested that the genus was closer to
Guitarra (although lacking their peculiar
microscleres), or Liosina, and he referred
Protophlitaspongia to the Desmacidonidae, but he
was probably referring mainly to his two
Micronesian species in doing this. Pulitzer-Finali
(1986: 138) followed similar reasoning, placing
the genus in Esperiopsidae, but none of these
species belong to Protophlitaspongia in any case.
Further clues as to the most appropriate placement of this genus can be seen from other characteristics: the palmate isochelae in E. (P.)
bargibanti, the accolada toxas in E. (P.) oxeata,
and the plumose ectosomal skeleton, composed
of auxiliary spicule brushes (typical of
Microcionidae, Axinellidae and Raspailiidae) in
all species. The present interpretation of Protophlitaspongia suggests affinities with Echinoclathria and Holopsanuna microcionids and the
reticulate Raspailiidae, and in the distribution of
its megascleres (i.e., auxiliary spicules in fibres
and peripheral skeleton, principal spicules
echinating fibres) it is assigned to Echinochalina.
Pseudanchinoe Burton, 1929
(Fig. 22D-E)
Pseudanchinoe Burton, 1929a: 433; de Laubenfels,
1936a: 109.
TYPE SPECIES. Stylostichon toxiferum Topsent,
1913a: 621 (by original designation and monotypy)
(holotype MNHNLBIMDT1612).
Massive subspherical growth form. Surface
uneven, apical surface conules. Choanosomal
skeleton plumoreticulate, with spongin fibres
forming primary plumose ascending columns,
partially interconnected by transverse secondary
fibres, both cored by multispicular tracts of
choanosomal principal subtylostyles and heavily
echinated by acanthostyles. Subectosomal
skeleton plumose, non-anastomosing, composed
of multispicular plumose tracts of choanosomal
principal spicules from peripheral fibres surrounded by plumose bundles of subectosomal
auxiliary subtylostyles. Ectosomal skeleton with
second tier of subectosomal auxiliary subtylostyles forming dense palisade on surface.
Megascleres choanosomal principal subtylostyles with or without basal spines, subectosomal
auxiliary subtylostyles with microspined bases,
and echinating acanthostyles varying from
smaller forms with even spination to larger forms
with aspinose points. Microscleres oxhorn and
accolada toxas. Chelae absent.
REMARKS. Burton (1929a) recorded palmate
isochelae in the type species but none were found
in the holotype nor did Topsent (1913a) record
any in his original description. In fact the absence
of isochelae and the possession of plumoreticulate skeletal architecture are virtually the only
two diagnostic features that distinguish
Pseudanchinoe from other Clathria (s.s.).
Similarly, Topsent (1913a) stated that there were
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microspines on the points of toxas, but these were
not seen in the holotype.
The type species of Pseudanchinoe is intermediate between Clathria and Thalysias, having
the ectosomal structure of the latter (i.e., with a
thick continuous palisade of erect spicules overlaying a similar subectosomal skeleton), but with
spicule composition of the former (single
category of auxiliary styles), further supporting
the decision to merge these two genera in a single
taxon (Clathria s.1.). In skeletal structure and
diversity of megascleres the genus could be placed
in the series: Dendrocia (single category of structural spicule in the entire skeleton), Clathria (one
category of auxiliary spicule and one category of
principal spicule), Pseudanchinoe (one category
of auxiliary spicule forming a continuous ectosomal palisade), Thalysias (two categories of
auxiliary spicules the smaller forming a continuous ectosomal palisade, and one category of
principal spicule), and Antho (some of which
have two categories of auxiliary spicules and two
categories of principal spicules).
Burton (1929a) remarked on the apparent
similarities between Pseudanchinoe and Anchinoe
Gray (= Phorbas Duchassaing & Michelotti;
Anchinoidae). The latter genus has plumose
columns of intermingled acanthostyles and tornotes (or oxeas), which are echinated by acanthostyles, but these similarities are superficial.
Anchinoidae have tangential ectosomal diactinal
megascleres and plumose columns of
choanosomal megascleres in which acanthostyles predominate (e.g., Van Soest, 1984b: 86). Of
the numerous species assigned to Pseudanchinoe
(e.g., de Laubenfels, 1936a: 109) most are interpreted here as belonging to Clathria, having
secondarily lost their isochelae. Australian examples are C. (C.) caelata Hallmann, C. (C.)
costifera Hallmann, C. (C.) inanchorata Ridley
& Dendy, and C. (C.) partita Hallmann. Two
other species belong to Clathria (Thalysias) (C.
dentata Topsent and C. fascicularis Topsent,
which are synonymous) (Van Soest, 19846). Of
contemporary authors only Koltun (1976) recognises this genus whereas Van Soest (1984b) and
(Wiedenmayer, 1989) merged it with Clathria. It is
included here in Clathria (Clathria).
Qasimella Thomas, 1974
Qasimella Thomas, 1974: 311.
TYPE SPECIES. Qasimella indica Thomas, 1974: 311
(by original designation)(holotype CMFRIT84/1 not
seen).
Specialised tubular growth form, with apical
oscules and central cavity running longitudinally
through body. Surface smooth, even.
Choanosomal fibres apparently absent, and
skeletal structure poorly defined with more-orless longitudinal tracts of choanosomal principal
subtylostyles bound together at nodes by collagen. Echinating acanthostyles absent. Subectosomal skeleton rudimentary, composed of
subectosomal auxiliary subtylostyles lying just
below surface, not protruding through it. Ectosomal skeleton absent. Megascleres
choanosomal principal subtylostyles with spined
bases, and slender smooth subectosomal
auxiliary subtylostyles. Microscleres described
as 'arcuate' isochelae and wing-shaped toxas.
REMARKS. The holotype and paratypes
(CMFRI T8412) of Q. indica are housed at the
Central Marine Fisheries Research Institute,
Mandapan Camp, Cochin, India. A slide of the
holotype is housed at the IM (A.K. Mandal, pers.
comm.).
The genus is monotypic and known only from
its original description (Thomas, 1974). It is an
unusual, very thin-walled hollow tubular sponge
with pseudo-syconoid construction, and attached
to the substrate by short peduncles reminiscent of
Aulospongus Norman (Raspailiidae). Apart from
spicule diversity and geometry, which place the
type species in Microcionidae, we know of few
other characters that give clues as to its affinities
with other microcionids. From its skeletal structure, or lack of it, Qasimella shows similarities to
Artemisina. It is also possible that Q. indica is a
larval sponge, but until type material is seen its
published description suggests that it has affinities to Artemisina. The chelae have been
described as 'arcuate', but this cannot be confirmed and must be regarded as suspect (e.g., they
may be merely modified, curved, palmate ones).
Quizciona de Laubenfels, 1936
(Fig. 22F-G)
Quizciona de Laubenfels, 1936a: 111.
TYPE SPECIES. Microciona heterospiculata
Brondsted, 1924: 465 (by original designation) (fragment of type BMNH1901.12.26.13).
Encrusting growth form. Surface hispid, uneven. Choanosomal skeleton differentiated into
two components: renieroid basal layer composed
of spongin fibres lying on substrate cored by
bispicular tracts of acanthostyles forming triangular meshes; basal fibres ascending non-anas-
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71
tomosing fibre nodes echinated by multispicular
plumose tracts of both choanosomal principal
subtylostyles and echinating acanthostyles (of
same morphology as basal renieroid skeleton
spicules) embedded and erect on substrate. Ectosomal skeleton paratangential, composed of
two size classes of subectosomal auxiliary subtylostyles, forming occasional brushes and scattered in mesohyl matrix near periphery.
Megascleres longer choanosomal principal subtylostyles with heavily microspined bases,
shorter acanthostyles with aspinose points (forming both renieroid basal skeleton and echinating
fibre nodes), and two size classes of auxiliary
subtylostyles with smooth or spined bases.
Microscleres palmate isochelae, cleistochelae
and small accolada toxas.
REMARKS. The original description differs
slightly from the definition above. In particular,
the very small, thin toxas were overlooked, as was
a renieroid basal skeleton typical of de
Laubenfels' (1936a) plocamiform sponges.
Brondsted's (1924) description of Quizciona suggests that it is close to Microciona with a
hymedesmoid basal skeleton and plumose ascending fibre nodes but having cleistochelae in
addition to palmate isochelae. The type material
shows the species more closely related to Antho
with a renieroid basal skeleton. Smaller acanthostyles and larger principal styles echinating basal
fibres is similar to Plocamilla, whereas acanthostyles rather than acanthostrongyles coring the
basal fibres is similar to Antho, and this mix of
characters illustrates the difficulty in separating
these two taxa.
Cleistochelae have been recorded in several
microcionids (Microciona cleistochela Topsent,
M. clathrata Whitelegge, M. chelifera Levi,
Clathria simpsoni Van Soest, C. toxipraedita
Topsent and Colloclathria ramosa Dendy).
Alander (1942) and Van Soest (1984b) suggested
that cleistochelae are modified palmate isochelae
(with fused chelate teeth) which have arisen
several times independently within the
Microcionidae, and consequently they do not
have primary taxonomic significance. Interestingly, Brondsted (1924) also remarked on the
similarities between cleistochelae of
Microcionidae and clavidiscs of the hypercalcified sponge Merlia Kirkpatrick, supporting current theories on the origin and affinities of the
`sclerosponges' with the demosponges (e.g.,
Vacelet, 1985).
Bergquist & Fromont (1988) merged M.
heterospiculata with Clathria mortensenii and
suggested that Quizciona should be abandoned
on the basis that they did not find cleistochelae in
their material, contrary to de Laubenfels' (1936a)
description. This synonymy is not upheld here,
although Bergquist's (1961a: 39) record of M.
heterospiculata from northern New Zealand may
be Clathria mortensenii. I interpret Quizciona,
based strictly on the type specimen, as a synonym
of Antho (Antho).
Ramoses de Laubenfels, 1936
(Fig. 22H-I)
Ramoses de Laubenfels, 1936a: 109.
TYPE SPECIES. Clathria pauper Brondsted, 1927: 3
(by original designation) (fragment of type
BMNH1930.11.5.2).
Arborescent, tubulo-digitate growth form. Surface shaggy, uneven, microscopically hispid.
Choanosomal skeleton irregularly plumo-reticulate, with vaguely ascending multispicular
primary tracts interconnected by transverse
paucispicular secondary tracts cored by both
smooth and partially spined choanosomal principal styles and echinated by plumose bundles of
smaller acanthostyles. Spongin fibres not seen
but spicules united by moderate quantities of
collagen. Subectosomal skeleton plumose, with
bundles of subectosomal auxiliary subtylostyles
protruding through surface. No special ectosomal
skeleton. Megascleres longer entirely smooth
choanosomal principal styles, slightly shorter
principal subtylostyles with spines on bases and
distal end of shaft, small slender echinating acanthostyles with even spination, and subectosomal
auxiliary subtylostyles with spined bases.
Microscleres accolada toxas, some with contort
centres. Chelae absent.
REMARKS. The holotype of C. pauper has not
yet been discovered in the Brondsted collection
at UZM (0. Tendal, pers. comm.), but a fragment
of it is held at the BMNH. A fragment of C.
pauper (with second label stating Trotoclathria
antarctica, 29.1.19?? [illegible], 340 fathoms, M
Burton') is in Sydney (AMZ2239). It is possible
that this material is also a fragment of the
holotype, but its status and origin are not certain.
Brondsted's (1927) original description conforms
closely to the type material, although he appears
to have overlooked the thin accolada toxa
microscleres (some with asymmetrical contort
central curvature).
^
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FIG. 22. Type species of microcionid genera. A, Plocamiopsis (P. signata Topsent, MNHNDT947). B-C,
Protophlitaspongia (Siphonochalina bispiculata Dendy, NMVG2319). D-E, Pseudanchinoe (Stylostichon
toxiferum Topsent, MNHNDT1612). F-G, Quizciona (Microciona heterospiculata Brondsted,
BMNH1901.12.26.13). H-I, Ramoses (Clathria pauper Brondsted, BMNH1930.11.5.2).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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Ramoses is similar to Pseudanchinoe in lacking
chelae microscleres, and Koltun (1976: 155)
synonymised the type species of Ramoses with P.
toxiferuin Topsent, the type species of
Pseudanchinoe (both species are from the Ross
Sea, Antarctica). This synonymy is confirmed
here from type specimens and both species are
lipochelous Clathria (Clathria). Koltun (1976)
suggested that C. pauper was a synonym of
Microciona toxifera (Topsent), also from the Antarctic, but this synonymy is rejected. Comparison of relevant type specimens shows that the
two species differ substantially in their skeletal
structure, spicule size and spination on
megascleres.
REMARKS. The type species is from the Indowest Pacific region where most species occur.
The primary and only consistent morphological
feature that distinguishes Rhaphidophlus from
other Clathria-like taxa is the specialised ectosomal skeleton, consisting of two differentiated
categories of auxiliary subtylostyles which form
brushes on the surface (either as discrete brushes
(s.s.)or in a continuous palisade). This is identical
to the condition described for Thalysias (which
has seniority).
Hallmann (1912), followed most recently by
Van Soest (1984b), suggested that the two sizes
of auxiliary megascleres may represent age differences in spicules, in which case the genus has
a distinct localisation of adult and juvenile
megascleres. This assertion, however, does not
Rhaphidophlus Ehlers, 187
yet have any empirical (experimental) support.
(Fig. 23A-C)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
This difference in size/age of auxiliary spicules is
crucial to the definition of Rhaphidophlus, beRhapidophlus Ehlers, 1870: 19,31; Ridley, 1884a: 449cause some species of Clathria, Microciona and
453; Ridley & Dendy, 1887: 151; Topsent, 1894a:
Dendrocia have a dermal skeleton of similar
19, 1925: 658, 1928a: 61, 1932: 98; Thiele, 1903a:
structure but containing only one sort of auxiliary
957; Dendy, 1905:170; Hallmann, 1912: 175,1920:
spicule (C. impelfecta, C. striata, C. pyramida).
769; Burton, 1932a: 320; de Laubenfels, 1936a:
112; Levi, 1960a: 56; Bergquist, 1965: 168; Similarly, some species of Antho (e.g., A. ridleyi)
Simpson, 1968a: 101, 104-106; Wiedenmayer,
also have relatively dense ectosomal brushes, but
1977: 140; Van Soest, 1984b: 109; Bergquist &
these consist of a single undifferentiated category
Fromont, 1988: 118.
of subectosomal megasclere. Hallmann (1912)
debated the value of ectosomal specialisation as
TYPE SPECIES. Spongia crat it ia Esper, 1797: 195 (by a generic character, concluding that although the
original designation) (fragment of type ZMB4577).
distinction between the two genera may eventualArborescent growth form. Surface microscopi- ly breakdown, they can be consistently differencally hispid, conulose. Choanosomal skeleton ir- tiated on composition of peripheral skeleton
regularly reticulate, with heavy spongin fibres rather than its development or density. These
cored by multispicular tracts of choanosomal conclusions are supported here.
Another character predominant in Thalysias
principal subtylostyles which terminate in
plumose tracts on peripheral fibres. Plumose mul- (including Rhaphidophlus) is the presence of
tispicular tracts of echinating acanthostyles more than one size category of isochelae, of
clumped around fibre nodes and also protruding which one or more may be contort (Fig. 5F). This
from fibres at regular intervals. Subectosomal feature is not consistent among species, nor is it
skeleton plumose, with well-developed multi- exclusive to the genus. Contort chelae are comspicular columns of subectosomal auxiliary sub- mon in Clathria (Thalysias) (e.g., Spongia
tylostyles arising from ends of principal spicule abietina Lamarck, Spongia cactiforinis Lamarck,
brushes in peripheral skeleton. Ectosome with Rhaphidophlus cervicornis Thiele, R. spiculosus
bundles of erect ectosomal auxiliary subtylos- Dendy, R. topsenti Thiele, Clathria fasciculatus
tyles overlaying subectosomal plumose brushes Wilson and C. spiculosus var. macilenta
of spicules. Megascleres entirely smooth hastate Hentschel), Clathria (Clathria) and Clathria
choanosomal principal subtylostyles, larger (Microciona) (e.g., Dictyociona adioristica de
smooth subectosomal auxiliary subtylostyles, Laubenfels, Clathria mixta Hentschel, C. bulshorter thinner smooth ectosomal auxiliary sub- botoxa Van Soest, Fisherispongia ferrea de
tylostyles, and echinating acanthostyles with Laubenfels, Esperiopsis obliqua George & Wilaspinose necks. Microscleres palmate isochelae son, Wilsonella conectens Hallmann, and M.
of two size categories, and two forms of toxas prolifera (sensu Wilson)), Clathria (Wilsonella)
(small wing-shaped and larger asymmetrical (e.g., Microciona tuberosa Bowerbank,
Clathriopsanuna ^reticulata Lendenfeld),
sinuous toxas).
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Clathria (Axociella) (e.g., Esperiopsis
canaliculata Whitelegge), Antho (Antho) (e.g.,
Holoplocamia penneyi de Laubenfels) and
Holopsamma (e.g., Plectispa macropora Lendenfeld). It is also known to occur in Cornulum
Carter (Iophonidae) (e.g., C. johnsoni (de
Laubenfels)) and consequently it cannot be given
much diagnostic value above the species level.
There are several other characters used by
authors to define genera present in many species
of Thalysias (including Rhaphidophlus), but are
not apomorphies occurring throughout related
groups in all combinations (e.g., encrusting
growth form and hymedesmoid architecture (e.g.,
Leptoclathria lambda Levi); absence of echinating megascleres (e.g., Axociella arteria de
Laubenfels), absence of microscleres (e.g.,
Clathria fascicularis Topsent), and modified
isochelae (e.g., Colloclathria ramosa Dendy)). A
systematics based on these features conflicts with
one based on ectosomal specialisation and dispersal of structural megascleres within the skeleton.
By comparison, Dendy (1905), Wilson (1925),
de Laubenfels (1936a), Hartman (1955), Wells et
al. (1960), Bergquist (1965), Hooper (1990a),
Hooper et al. (1990, 1992) and Hooper & Levi
(1993a) amalgamated Thalysias (including
Rhaphidophlus) and Clathria (including
Microciona) on the basis that dermal specialisation can vary intraspecifically, especially in relation to growth form and age of an individual.
These authors suggest that this feature is probably
not a sound base on which to separate genera. In
contrast, Simpson (1968a), Wiedenmayer (1977)
and Bergquist & Fromont (1988) maintain these
genera separately, although they do state that
dermal specialisation may not be important at the
generic level (i.e., they offer a convenient classification rather than one based on phylogeny).
Simpson (1968a) showed that despite very close
morphological similarities between Clathria and
Rhaphidophlus, there were cytological differences between the taxa which he considered were
sufficient to separate them at the generic level of
classification. Thus, Simpson's (1968a) cytological data supports the conclusion above based on
morphological evidence that the only morphological character of consistent diagnostic importance, and which correlates to some extent
with supposed biological differences between
those genera, is the possession of ectosomal
specialisation. This conclusion is upheld in this
study and used at the subgeneric level.
There are several nomenclatural complexities
that still exist for Rhaphidophlus, and these require further discussion.
1) Van Soest (1984b: 91) argued that Thalysias
should be abandoned in favour of Rhaphidophlus,
following Levi (1960a: 52). His argument was
based on the premise that Carter (1876: 311)
designated the 'representative' of Thalysias as
Thalysias subtriangularis (Duchassaing, 1850),
and he suggested that although Carter (1876) did
not actually use the words 'type' or 'type species',
there was no doubt of his intentions (T subtriangularis is a species of Xestospongia; Wiedenmayer, 1977: 255; Van Soest et al., 1983: 199).
Van Soest (1984b) also suggested that the use of
Rhaphidophlus over Thalysias was a pragmatic
solution since the name was in current usage by
the majority of contemporary authors. Conversely, Wiedenmayer (1977: 140), Hooper (1990a),
Hooper et al. (1990, 1992) and Hooper & Levi
(1993a) note that Carter (1876) merely compared
three small specimens from the North Atlantic
with Thalysias, but left them unnamed. Wiedenmayer (1977) states that although Carter did cite
Duchassaing's (1850: pl. 17, fig. 1) figure of
Thalysias subtriangularis, Carter's action cannot
be construed as a subsequent designation of a type
species under Articles 67c and 69a of the ICZN
(Anonymous, 1985). Therefore, de Laubenfels'
(1936a: 104) subsequent designation of Thalysias
virgultosa (Lamarck, 1814) as type species of
Rhaphidophlus is valid, and the genus is a junior
synonym of Thalysias.
2) Simpson (1968a: 98) suggested that arguments supporting or refuting the choice of one
genus name over another were irrelevant because
Rhaphidophlus,Thalysias and Axocielita showed
different cytological features, apparently justifying their recognition at the generic level. There
are no arguments based on skeletal evidence
which can contend with Simpson's hypothesis
since his cytological data do not correlate with
any skeletal features. However, Simpson's
(1968a) conclusions are based on only three
species, one in each genus, and it could be interpreted that the differences he observed may be
applicable only at the species level.
Simpson (1968a) found that Thalysias was distinct from both Microciona (cf. Hartman, 1955;
Wells et al., 1960), Rhaphidophlus and Clathria
(cf. Levi, 1960a). Thalysias was related to
Rhaphidophlus, but cytologically distinct, which
he stated was also apparently reflected in ectosomal cytological and morphological differences between the two genera. Rhaphidophlus
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU
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75
(sensu Simpson) had a specialised and extensive
region containing a secondary fibre system and a
continuous ectosomal skeleton, whereas
Thalysias had fibre cell tracts organised into dermal columns which produce tufts of ectosomal
styles. However, no similar correlations were observed in any other species examined (Van Soest,
1984b; Hooper, 1990a). Simpson's (1968a) system is intrinsically unworkable as it presently
stands, given that the existing systematics is
based on skeletal attributes, and essentially
Thalysias and Rhaphidophlus differ only in their
nomenclature.
3) Simpson (1968a) suggested that Axocielita
was distinct from, but most closely related to
Thalysias, but this conclusion is not supported by
their morphology. Axocielita similis (Stephens)
has a hymedesmoid skeleton with plumose fibre
nodes, each node cored by plumose spicule tracts,
echinating acanthostyles are absent and only one
size class of auxiliary spicule is present, suggesting that the species is more closely related to
Microciona than Thalysias (Hooper, 1990a).
Simpson (1968a: 113) also agreed that echinating acanthostyles are of minor diagnostic importance. The implication of these data is that
Axocielita harttnani Simpson and A. simi/is
(Stephens) are either not cogeneric, which is not
indicated by their morphological characters, or
that morphological systematics is not corroborated by cell biology, and this is the main
obstacle in using Simpson's results.
4) Simpson (1968a) also suggested that
Clathria was more closely related to
Rhaphidophlus and Thalysias in its cytological
characteristics, than it was to the morphologically
more-similar Microciona. This result conflicted
with the system proposed by Levi (1960a), and
developed further by Van Soest (1984b), which
distinguished microcionid genera on the basis of
their skeletal architecture and ectosomal characteristics, respectively. Nevertheless, it suggests
that thinly encrusting microcionid sponges may
have different cytological characteristics than
ramose or massive forms, and this poses questions concerning environmental influences on
cellular behaviour.
5) Hallmann (1920: 769) preferred Tenacia
Schmidt (1870) to Rhaphidophlus Ehlers (1870)
because t Tenacia had been firmly established by
the redescription of T. clathrata Schmidt (Wilson,
1902: 397), whereas Rhaphidophlus was imperfectly known only from Ehlers' (1870: 18, 31)
inadequate redescription of Spongia cratitia
Esper. The argument is irrelevant, since S. cratitia
is recognisable, and Thiele (1903a: 957) has already redescribed portions of the type material
(ZMB4577, 4578), even though the whereabouts
of the actual type specimen is presently unknown.
Hallmann (1920) also suggested that Tenacia
should be preferred to Rhaphidophlus because
Schmidt's publication was abstracted in the
Zoological Record for 1870, whereas Ehlers
work was abstracted in 1872. There is no doubt
that the genera are synonymous (Hartman, 1955:
176; Levi, 1960a: 56; Wiedenmayer, 1977: 140;
Van Soest, 1984b: 91) but Rhaphidophlus has
seniority over Tenacia, and Thalysias has
seniority over both. It is included in Clathria
(Thalysias).
Seriatula Gray, 1867
Seriatula Gray, 1867: 515; de Laubenfels, 1936a: 122.
(Refer to Ophlitaspongia).
Sophax Gray, 1867
(Fig. 23D-E)
Sophax Gray, 1867: 521; de Laubenfels, 1936a: 112.
TYPE SPECIES. Microciona fallax Bowerbank, 1866:
128 (by monotypy) (lectotype BMNH1910.1.1.71;
paralectotype BMNH1930.7.3.198; fragment of lectotype USNM5047).
Encrusting growth form. Surface uneven,
microscopically hispid. Choanosomal skeleton
plumose (slightly plumoreticulate), with spongin
fibres reduced to basal layer of spongin on substrate with ascending spongin fibre nodes cored
by multispicular columns of choanosomal principal subtylostyles, and echinated by acanthostyles. Ascending fibres interconnected by sparse
transverse spongin fibres, aspiculose or with few
coring principal spicules. Ectosomal skeleton
with tangential, paratangential or poorly
developed brushes of subectosomal auxiliary
subtylostyles. Megascleres long choanosomal
principal subtylostyles with heavily microspined
bases, entirely smooth, flexuous, sinuous or
straight subectosomal auxiliary subtylostyles,
and echinating acanthostyles of vaiiable size and
evenly distributed spines. Microscleres absent.
REMARKS. Gray (1867) erected Sophax for M.
fallax, without further comment or comparison
with other Microciona species. The type species
has a Microciona-like hymedesmoid skeleton
with erect spongin fibre nodes (similar to other
nominal genera such as Abila, Axosuberites,
Stylotellopsis, and Thalyseurypon), but unlike
these it also has some transverse fibres intercon-
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76zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 23. Type species of microcionid genera. A-C, Rhaphidophlus (Spongia cratitia Esper, ZMB4577). D-E,
Sophax (Microciona fallax Bowerbank, BMNH1910.1.1.71). F-G, Stylotellopsis (S. amabilis Thiele,
ZMB3309). H-I, Tablis (Echinochalina anomala Hallmann, AMG10548).
77
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
necting multispicular tracts, producing a slightly
plumoreticulate architecture. Sophax is referred
here to Clathria (Microciona).
Stylotellopsis Thiele, 1905
(Fig. 23F-G)
species of Clathria (Thalysias). De Laubenfels
(1936a) included the genus with Eurypon
(Raspailiidae), but this is not upheld here:
similarities between Stylotellopsis and Thalysias
in their peripheral skeletal structure and the
geometry of auxiliary spicules are closer than
with the raspailiids (see Hooper, 1991).
Stylotellopsis Thiele, 1905: 456; de Laubenfels, 1936a:
112.
Tablis de Laubenfels, 1936
TYPE SPECIES. Stylotellopsis zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
am abilis Thiele, 1905:
456 (by monotypy) (holotype ZMB3309).
(Fig. 23H-I)
Thickly encrusting growth form. Surface
hispid, uneven. Choanosomal skeleton hymedesmoid, with basal layer of spongin fibres lying on
substrate and plumose non-anastomosing fibre
nodes. Spongin fibre nodes cored by large
echinating acanthostyles standing perpendicular
to substrate, forming multispicular plumose tufts
on basal skeleton. Subectosomal skeleton
plumose, with multispicular columns of subectosomal auxiliary subtylostyles arising from ends
of echinating spicule brushes, producing
dendritic whispy tracts especially near periphery.
Ectosomal skeleton without special spiculation
but erect subectosomal auxiliary styles form
nearly continuous palisade on surface. Mesohyl
has heavy deposits of loose spongin. Megascleres
single category of smooth subectsomal auxiliary
styles with tapering bases and hastate points, and
large echinating acanthostyles with swollen
bases, large spines, and aspinose points. Principal
spicules undifferentiated from auxiliary spicules.
Microscleres absent.
REMARKS. From published descriptions
(Thiele, 1905; Topsent, 1928a; Koltun, 1964a)
Stylotellopsis appears to be similar to Thalysias
in having differentiated subectosomal and ectosomal auxiliary spicules (i.e., a specialised ectosomal skeleton), and Van Soest (1984b)
synonymised the two genera on this basis. Re-examination of type material found that auxiliary
spicules range greatly in their length, and principal spicules are very similar to auxiliary
spicules scattered throughout the skeleton. The
genus has a `microcionid' choanosomal skeletal
structure (i.e., hymedesmoid with plumose fibre
nodes), and an unusual subectosomal skeleton
composed of dendritic or sinuous auxiliary
spicule tracts. It also lacks microscleres (although
several arcuate isochelae were seen in histological preparations of the holotype, but these are
probably foreign). Stylotellopsis is interpreted
here as being an encrusting (hymedesmoid)
Tab/is de Laubenfels, 1936a: 76.
TYPE SPECIES. Echinochalina anom ala Hallmann,
1912: 292 (by original designation) (holotype
AMG10548).
Massive, lobate-digitate growth form. Surface
'honeycomb' reticulate. Choanosomal skeleton
reticulate, with heavy primary and secondary
spongin fibres forming wide-meshed reticulation
cored by multispicular tracts of subectosomal
auxiliary subtylostyles. Coring spicules in
primary fibres compressed, more dispersed in
secondary fibres. Fibres heavily echinated by
principal styles. Subectosomal skeleton with
scattered auxiliary megascleres dispersed between fibre meshes and protruding through surface in paratangential bundles. Megascleres
entirely smooth principal styles-subtylostyles,
and smooth subectosomal auxiliary subtylostyles, often secondarily modified to tornotes.
Microscleres absent.
REMARKS. Tablis was erected for Whitelegge's
(1907) specimen of Echinochalina glabra from
Woolongong, NSW, a junior homonym of Ridley
& Dendy's (1887) species renamed by Hallmann
(1912) to E. anomala. This species has also been
recorded from the Great Barrier Reef (Burton,
1934a). Three other species were referred to
Tab/is by de Laubenfels (1936a): Echinochalina
glabra Ridley & Dendy (a synonym of Spongia
barba Lamarck), E. reticulata Whitelegge, both
from SW Australia, and Spongia clavosa
Lamarck. The last mentioned species is unknown: it was not mentioned in Topsent's (1933)
revision of Lamarck's species nor found in the
Lamarck collections (MNHN). These other 2
species differ from E. anomala in having
microscleres but lacking an Holopsamma-like
'honeycombed reticulate' growth form. Tab/is is
returned here to Echinochalina (Echinochalina).
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Tenacia Schmidt, 1870
(Fig. 24A-C)
TYPE SPECIES. Esperiopsis ^canaliculata
Whitelegge, 1906: 471 (by monotypy) (lectotype
AMG4325).
Tenacia Schmidt, 1870: 56; Hallmann, 1920: 769;
Topsent, 1920b: 17; Burton & Rao, 1932:337; Levi,
1960a: 56.
Erect, branching, digitate growth form. Surface even, hispid, with oscules dispersed over
lateral margins of branches. Choanosomal
skeleton irregularly reticulate, nearly radial, with
compressed axial and plumose extra-axial
skeletons. In axis heavy spongin fibres cored by
multispicular tracts of choanosomal principal
styles, and in extra-axial skeleton principal styles
protrude from peripheral fibres forming radial or
plumose tracts, lying immediately below and supporting subectosomal skeleton. Echinating
megascleres absent. Subectosomal skeleton
plumose, with multispicular ascending tracts of
subectosomal auxiliary styles at ends of
peripheral choanosomal fibres. Ectosomal
skeleton with thick continuous palisade of
smaller ectosomal auxiliary styles standing perpendicular to surface. Megascleres smooth
fusiform choanosomal principal styles, subectosomal auxiliary styles-subtylostyles with
smooth or microspined bases, and ectosomal
auxiliary styles-subtylostyles with microspined
bases. Microscleres palmate isochelae of two
sizes, including contorted forms, and oxhorn and
wing-shaped toxas.
TYPE SPECIES. Tenacia clathrata Schmidt, 1870: 56
(by monotypy) (fragment of type BMNH1870.5.3.156); = Spongia virgultosa Lamarck, 1814: 444
(fragment of type MNHNLBIMDNBE1344, 1338).
Erect, arborescent, reticulate branching growth
form. Surface highly conulose, uneven, microscopically hispid. Choanosomal skeleton irregularly reticulate, with well developed spongin
fibres differentiated into ascending primary and
transverse secondary fibres. Principal spicules
confined to axis of primary fibres, but absent
completely from secondary connecting fibres,
and all fibres cored by paucispicular tracts of
subectosomal auxiliary subtylostyles, and heavily echinated by small acanthostyles. Subectosomal skeleton plumose, with tracts of
subectosomal auxiliary subtylostyles originating
from deeper regions of choanosomal skeleton;
subectosomal auxiliary subtylostyles also scattered abundantly throughout mesohyl between
fibre meshes. Ectosome plumose, with brushes of
smaller ectosomal auxiliary subtylostyles arising
from ends of subdermal spicule brushes.
Megascleres smooth choanosomal principal
styles, smooth subectosomal auxiliary subtylostyles, smooth ectosomal auxiliary subtylostyles,
and short thick echinating acanthostyles with
aspinose points and necks. Microscleres palmate
isochelae of two sizes, and wing-shaped, accolada and sinuous asymmetrical toxas.
REMARKS. Tenacia and Rhaphidophlus are
synonyms (Hallmann, 1920; Topsent, 1932; Levi,
1960a), and use of one name over another is
merely a nomenclatural decision (see remarks for
Rhaphidophlus). Topsent's (1932: 97) synonymy
of T clathrata and Spongia juniperina Lamarck
is not upheld here: the former is considered here
to be a synonym of T virgultosa and restricted to
Caribbean and NE. Atlantic populations, whereas
T. juniperina (including T clathrata of Hallmann,
1912) is known only from the Indian Ocean. The
genus is synonymised with Clathria (Thalysias).
Tenaciella Hallmann, 1920
(Fig. 24D-E)
Tenaciella Hallmann, 1920: 772; de Laubenfels,
1936a: 126.
REMARKS. Hallmann (1920) erected Tenaciella
for species like Tenacia(= Thalysias), but lacking
echinating megascleres. Wells et al. (1960) also
referred Esperiopsis obliqua George & Wilson to
Tenaciella, but this species lacks dermal
specialisation and is more closely allied with
Axocielita (= Clathria). Ristau (1978) suggested
that Artemisina archegona Ristau (USNM
24528) was similar to the type species of
Tenaciella in having prominent subectosomal
(extra-axial) spicule columns, and he suggested
that the two species differed only in growth form,
shape of the toxa microscleres and the appreciably more organised skeletal architecture in T
canaliculata. However, differences observed in
types of these species are more noticable than
their similarities (e.g., ectosomal skeleton, fibre
reticulation, degree of axial and extra-axial differentiation), and their supposed affinities (Ristau, 1978), are superficial.
In its skeletal structure, differentiation of axial
and extra-axial skeletons, fibre characteristics,
the possession of similar extra-axial radial
bundles of megascleres, and having a continuous
ectosomal palisade of spicules the type species
shows close similarities to Ceratopsion axifera
(Hentschel) (Raspailiidae) from the Arafura Sea
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
79
(Hooper, 1991). In fact the resemblance between
these two species is remarkable (although
Esperiopsis canaliculata obviously belongs to
Microcionidae, having isochelae and toxas,
whereas Ceratopsion belongs to Raspailiidae in
having long extra-axial spicules surrounded by
bundles of ectosomal spicules and lacking chela
or toxa microscleres). These structural
similarities suggest a closer relationship between
the two families, as proposed by Hooper (1991)
in returning Raspailiidae to the Poecilosclerida,
and subsequently supported by chemotaxonomic
evidence (Hooper et al., 1992).
In skeletal architecture E. canaliculata is identical to E. cylindrica, the type species of Axociella. Consequently, both Tenaciella and Axociella
are synonymised here (the latter being the most
senior available name), both having a compressed
reticulate axis and plumose or plumo-reticulate
extra-axial (subectosomal) skeletons, isochelae
and toxa microscleres, lacking echinating
spicules, and referred to Clathria (Axociella).
Thalassodendron Lendenfeld, 1888
Thalassodendron Lendenfeld, 1888: 222; de Laubenfels, 1936a: 112.
TYPE SPECIES. Thalassodendron typica Lendenfeld,
1888: 223 (by indication) (holotype unknown).
Flabellate, cup-shaped growth form. Surface
striated longitudinally. Choanosomal skeleton
reticulate, with differentiated primary ascending
and secondary transverse connecting spongin
fibres; primary fibres cored by multispicular
tracts of choanosomal principal styles, lightly
echinated by acanthostyles (often secondarily incorporated into fibres), whereas secondary fibres
aspiculose. Subectosomal and ectosomal
skeletons unknown. Megascleres include smooth
choanosomal principal styles, and short stout
echinating acanthostyles. Microscleres unknown.
REMARKS. This diagnosis is based on
Lendenfeld's (1888) description of the type
species which is virtually unrecognisable (de
Laubenfels, 1936a). It is suspected that Thalassodendron typica Lendenfeld is a synonym of
Echinonema typicum Carter, given Lendenfeld's
propensity for describing other authors' species
as his own 'new species'. However, it is not
possible to associate any type materialwith the
name `Thalassodendron typicum' and this
synonymy remains doubtful. Echinonema
typicum is also a synonym of Spongia cactifortnis
Lamarck, in which case Thalassodendron would
belong to Clathria (Thalysias) . Thalassodendron
typica of Whitelegge (1901: 86; holotype
AMZ958) is not the same as Lendenfeld's
species, being a synonym of Echinodictyum
mesenterinum (Lamarck) (Raspailiidae)
(Hooper, 1991: 1379).
Thalyseurypon de Laubenfels, 1936
(Fig. 24F-G)
Thalyseurypon de Laubenfels, 1936a: 107; Wiedenmayer, 1977: 143.
TYPE SPECIES. Spongia raphanus Lamarck, 1814:
444 (by original designation) (holotype
MNHNLBIMDT572).
Arborescent, reticulate, bushy growth form.
Surface conulose, not hispid. Choanosomal
skeleton more-or-less regularly reticulate, with
heavy spongin fibres undifferentiated into
primary or secondary components. Choanosomal
fibres mostly aspiculose, or with irregularly
paucispicular tracts of choanosomal principal
subtylostyles, sparsely echinated by acanthostyles. Subectosomal skeleton poorly developed,
consisting only of extra-fibre paratangential
tracts of subectosomal auxiliary subtylostyles,
becoming tangential in ectosomal region.
Megascleres entirely smooth fusiform choanosomal principal subtylostyles, smooth subectosomal auxiliary subtylostyles, and small
vestigially spined acanthostyles. Microscleres
absent.
REMARKS. Thalyseurypon was established for
8 species differing from Clathria only in lacking
microscleres. Van Soest (1984b) proposed to
merge the genus with Clathria, whereas Wiedenmayer (1977) synonymised it with Pandaros suggesting that the skeletal architecture and absence
of microscleres in Spongia raphanus (sensu Topsent, 1932: 100) was similar to P. acanthifolium
Duch. & Mich. However, re-examination of type
material in both type specimens indicates that S.
raphanus is most appropriately placed in
Clathria (Clathria).
Thalysias Duchassaing & Michelotti, 1864
(Figs 24A-C, 25A-B, 176-177)
1) Thalysias Duchassaing & Michelotti, 1864: 82; de
Laubenfels, 1936a: 104; Hartman, 1955: 172;
Wiedenmayer, 1977: 140.
2) Thalysias; Carter, 1876: 311; de Laubenfels, 1954:
137; Levi, 1960a: 52; Simpson, 1968a: 98.
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TYPE SPECIES. 1) Interpretation by Wiedenmayer
(1977), (i.e., Thalysias s.s.): Spongia virgultosa
Lamarck, 1814 (by subsequent designation; de
Laubenfels, 1936a: 104) (holotype MNHN missing,
fragment BMNH1954.2.20.67); junior synonym of
Spongia juniperina Lamarck, 1814 (lectotype
MNHNLBIMDT570) (de Laubenfels, 1936a: 104). 2)
Interpretation by Van Soest (1984b), i.e., Thalysias of
authors: Thalysias subtriangularis Duchassaing, 1850
(by subsequent designation; Carter, 1876: 311); = Xestospongia subtriangularis (Wiedenmayer, 1977: 255).
DEFINITION OF TYPE SPECIES. As for
Tenacia.
REMARKS. The nomenclatural complexities of
Thalysias have been discussed above (see
Rhaphidophlus and Tenacia). According to Van
Soest (1984b), Carter's (1876) statement that T
subtriangularis was 'representative' of the genus
means that Thalysias sensu Carter is a haplosclerid, but this is not a valid subsequent designation. Wiedenmayer (1977) stated that de
Laubenfels' (1936a) subsequent designation of T.
virgultosa as the type, which makes Thalysias
(established 1864) a senior synonym of
Rhaphidophlus (established 1870). Moreover,
Tenacia is an objective synonym of Thalysias by
synonymy of their respective type species.
This has been confirmed by type material (although the holotype of Spongia virgultosa is only
represented by a fragment in the BMNH).
Thalysias is used here as the earliest available
subspecific name for Clathria with differentiated
ectosomal and subectosomal spiculation (i.e.,
specialised ectosomal skeleton).
Wetmoreus de Laubenfels, 1936
Wetmoreus de Laubenfels, 1936a: 112.
TYPE SPECIES. Microciona novaezealandica
Brondsted, 1924: 463 (by original designation)
(holotype UZM not found).
Encrusting growth form. Surface shaggy,
hispid. Choanosomal skeleton hymedesmoid,
with spongin fibres reduced to basal layer of
spongin on substrate with plumose non-anastomosing fibre nodes. Fibre nodes cored by erect
multispicular bundles of choanosomal principal
subtylostyles forming ascending plumose skeletal columns, and echinating acanthostyles at oblique angles to skeletal columns, usually forming
brushes. Subectosomal skeleton paratangential or
plumose, with single category of subectosomal
auxiliary subtylostyles forming light dermal
brushes erect on surface, or scattered individually
on surface and throughout mesohyl. Megascleres
choanosomal principal subtylostyles with
prominent basal spines, smooth subectosomal
auxiliary subtylostyles-styles, and small evenly
spined echinating acanthostyles. Microscleres allegedly include both palmate and arcuate forms.
Toxas absent.
REMARKS. This definition is from Brondsted's
(1924) description of the type.
The holotype is possibly extant in Brondsted's
collection at the UZM, but has not been located
(0. Tendal, pers. comm.). Wetmoreus differs from
Microciona (s.s.) in allegedly having both palmate and arcuate isochelae, and lacking toxas
(see remarks for Paradoryx), but this cannot be
verified. It is possible that both forms of chelae
are merely modified (curved, thickened) palmate
forms. Wetmoreus is included here in Microciona
based on the possession of a hymedesmoid basal
skeleton with plumose fibre nodes.
Wilsonella Carter, 1885
(Fig. 24H-I)
Wilsonella Carter, I885f: 366; Hallmann, 1912: 237,
1920: 768; Topsent, 1928a: 62; de Laubenfels,
1936a: 109.
TYPE SPECIES. Wilsonella australiensis Carter,
1885f: 366 (by monotypy) (holotype
BMNH1886.12.15.43).
Erect, massive or flabelliform growth forms.
Surface prominently conulose at apex of sponge,
with conspicuously raised oscules. Choanosomal
skeleton reticulate, with moderately light spongin
fibres forming irregular meshes with distinct
primary ascending and secondary transverse
lines. Primary fibres cored by paucispicular tracts
of robust auxiliary subtylostyles and abundant
detritus, and heavily echinating by acanthostyles
particularly at fibre nodes. Secondary connecting
fibres with paucispicular tracts, little detritus and
lightly echinated. Ectosomal skeleton lightly
arenaceous, mostly with spicule fragments, lacking specialised spiculation but with light tangential or paratangential tracts of more slender
subectosomal auxiliary subtylostyles.
Megascleres fusiform robust auxiliary subtylostyles with smooth bases and hastate or telescoped
points (inside fibres), more slender auxiliary subtylostyles with spines on both bases and points
(outside fibres), and echinating acanthostyles
with even spination. Microscleres palmate
isochelae and wing-shaped toxas.
REMARKS. Wilsonella is characterised by
auxiliary megascleres coring fibres (i.e., without
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
^
81
true choanosomal principal spicules), incorporation of sand and foreign spicule detritus into the
skeleton, and lacking any differentiation between
ectosomal and subectosomal skeletons. Wilsonella australiensis is unusual amongst 'sandy'
microcionid sponges in that subectosomal
auxiliary styles outside fibres (e.g., in the ectosomal skeleton) are geometrically very similar
to those coring fibres differing only in having
spines on both their bases and points. These
spicules are characteristic of the type species
(although they are also known in several other
microcionids, such as Clathria (C.) chelifera and
Clathria (T) major). Even though spicules coring
the fibres cannot be construed as being true principal megascleres, it must be concluded that this
species has two distinct categories of auxiliary
spicules. In this respect it is contrasted with
Clathria (Dendrocia), which has a completely
plumose skeletal architecture, including ectosomal and subectosomal differentiation, but
only a single undifferentiated category of structural megascleres throughout the skeleton.
Species referred to Paradoryx by Hallmann
(1920) are similar to Wilsonella in having only
auxiliary megascleres in the skeleton, but these
have arcuate-like chelae instead of palmate
isochelae, more than one category of structural
megasclere (i.e., Clathria (Clathria)), or only 1
category of auxiliary spicules and a plumose architecture (i.e., Clathria (Dendrocia)). In contrast, the type species of Wilsonella incorporates
detritus into spongin fibres, and this character is
interpreted as probably being phylogenetic rather
than merely ecological despite being
homoplasious througout Porifera (known also in
Phorospongiidae, Raspailiidae, Ircinidae,
Dysideidae, etc.), because it is correlated with
distribution of megascleres within skeletal
regions as described above. Contemporary
authors (e.g., Levi, 1967b) have included Wilsonella as a synonym of Clathria, whereas in the
present work the type species of Wilsonella is
synonymised with Clathriopsamnza (both with
detritus in the skeleton), the former being the
senior name, and used as at the subgenus level,
Clathria (Wilsonella).
GENERA EXCLUDED FROM
MICROCIONIDAE
Acarnus Gray, 1867
Acarnus Gray, 1867: 544; Hooper, 1987: 71; Hiemstra
& Hooper, 1991: 434; Van Soest et al., 1991: 49;
Hooper & Levi, 1993a: 1222 (for full synonymy see
Hooper, 1987).
TYPE SPECIES. Acarnus innominatus Gray, 1867:
544 (by monotypy) (holotype BMNH not found).
Thickly encrusting, massive, flabellate or
vasiform growth forms. Surface smooth or
minutely hispid, uneven, often sculptured with
subdermal canals in encrusting forms.
Choanosomal skeleton with short plumose tracts
connected by renieroid reticulation of spongin
fibres, reduced to plumo-reticulate skeletal tracts,
or further reduced to plumose-halichondroid
skeleton in encrusting forms. Skeletal tracts composed of uni- or paucispicular tracts of smooth
choanosomal principal styles or subtylostyles,
sometimes with spined bases. Fibres usually
heavily invested with spongin, with granular collagenous around nodes of skeletal tracts, and
fibres echinated by smooth and/or spined
cladotylotes of one or two size categories, with or
without additional, smaller echinating acanthostyles. Ectosomal skeleton with a tangential or
paratangential layer of basally spined tylotes.
Microscleres palmate isochelae and up to three
distinct forms of toxas: oxhom, wing-shaped and
accolada toxas.
REMARKS. The type species concept follows
Van Soest (1984b: 61) based on material from the
Caribbean given that the holotype is missing from
the BMNH. The type has two categories of
cladotylote megascleres, a larger smooth and
smaller spined variety, whereas other species are
known to have various combinations of those
spicules (Van Soest et al., 1991), together with
acanthostyles in some species (A. (Acanthacarnus) Levi).
Although the genus has now been exhaustively
discussed (Hooper, 1987; Hiemstra & Hooper,
1991; Van Soest et al., 1991), its family placement
is still debatable. In its microsclere complement
(palmate isochelae, diverse toxas), differentiated
principal and auxiliary spicules, echinating acanthostyles in some species (as well as having a
highly modified second category of echinating
spicules (cladotylotes) which are unique to the
genus, derived from either acanthostyles
(Hooper, 1987) or ectosomal tylotes (Hiemstra &
Hooper, 1991)), the genus appears to have affinities with the Microcionidae (e.g., Burton,
1959; Levi, 1960a; Vacelet et al., 1976). Other
authors (de Laubenfels, 1936a; Tanita, 1963;
Hechtel, 1965; Bakus, 1966; Thomas, 1970a,
1973; Hoshino, 1981; Van Soest, 1984b; Hooper,
1987; Hiemstra & Hooper, 1991; Van Soest et al.,
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FIG. 24. Type species of microcionid genera. A-C, Tenacia (Spongia juniperina Lamarck, MNHNDT570). D-E,
Tenaciella (Esperiopsis canaliculata Whitelegge, AMG4325). F-G, Thalyseurypon (Spongia raphanus
Lamarck, MNHNDT572). H-I, Wilsonella (W. australiensis Carter, BMNH1886.12.15.43).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP
^
83
1991; Hooper & Levi, 1993a) suggest that true
diactinal ectosomal spicules (tylotes with terminal spines and swollen tips) and a renieroid
reticulation indicate relationship to Myxillidae
(in the sense of Hartman, 1982). Hajdu et al.
(1994) resurrected Iophonidae for Acarnus,
Megaciella, Melonchela and others with
microcionid-like spiculation (terminally spined
megascleres of diverse categories, palmate
isochelae, toxas) as well as ectosomal tylotes
(previously considered diagnostic for the Myxillidae; Van Soest, 1984b). This revised interpretation de-emphasises the primary importance
placed on skeletal structure (Bergquist &
Fromont, 1988) for example), and allows for the
inclusion of renieroid reticulate skeletons in
several families (Iophonidae, Microcionidae,
Raspailiidae, Phoriospongiidae, Cornuliidae,
etc.). Skeletal structure would, therefore, be a
highly homoplasious character but this interpretation does allow a consistent differentiation
between Microcionidae and Iophonidae based on
monactinal versus diactinal-derived ectosomal
spicules, whilst also acknowledging their affinities based on the possession of similar
microscleres through their inclusion together in
Microcionina.
Amphilectus Vosmaer, 1880
Amphilectus Vosmaer, 1880: 109; Ridley & Dendy,
1887: 123; Burton, 1929a: 428; Levi, 1960a: 55.
TYPE SPECIES. Isodiciya gracilis Bowerbank, 1866:
149 (by subsequent designation of Burton, 1929a: 428)
(holotype BMNH1877.5.21.754).
Arborescent, dichotomously branched, stipitate
growth form. Surface even, hispid. Choanosomal
skeleton reticulate, with multispicular ascending
primary fibres and uni- or paucispicular
transverse connecting fibres, both cored by small
styles. Subectosomal region with tracts of
spicules projecting through surface. Ectosome
membraneous, without specialised spiculation.
Echinating megascleres absent. Megascleres
small smooth styles of a single category.
Microscleres palmate isochelae. Toxas absent.
REMARKS. Vosmaer (1880: 109) established
Amphdectus for a heterogeneous assemblage of
42 poecilosclerid species, most of which were
related to Mycale, Esperiopsis (Mycalidae), Desmacidon (Phoriospongiidae) or Myxilla (Myxillidae). Ridley & Dendy (1887) restricted the
genus to taxa with smooth styles and palmate
isochelae, but even so, they remarked that the
taxon was undoubtedly artificial. Burton (1929a)
designated Vosmaer's first-named species as
genotype, and suggested that in the strict sense
(i.e., the above diagnosis) the genus had affinities
with Esperiopsis. Levi (1960a) decided to abandon Amphilectus because, in the sense of Vosmaer
(1880), it was too vague and served only as a
catch-all taxon. In the broad sense Amphilectus
contains some microcionid species (e.g.,
Microciona annata Bowerbank) whereas in the
strict sense it fits with the concept of Myxillidae
(Bergquist & Fromont, 1988; Hajdu et al., 1994).
Caulospongia Kent, 1871
Caulospongia Kent, 1871: 616; Burton, 1930c: 673; de
Laubenfels, I 936a: 118.
Plectrodendron Lendenfeld, 1888: 66; Hallmann,
1914a: 306.
TYPE SPECIES. Caulospongia verticillata Kent,
1871: 616 (by subsequent designation of Hallmann,
1914a: 306) (holotype BMNH1895.7.16.1); =Spongia
perfoliata Lamarck, 1814: 439 (Topsent, 1932: 85)
(lectotype MNHNLBIMDT582).
Distinctive foliose growth form with lamellae
arranged in whorls, or in plates, around an erect
stalk. Surface even, minutely hispid. Choanosomal skeleton plumoreticulate, with well
developed spongin fibres cored by pauci- or multispicular tracts of choanosomal principal tylostyles; longitudinal primary fibres form dendritic
branches through axis, and ascending or oblique
secondary fibres produce a nearly regular secondary reticulation. Subectosomal skeleton plumose,
with erect brushes of choanosomal megascleres
protruding through ectosome. Ectosome with
tangential or paratangential crust of smaller tylostyles. Megascleres tylostyles of two sizes but with
same geometry. Microscleres absent.
REMARKS. Caulospongia has been included in
Suberitidae (Hadromerida) by most authors since
Burton (1930c). The type species is distinctive in
growth form, and although it is common in NW
Australian coastal waters it has been recorded in
the literature only infrequently (Lamarck, 1814;
Kent, 1871; Topsent, 1932; Hooper, 1984a). De
Laubenfels (1936a: 118) defined Caulospongia
in Ophlitaspongiidae presumably based on
similarities to ophlitaspongiids (most now included in Microcionidae) in skeletal architecture
(differentiated axial and extra-axial regions),
localisation of spicules to different parts of the
skeleton, and growth form (erect, whereas
'typical' suberitids are massive). However, these
similarities are convergent. The presence of true
tylostyles in Caulospongia (which are charac-
^
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84zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 25. Type species of microcionid genera. A-B, Thalysias (Spongia virgultosa Lamarck, BMNH1954.2.20.67).
teristic of suberitids, as opposed to subtylote
styles seen in many microcionids), and the absence of microscleres (which often provide useful
clues on phylogenetic affinities) suggest that the
genus should be retained in Suberitidae.
Plectrodendron (type species Plectrodendron
elegans Lendenfeld, 1888: 66, by monotypy) was
also referred to Caulospongia by Hallmann
(1914a: 306).
Megaciella Hallmann, 1920
(Fig. 19A-B)
Megaciella Hallmann, 1920: 772.
TYPE SPECIES. Amphilectus pilosus Ridley &
Dendy, 1886: 350 (by original designation) (lectotype
BMNH1887.5.2.125).
Lobate flabellate growth form. Surface shaggy,
ridged, hispid. Choanosomal skeleton reticulate,
with ascending multi- or paucispicular tracts of
choanosomal principal styles, interconnected by
secondary uni- or bispicular tracts within light
spongin, producing irregular wide meshes.
Echinating spicules absent. Subectosomal
skeleton radial or plumose, with erect
choanosomal principal styles protruding from
peripheral fibres through surface. Ectosome with
tangential or paratangential layer of tylotes, often
in bundles. Megascleres very large smooth
choanosomal principal styles, and ectosomal
tylotes, often curved or sinuous, with slightly
swollen microspined bases. Microscleres minute
palmate isochelae and two sizes of toxas (very
large accolada and wing-shaped).
REMARKS. This diagnosis is based on the lectotype which differs slightly from the original
description of Ridley & Dendy (1886, 1887).
Specifically, the so-called 'long thin centrally
curved oxeas' are very large accolada toxas with
slight central curvature and slightly reflexed hastate points, some of which exceed 100p,m long
(i.e., larger than the principal styles). Similarly,
these toxas are not associated with the ectosomal
skeleton but are scattered throughout the
mesohyl, whereas bundles of `oxeas' described
by Ridley & Dendy (1887) are actually bundles
of ectosomal tylotes. A second morph of toxas is
85
^
REVISION OF MICROCION1DAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
also present, being large and wing-shaped, and
these too have hastate points.
Hallmann (1920) suggested that a lax skeletal
construction and supposedly comparable spiculation indicated affinities between Megaciella and
Artemisina. He noted that Megaciella differed
from that genus in lacking ornamentation on
toxas, which he considered was a significant
diagnostic character at the generic level. However, in regard to this latter character, it has been
found that a number of microcionid species have
distally spined toxas (see remarks for Clathria),
and in any case, both genera have quite different
ectosomal skeletons. On the basis of its ectosomal
characteristics (consisting of basally spined diactinal tylote spicules), Van Soest (1984b) referred
Megaciella to the Myxillidae, whereas Artemisina (with a monactinal ectosomal skeleton)
was retained in Microcionidae. Under the revised
scheme proposed by Hajdu et al. (1994)
Megaciella is included in Iophonidae, with palmate isochelae and toxas which being the only
real synapomorphy between Megaciella and the
Microcionidae (i.e., Microcionina).
Melonchela Koltun, 1955
(Fig. 19C-E)
Melonchela Koltun, 1955a: 17, 1959: 187.
TYPE SPECIES. Melonchelaclathrata Koltun, 1955a:
17 (by original designation) (paratype BMNH1963.7.29.7).
Arborescent, reticulate planar, branching
growth form. Surface even, microscopically
hispid. Choanosomal skeleton plumose, with bundles
of independent ascending fibres cored by large
and small choanosomal principal styles. Ascending fibres not interconnected. Echinating acanthostyles absent. Subectosomal skeleton plumose,
with principal styles projecting through surface.
Ectosome with tylotes forming tangential layer or
erect brushes on surface. Megascleres large and
small choanosomal principal styles, with smooth
or microspined bases, and diactinal ectosomal
tylotes (swollen bases) and strongyles (rounded
bases) with microspined bases. Microscleres
abundant palmate cleistochelae, palmate
isochelae, small wing-shaped toxas, and oxhorn
toxas with spines, mucronate points, or telescoped points, or simply with subterminal ridge.
REMARKS. This species is remarkable in
several features: its erect planar reticulate growth
form (superficially resembling the microcion id
Clathria coppingeri and the raspaili id Echinodic-
tyum cancellatum); extremely large size range of
principal spicules, the largest protruding a long
way through fibre bundles, reminiscent of
Raspailiidae; the apparent lack of connecting
fibres between the ascending plumose spicule
tracts; a ridge-like subterminal ornamentation on
toxas; and extremely abundant tracts of chelae
microscleres throughout the mesohyl.
Diactinal ectosomal spicules (tylotes, strongyles and intermediates, varying in the degree of
swelling of their bases), palmate isochelae and
toxas indicates that the species has affinities to
Acarnus and Megaciella in the Iophonidae.
Naviculina Gray, 1867
(Fig. 19H-I)
Naviculina Gray, 1867:538; de Laubenfels, 1936a: 88.
TYPE SPECIES. Naviculina cliftoni Gray, 1867: 538
(by monotypy); for Ilymedesmia sp. nov.' of Bowerbank, 1864: 252 (fragment of type BMNH1877.5.21.270).
Growth form and surface details unknown. Ectosomal features unknown. Choanosomal
skeleton evenly reticulate, wide meshed, composed of tracts of subtylostyles bound together
with nodal spongin, with multispicular tracts
several spicules wide interconnected by uni- or
paucispicular tracts only one spicule wide, both
producing even triangular meshes. Megascleres
single category of subtylostyles. Microscleres
cleistochelate anisochelae, dispersed throughout
mesohyl particularly between fibre anastomoses.
REMARKS. Naviculina is monotypic, and the
type species, from Fremantle is characterised by
cleistochelae ('naviculoid spiculum' of Bowerbank, 1864), but nothing else was known about
the species. de Laubenfels (1936a) considered
Naviculina dubious, with alleged affinities to
Plocamiopsis (having cleistochelae), although
several other nominal genera are also known to
have cleistochelate microscleres, interpreted as
modified palmate isochelae (Colloclathria,
Plocamiopsis and Quizciona of the Microcionidae;
and Melonchela of the Iophonidae). The holotype
no longer exists. The holotype slide (Bowerbank,
1864; Gray, 1867) contains a small section of the
skeleton (Fig. 19H-I), but enough detail to indicate the Mycalidae, probably Arenochalina.
Paracornulum Hallmann, 1920
Paraconiulum Hallmann, 1920: 772.
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TYPE SPECIES. Cornu/um dubium Hentschel, 1912:
346 (by original designation) (holotype SMF964).
Encrusting growth form. Surface smooth, even.
Choanosomal skeleton hymedesmoid, with spongin fibres lying on substrate and bases of acanthostyles embedded and spicules standing
perpendicular to surface. Subectosomal skeleton
radial or plumose, with ascending tracts of erect
tornotes. Acanthostyles also scattered throughout
mesohyl. Ectosome with tangential tracts of tornotes. Megascleres radial tornotes (with fainly
rugose bases) and principal acanthostyles (with
some oxeote or strongylote modifications).
Microscleres palmate isochelae and oxhorn
toxas.
REMARKS. Paracornulum does not appear to be
as closely related to microcionids as inferred by
Hallmann (1920), showing similarities in possession of echinating acanthostyles, hymedesmoid
basal spongin fibres, palmate isochelae and toxas.
Two other species referred by Hallmann to this
genus (Cliona purpurea Hancock and Suberites
fuliginosus Carter) are very poorly known and
further comment on those taxa must await
redescription of relevant type material. Based on
its ectosomal characteristics and megasclere
spiculation Paracornulutn was subsequently
referred to Cornulidae (Levi & Levi, 1983a: 966).
The type species appears close to Cornulum
Carter, lophon Gray and Zyzzya de Laubenfels
because the ectosomal tornotes are reduced
tylotes with rugose (i.e., vestigially microspined)
bases. These genera are referred to Iophonidae
(Hajdu et al., 1994).
Paresperia Burton, 1930
Paresperia Burton, 1930a: 501.
TYPE SPECIES. Paresperia intermedia Burton,
1930a: 501 (by monotypy) (holotype BMNH1910.1.1.912).
Irregularly massive, low growth form. Surface
even, smooth. Choanosomal skeleton reticulate,
with loose, unispicular, irregular reticulation of
light spongin fibres cored by auxiliary subtylostyles, of same geometry as those in subectosomal
and ectosomal skeletons. Echinating acanthostyles absent. Subectosomal and ectosomal
skeletons lack specialised spiculation, but have
loose tangential reticulation of subectosomal
auxiliary megascleres. Megascleres only
auxiliary subtylostyles, entirely acanthose or
only basally spined. Microscleres palmate
isochelae. Toxas absent.
REMARKS. Burton (1930a) assigned
Paresperia to the Microcionidae based on supposed affinities to Artemisina (sensu Burton),
having an irregular, confused skeletal architecture composed of undifferentiated choanosomal
and ectosomal megascleres. The genus differed
from Artemisina (s.s.) in having lightly and evenly spined auxiliary megascleres and lacking
toxas. Burton also noted that Paresperia had affinities with the Mycalidae, whereas Van Soest &
Stone (1986) suggested that the presence of a
unispicular choanosomal reticulation of acanthostyles and palmate isochelae placed the genus
closer to Esperiopsis (placement still controversial in either Mycalidae or Esperiopsidae) than to
other microcionids.
Querciclona de Laubenfels, 1936
Querciclona de Laubenfels, 1936a: 46.
TYPE SPECIES. Antherochalina quercifolia Keller,
1889: 383 (by original designation) (holotype
ZMB429).
Erect, arborescent, flabellate growth form. Sur-
face even, hispid, regularly porous with oscules.
Choanosomal skeleton reticulate, with heavy
spongin fibres producing tight and irregular
meshes, fibres cored by choanosomal principal
styles which form primary ascending multispicular (eventually plumose) tracts, and secondary transverse uni- or paucispicular tracts,
together producing a subisodictyal reticulation.
Axial skeleton slightly different from extra-axial
region: near axis skeletal architecture isodictyal,
whereas towards periphery skeleton plumose.
Echinating megascleres absent. Subectosomal
skeleton plumose, with brushes of choanosomal
principal styles protruding through surface. Ectosomal skeleton with brushes of smaller
choanosomal styles surrounding one or few larger
subectosomal styles. Megascleres small smooth
choanosomal principal styles, and long smooth
subectosomal auxiliary styles. Microscleres absent.
REMARKS. Querciclona seems to be a case
where inadequate primary taxonomy has been
badly misinterpreted. The genus is restricted here
to include only A. quercifolia, and its association
with the Microcionidae rests solely upon the
original placement of the type species in Antherochalina (whereby many of the species
originally described for the genus by Lendenfeld
(1888) are true microcionids). By comparison, de
Laubenfels (1936a) originally intended Quer-
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
^
87
ciclona to include Haliclona-like species
(Haplosclerida), but this is only true for a second
species, Arenochalina arabica Keller (which was
subsequently transferred to A renosclera PulitzerFinali (1982), but is probably an arenaceous Callyspongia). Conversely, the type species belongs
to the Axinellidae.
Antherochalina quercifolia is structurally close
to Isociella (s.s.) and Phakellia of authors. It has
an almost regular isodictyal choanosomal
reticulation of spongin fibres cored by primary
and secondary skeletal lines, without any
pronounced compression of the axial skeleton but
with some differentiation of the axial and extraaxial skeletons, and it also lacks a specialised
ectosomal skeleton. Details of the holotype (reexamined and redescribed above) bear little
resemblance to Keller's (1889) original description, and it is concluded that Querciclona is most
appropriately referred to Phakellia (Axinellidae)
(Hooper & Levi, 1993b). A specimen label in M.
Burton's handwriting accompanying a fragment
of the holotype in the BMNH (BMNH1908.9.24.179) suggests further that A. quercifolia is a synonym of Phakellia donnani
(Bowerbank), but this synonymy is so far unsubstantiated.
Scopalina Schmidt, 1862
Scopalina Schmidt, 1862: 78; Gray, 1867: 535.
TYPE SPECIES. Scopalina lophyropoda Schmidt,
1862: 79 (by monotypy) (holotype LMJG15117/154).
Thickly encrusting growth form. Surface
prominently conulose. Choanosomal skeleton
hymedesmoid, with heavy spongin fibres forming basal layer of spongin on substrate, with
ascending non-anastomosing fibre nodes cored
by plumose brushes of auxiliary styles. Echinating megascleres absent. Ectosome lacks
specialised spiculation, although brushes of
auxiliary styles protrude through surface.
Mesohyl matrix heavy. Megascleres only a single
category of long auxiliary style. Microscleres
absent.
REMARKS. Gray (1867) and de Laubenfels
(1936a) suggested that Scopalina was
Microciona-like with plumose skeletal architecture and greatly reduced spiculation. Although
skeletal architecture is hymedesmoid containing
non-anastomosing ('microcionid') fibre nodes,
and the possession of only a single category of
auxiliary spicules could be construed as reduction
of the typical Microciona condition (analogous to
Dendrocia (Microcionidae) or Amphilectus s.l.
(Myxillidae)), no other characters support its inclusion in the Microcionidae. Boury-Esnault
(1971) and Uriz (1982) include it in the
Halichondrida, and suggested close affinites with
Stylinos.
KEY TO GENERA
1(0). Choanosomal skeleton more-or-less undifferentiated, unstructured ^ Artemisina
Choanosomal skeleton well structured, hymedesmoid to reticulate, but lacking any differentiated
components
2
Choanosomal skeleton well structured,
predominantly reticulate, differentiated into two
distinct components ^ 7
2(1). Choanosomal fibres or skeletal tracts cored by
one or more category of principal spicules . ^ 3
Choanosomal fibres or skeletal tracts cored by
auxiliary spicules but partially or wholly
5
replaced by detritus ^
Choanosomal fibres or skeletal tracts cored by
auxiliary spicules identical to those in ectosomal
and subectosomal skeletons ^ 6
Choanosomal fibres or skeletal tracts cored by
auxiliary spicules different from those in
peripheral skeleton ^ Panda ros
3(2). Choanosomal skeleton without any marked
axial compression or differentiated axial and
extra-axial regions
4
Choanosomal skeleton with noticeably compressed axis and well differentiated axial and
extra-axial (radial, plumose or plumoreticulate)
regions ^ Clathria (Axociella)
Choanosomal skeleton hymedesmoid or
microcionid, with basal layer of spongin lying
on substrate (with or without ascending, nonanastomosing fibre nodes), and bases of principal spicules standing perpendicular to
substrate ^ Clathria (Microciona)
Choanosomal skeleton evenly renieroid reticulate
throughout with well developed spongin fibres
cored by smooth principal styles
Clathria (Isociella)
^
4(3). With single category of (subectosomal)
auxiliary spicule on surface forming tangential,
paratangential or plumose tracts
^
Clathria (Clathria)
With two categories of auxiliary spicules, smaller
ectosomal spicules generally overlaying larger
subectosomal spicules forming discrete bundles
or continuous palisade on surface
^
Clathria (Thalysias)
5(2). Special category of (acantho)styles present
echinating fibres, differentiated from principal
spicules ^ Clathria (Wilsonella)
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Echinating spicules styles or acanthostyles undifferentiated from principal spicules coring sponHolopsamma
gin fibres
6(2). Special category of (acantho)styles present
echinating fibres, differentiated from principal
spicules ^ Clathria (Dendrocia)
Echinating spicules styles or acanthostyles representing principal spicules, but different from
those coring fibres
^ Echinochalina (Echinochalina)
Echinating spicules oxeas or anisoxeas representing principal spicules, but different from
those coring fibres
^ Echinochalina (Protophlitaspongia)
7(1). Primary skeleton renieroid cored by axially or
basally compressed tracts of acanthostyles,
secondary skeleton cored by smooth principal
styles in plumose, subisodictyal or plumoreticulate tracts ^ Antho (Antho)
Primary skeleton renieroid cored by axially or
basally compressed tracts of acanthostrongyles,
secondary skeleton cored by smooth principal
styles in plumose, subisodictyal or plumoreticuAn (ho (Plocamia)
late tracts
Primary skeleton axially compressed spongin
fibres cored by renieroid tracts of sparsely
spined principal styles intermingled with
plumose or plumoreticulate tracts of smooth
principal styles, overlaid by secondary extraaxial plumose skeleton cored by larger smooth
principal styles ^ Antho (lsopenectya)
Primary renieroid reticulate skeleton cored by
smooth principal styles and echinated by identical spicules, with secondary radial extra-axial
skeleton on exterior edge of skeleton only cored
by larger smooth principal styles
Echinoclathria
^
DESCRIPTION OF AUSTRALIAN SPECIES
Clathria Schmidt, 1862
Refer to subgenera for synonymy.
TYPE SPECIES. Clathria compressa Schmidt, 1862:
58 (designated Schmidt, 1864: 35).
DEFINITION. Monactinal auxiliary spicules in 1
or 2 categories forming ectosomal skeletons
ranging from sparse, mostly membraneous (C.
(Microciona)), sparse, tangential (C. (Clathria))
to relatively dense, erect (C. (Thalysias)).
Choanosomal skeletal tracts usually enclosed
within spongin fibres, sometimes simply with
nodal spongin; fibres cored by smooth, basally
spined or partially spined principal monactinal
megascleres, usually geometrically different
from auxiliary megascleres, sometimes secondarily lost and cored by single category of
auxiliary subtylostyle (C. (Dendrocia)), or sometimes replaced partially or fully by detritus in
fibres (C. (Wilsonella)). Echinating megascleres
partially or entirely acanthose, occasionally
smooth or vestigial spination, sometimes secondarily lost (C. (Axociella), C. (Isociella)).
Choanosomal structure ranges from leptoclathrid
to microcionid plumose (C. (Microciona)),
renieroid (C. (Isociella)), plumoreticulate or
reticulate, with (C. (Axociella)) or without compressed axis and radial extra-axial regions.
Microscleres include palmate isochelae and
modified forms, and toxas with smooth or spined
points, occasionally absent.
REMARKS. This definition is necessarily broad
to encompass the 7 subgenera included in
Clathria, showing a wide spectrum of character
states, most of which are interpreted as secondary
losses rather than unique apomorphies, and many
characters show intermediate states making it
virtually impossible to maintain generic boundaries recognised by earlier authors.
Clathria (Clathria) Schmidt, 1862
Clathria Schmidt, 1862: 57.
Allocia Hallmann, 1920: 768.
Antherochalina Lendenfeld, 1887b: 741.
Bipocillopsis Koltun, 1964a: 79.
Diclyociona Topsent, 1913a: 579.
Labacea de Laubenfels, 1936a: 125.
Ligrota de Laubenfels, 1936a: 125.
Litaspongia de Laubenfels, 1954: 162.
Marleyia Burton, 1931a: 346.
Ramoses de Laubenfels, 1936a: 109.
Thalyseurypon de Laubenfels, 1936a: 107.
TYPE SPECIES. Clathria compressa Schmidt, 1862:
58 (by subsequent designation of Schmidt, 1864).
DEFINITION. Ectosomal skeleton composed of
a single undifferentiated category of auxiliary
megasclere; choanosomal skeletal structure
plumoreticulate or reticulate, usually without
marked difference between axial and extra-axial
regions; spongin fibres cored by completely
smooth, basally spined or partially spined principal megascleres, geometrically differentiated
from auxiliary megascleres, but sometimes
secondarily lost; echinating megascleres entirely
or partially acanthose, occasionally smooth,
sometimes secondarily lost. Microscleres include
palmate isochelae and modified forms, and toxas
with smooth or spined points.
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89
TABLE 1. Comparison between present and published
records of Clathria (Clathria) a ngulifera
Dendy.Measurements in p.m, denoted as range (and
mean) of spicule length x spicule width (N=25).
SPICULE
Holotype (NMO2286)
(Victoria)
Specimens (N=2)
(SE.Queensland)
Choanosomal
principal styles
1444172.4)-211
x 3-(4.3)-6
Subectosomal
auxiliary styles
168-(175.0)-254
x 2.5-(3.6)-4.5
x 3.5-(4.1)-4.5
128-(141.3)-150
x 3-(3.8)-5
181-(195.8)-223
x 342.3)-5
58-(61.8)-72
x 2-(3.3)-5
Chelae
5.5-(7.1)-8.5
5-(6.7)-9
Toxas
18467.4)-101
x 0.541.7)-3.3
absent
Echinating
acanthostyles
58-(64.2)-72
REMARKS. Of the 154 named species described
in, or subsequently referred to Clathria, or one of
its synonyms listed above, 112 are retained in this
subgenus. There are 31 (2 new) species known
from Australian waters.
Clathria (Clathria) angulifera Dendy, 1896
(Figs 26-27, Plate 1A, Table 1)
Clathria anguhfera Dendy, 1896: 32; Ayling et al.,
1982: 100; Hooper & Wiedenmayer, 1994: 258.
Thalysias angulifera; de Laubenfels, 1936a: 104.
cf. Clathria conectens Hallmann, 1912: 247.
MATERIAL. HOLOTYPE: NMVG2286 (fragment
BMNH1902.10.18.323): Outside Port Phillip Heads,
Vic, 38°09'S, 144°52'E, coll. J.B. Wilson (dredge).
OTHER MATERIAL: Queensland — QMG303230,
QMG303960.
HABITAT DISTRIBUTION. Submerged rock reef;
10-30m depth; Bass Strait (Vic), N. Stradbroke I. and
Noosa Heads (SEQ) (Fig. 260).
DESCRIPTION. Shape. Thickly lamellate, ap-
proximately 9 cm long, 30mm wide, 50mm high,
with numerous bulbous lobate digits up to 15mm
long.
Colour Bright yellow-orange alive (Munsell
2.5Y 8/10), grey-brown in ethanol.
Oscules. Occasional large oscules, 1.5 3mm
diameter, on edges and between lobes.
Texture and surface characteristics. Firm, compressible; glabrous skin-like surface.
Ectosome and subectosome. Ectosomal skeleton
membraneous, with loose, irregular, tangential,
occasionally paratangential or erect tracts of subectosomal auxiliary subtylostyles; rarely protruding above surface. Subectosomal portion of
peripheral skeleton slightly plumose, with sparse
-
diverging brushes of auxiliary megascleres which
arise from ascending choanosomal tracts.
Choanosome. Choanosomal skeleton regularly
reticulate, cavernous, vaguely renieroid; fibre
anastomoses produce wide oval, rectangular or
sometimes slightly triangular meshes; spongin
fibres thin, only lightly invested with spongin,
barely differentiated from mesohyl matrix; spongin fibres cored by uni- or paucispicular tracts of
choanosomal principal megascleres; echinating
acanthostyles sparsely dispersed on fibres;
mesohyl matrix heavy, lightly pigmented, containing few straight or sinuous subectosomal
auxiliary megascleres and abundant spherical incubated larvae (275-34511m diameter) with well
differentiated cellular development.
Megascleres (Table 1). Choanosomal principal
styles short, thin, fusiform, straight or slightly
curved towards basal end, with rounded or only
slightly subtylote, smooth bases.
Subectosomal auxiliary subtylostyles straight,
thin, relatively long, almost hastate, with smooth
and only slightly subtylote bases.
Acanthostyles small, thin, prominently subtylote, with rudimentary granular spination and
small aspinose 'neck' near basal constriction.
Microscleres (Table 1). Palmate isochelae very
small, weakly silicified, unmodified.
Toxas not common, relatively large, v-shaped,
with sharply angular central curvature and
straight arms.
REMARKS. The thickly lamellate, lobate growth
form, glabrous surface, relatively open, slightly
renieroid, paucispicular skeletal structure (cf.
Dendy, 1896), and aspects of spiculation in C.
(C.) angulifera are quite distinctive and unusual
amongst microcionids (cf. Hallmann, 1912).
Many of these features are similar to those in C.
(T)aphylla sp. nov. from the Houtman Abrolhos,
WA, although the latter has a specialised ectosomal skeleton and hence is included in C.
(Thalysias) rather than C. (Clathria). These
species together with C. (C.) hispidula are sister
species belonging to a species group termed here
`angulifera' group. They are compared in detail
in discussion on C. (T) aphylla.
This species is also slightly reminiscent of C.
(C.) conectens, differing in that megascleres
coring fibres are true principal spicules rather
than just undifferentiated auxiliary spicules (albiet these differences are not as well marked in C.
(C.) angulifera as in many other species of
Clathria). This reduction in coring spicules was
interpreted by Hallmann (1912) to be charac-
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FIG. 26. Clathria (Clathria) angulifera Dendy (holotype NMVG2286). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Palmate isochelae. E, V-shaped toxas. F,
Section through peripheral skeleton. G, Australian distribution. H, Holotype.
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FIG. 27. Clathria (Clathria) angulifera Dendy (A-B, G, holotype NMVG2286; C-E, QMG303230). A,
Choanosomal skeleton. B, Fibre characteristics (x456). C, Ectosomal paratangential skeleton. D, Palmate
isochelae. E, Echinating acanthostyles. F, Acanthcstyle spines. G, Principal styles and v-shaped toxas.
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teristic of C. (Wilsonella) (although this is shown
here to be an erroneous interpretation of Wilsonella). De Laubenfels (1936a) also transferred
C. (C.) angulifera to Thalysias, but this was completely unjustified.
Although known only from 2 widely separated
localities it is probable that it is more widely
distributed within shallow coastal waters in
southeastern Australia.
Clathria (Clathria) arcuophora Whitelegge,
1907 (Figs 28-29)
Clathria arcuophora Whitclegge, 1907: 491,500-501,
p1.45, fig.29; Hallmann, 1912: 211, 229, 234- 237,
260, 263, text-fig. 49-49a; Hooper & Wiedenmayer,
1994: 258.
Thalysias arcuophora; de Laubenfels, 1936a: 105.
Microciona arcuophora; Vosmaer, 1935a: 611, 649,
665.
MATERIAL. LECTOTYPE: AMG4346: Off Barranjoey, S. coast of NSW, 33 °35'S, 151 °20'E, 50-66m
depth, coll. FIV `Thetis' (dredge). PARALECTOTYPE: AMZ1209: Off Botany Bay, NSW, 34°00'S,
151°11'E, 40-46m depth, coll. FIV `Thetis' (dredge).
HABITAT DISTRIBUTION. Depth range 30-90m;
substrate unknown; central and S. coast of NSW (Fig.
28G).
DESCRIPTION. Shape. Thinly flabelli form, 170mm
long, 120mm maximum breadth, up to 4mm thick,
with long cylindrical stalk, 90mm long, 13mm
diameter, and rounded or slightly lobate margins.
Colour Grey-brown in dry state.
Oscules. Surface covered with evenly distributed
minute oscules, up to 1.5mm diameter.
Texture and surface characteristics. Harsh in dry
state; surface optically smooth.
Ectosome and subectosome. Ectosome microscopically hispid, with regularly distributed
choanosomal principal styles forming erect
plumose brushes, and protruding from peripheral
fibre endings; tangential layer of subectosomal
auxiliary spicules lying on or just below surface,
at base of principal styles. Subectosome with only
slightly plumose tracts of choanosomal
megascleres, virtually undifferentiated from
choanosomal skeleton.
Choanosome. Choanosomal skeleton more-orless regularly reticulate, sub-renieroid; axial
skeleton slightly compressed, peripheral skeleton
slightly plumo-reticulate; branching between moderately heavy spongin fibres produces ovoid to
square meshes, and fibres differentiated into
primary ascending and secondary transverse components; primary fibres with pauci- to multispicular
core of choanosomal principal styles; secondary
fibres with uni- or bispicular tracts of principal
spicules; echinating acanthostyles sparse, slightly more abundant on peripheral fibres; mesohyl
matrix light, with only few subectosomal
auxiliary megascleres dispersed between fibres.
Megascleres. Choanosomal principal styles
thick, slightly curved, slightly fusiform, with
rounded smooth bases. Length 1764258.4)444p,m, width 12-(22.2)-27p.m.
Subectosomal auxiliary subtylostyles small,
thin, straight, with smooth or microspined bases,
bases slightly subtylote, points fusiform. Length
1924231.1)-276pm, width 3.5-(4.8)-6pm.
Acanthostyles small, slightly subtylote, with
relatively even granular spination. Length 68(93.2)-104pm, width 6.547.5)-10p.m.
Microscleres. Palmate isochelae large, unmodified. Length 16-(21.4)-26p.m.
Toxas intermediate between oxeote and oxhorn, thick, gently curved at centre, with straight
arms or slightly reflexed points. Length 28(104.0)-132p.m, width 1.5-(4.8)-6pm.
REMARKS. This species is distinctive by its
renieroid choanosomal skeleton, plumo-reticulate subectosomal skeleton, distinctive spongin
fibres which contain only few but very thick
coring spicules, and an ectosomal region which
is dominated by plumose brushes of principal
spicules. The renieroid skeletal construction is
superficially similar to Antho (lsopenectya) and
Clathria(lsociella)), and several other (otherwise
unrelated) groups (some Raspailiidae (e.g., Ectyoplasia), Axinellidae (e.g., Axinella aruensis
(Hentschel, 1912)), Iophonidae (e.g., Acarnus)
and Myxillidae (Lissodendoryx) (see also
remarks for C. (C.) crassa)). Clathria (C.) arcuophora is most reminiscent of Ectoplasia frondosa (Lendenfeld) (Raspailiidae; see Hooper,
1991: figs 47-48), which was originally described
in the Microcionidae, having closely comparable
architecture, fibre characteristics and geometry
of some spicules. Within the Microcionidae, C.
(C.) arcuophora is also similar to C. (C.)
biclathrata in spicule geometry and fibre characteristics, although there are substantial differences between them in skeletal construction. The
species should also be compared to C. (C.)
striata, which differs mainly in the morphology
of its toxa.
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FIG. 28. Clathria (Clathria) arcuophora Whitelegge (paralectotype AMZI 209). A, Choanosomal principal
styles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Palmate isochelae. E, Oxeote
and oxhorn toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype AMG4346.
Clathria (Clathria) biclathrata sp. nov.
(Figs 30-31, Table 2)
Microciona clathrata Whitelegge, 1907: 491-494,
p1.46, fig. 38-38a; Vosmaer, 1935a: 608; Hooper
& Wiedenmayer, 1994: 258.
Clathria clathrata; Hallmann, 1912: 209.
Dictyociona clathrata; de Laubenfels, 1936a: 110.
Thalysias clathrata; de Laubenfels, 1953: 527.
Not Tenacia clathrata Schmidt, 1870: 56,80.
Not Clathria clathrata; Vosmaer, 1880: 153; Ridley &
Dendy, 1887: 147; Wilson, 1902: 397-398; Alcolado, 1976: 5.
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FIG. 29. Clathria (Clathria) arcuophora Whitelegge (paralectotype AMZ1209). A, Choanosomal skeleton. B,
Fibre characteristics (x409). C, Echinating acanthostyle. D, Acanthostyle spination. E, Base of auxiliary
subtylostyle. F, Palmate isochelae. G, Oxeote toxas. H, Oxhorn toxas.
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TABLE 2. Comparison between present and published
records of Clathria(Clathria)biclathrata, sp.nov. All
measurements are given in pm, denoted as range (and
mean) of spicule length x spicule width (N=25).
SPICULE
Choanosomal
principal
styles
Lectotype
(AMG4355)
Paralectotype
(AMG10530)
2534372 . 3)446 x 18(22.8)-33
226(337.5)423 x 17(22.6)-28
Specimen
(N=I)
2334319.8)418 x 14(17.8)-24
Subectosomal I 32-(214.0)125-(165.4)1444197.0)auxiliary
293 x 2-(4.9)-7 243 x 3-(5.5)-7 253 x 3-(4.6)-7
styles
734158.4)-231 1144161.1)-193 1134150.2)-193
Echinating
acanthostyles x 4.5-(14.7)-I9 x 11-(13.6)-18 x 8-(12.4)-I6
Chelae
7-(10.4)-I4
6-(8.6)-11
4-(7.8)-12
Toxas I
28-(93.0)-149
x 0.8-(2.4)-3.5
Toxas II
4-(10.2)-I8
x 0.841.7)-2.0
434102.8)-141
x I.8-(2.8)-3.5
6-(I8.0)-36
38-(66.6)-104
x 1-(2.7)-4
3-(12.4)-24
MATERIAL. LECTOTYPE: AMG4355: (dry) Off
Woolongong, NSW, 34 °25'S, 151 ° 10'E, 110-112m
depth, coll. FIV 'Thetis' (dredge). PARALECTOTYPES: AMG10530: (dry) unknown locality,
NSW, coll. FIV `Thetis' (dredge). AMG10531
(presently missing): (label `Dictyociona clathrata,
cotype'). OTHER MATERIAL: NEW SOUTH
WALES - AMZ994.
HABITAT DISTRIBUTION. Up to 112m depth, substrate unknown; S. coast of NSW (Fig. 30H).
DESCRIPTION. Shape. Clavulate to subspherical, bushy, honeycombed mass, 85-105mm high,
30-55mm wide, up to 40mm maximum thickness,
composed of numerous thin, cylindrical, tightly
anastomosing branches; small cylindrical stalk,
8-15mm long, up to 8mm diameter.
Colour Live colouration unknown, grey-brown
to dark brown in dry state.
Oscules. Small oscules, 1-2.5mm diameter, on
edges of surface microconules.
Texture and surface characteristics. Surface very
hispid, minutely porous, with numerous slightly
elevated microconules; texture unusually tough.
Ectosome and subectosome. Ectosomal skeleton
a tangential layer of subectosomal auxiliary
styles in multispicular tracts, with numerous
choanosomal principal spicules protruding and
extending a long way through surface; subdermal
skeleton, if present, totally obscured by dense
mass of erect choanosomal megascleres.
Choanosome. Choanosomal skeletal architecture
irregularly reticulate, slightly renieroid, with
heavy spongin fibres forming tight meshes, and
some compression of axial fibres; spongin fibres
not clearly divisible into primary or secondary
components, but merely ascend and diverge
towards surface; fibres mostly aspicular, sometimes with one or few choanosomal principal
subtylostyles in core, heavily echinated by both
acanthostyles and choanosomal principal subtylostyles (the latter `spicate' in arrangement),
and some intermediate spicules with rudimentary
spines on shaft; echinating megascleres most
abundant on peripheral fibres; mesohyl matrix
heavy, with few subectosomal auxiliary subtylostyles between meshes.
Megascleres (Table 2). Choanosomal principal
subtylostyles slightly curved or straight,
fusiform, slightly constricted at base, heavily
spined bases (smaller examples may also have
scattered spines on shafts).
Subectosomal auxiliary subtylostyles short,
fusiform, straight, with slightly subtylote
microspined bases.
Acanthostyles relatively long, thick, slightly
curved, with prominent subtylote bases, with
evenly distributed large spines over entire spicule
or with an aspinose region proximal to base.
Microscleres (Table 2). Palmate isochelae, some
twisted.
Toxas include larger thick wing-shaped forms
with large central curvature, slightly reflexed
arms and microspined points, and smaller oxhorn
forms, the smallest with abbreviated arms.
REMARKS. This species is characterised by its
compressed skeletal architecture, fibre and ectosomal features, and it is unlikely to be confused
with other members of Hallmann's (1912)
spicata group of microcionid species (cf. Hooper
et al., 1990) which have principal spicules
protruding through fibres and surface skeletons
but few within fibres themselves. A feature overlooked by previous authors is the presence of
spinous extremities on toxas, which are virtually
identical to those of type species of Clathria, C.
(C.) compressa, also occurring in C. (C.) juncea,
C. (C.) lobata, and Artemisina suberitoides. The
geometry of spicules in C. (C.) biclathrata is
similar to those in C. (C.) arcuophora, although
these species differ quite substantially in their
skeletal architecture.
Clathria (Clathria) caelata Hallmann, 1912
(Figs 32-33, Table 3)
Clathria caelata Hallmann, 1912: 139, 177, 206, 211-
216, p1.33, fig.4, text-fig.43; Hooper & Wiedenmayer, 1994: 258.
Clathria coelata; Burton & Rao, 1932: 336 [lapsus].
Pseudanchinoe caelata; de Laubenfels, 1936a: 109.
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FIG. 30. Clathria (Clathria) biclathrata sp.nov. (lectotype AMG4355). A, Choanosomal principal subtylostyle
and spined subtylostyle. B, Echinating acanthostyles. C, Subectosomal auxiliary subtylostyles. D, Palmate
isochelae. E, Wing-shaped toxas. F, Oxhorn toxas. G, Section through peripheral skeleton. H, Australian
distribution. I, Lectotype AMG4355.
Not Clathria inanchorata Ridley & Dendy, 1886:475;
Ridley & Dendy, 1887: 150, p1.28, fig.4, p1.29, figs
13,13a.
cf. Microciona prolifera; Vosmaer, 1935a: 611, 648,
664.
MATERIAL. LECTOTYPE: AMZ778: (wet) 64km
W. of Kingston, SA, 36 °50'S, 139 °05'E, 60m depth,
coll. FIV 'Endeavour' (dredge; label `4th
consignment'). PARALECTOTYPES: AME53: (dry)
same locality. AMZ952-953: unspecified locality, W.
coast Tas. ('ref. G255'). OTHER MATERIAL: TASAME2273.
HABITAT DISTRIBUTION. Depth 53-106m; substrate unknown; Kingston (SA), Bass Strait (Vic), Cape
Barren, W coast (Tas) (Hallmann, 1912).
DESCRIPTION. Shape. Planar or multiplanar,
digitate fans, 110-150mm high, 70-90mm wide,
with short cylindrical stalk, 8-17mm long, 5-
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FIG. 31. Clathria (Clathria) biclathrata sp.nov. (lectotype AMG4355). A, Choanosomal skeleton. B, Ectosomal
skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spination. F, Wing-shaped toxa.
G, Oxhorn toxas. H, Palmate isochelae.
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FIG. 32. Clathria (Clathria) caelata Hallmann (paralectotype AME53). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyle. C, Intermediate echinating and principal style. D, Echinating acanthostyle.
E, Accolada toxa. F, Oxhorn toxas. G, Section through peripheral skeleton. H, Australian distribution. I,
Paralectotype AMZ953. J, Paralectotype AMZ952.
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FIG. 33. Clathria (Clathria) caelata Hallmann (A-B, Lectotype AMZ778; C-G, E2273). A, Choanosomal
skeleton. B, Fibre characteristics. C, Ectosomal skeleton. D, Accolada toxa. E, Oxhorn toxas, F, Echinating
acanthostyles. G, Acanthostyle spination.
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lOmm diameter, rounded lobate, or digitate margins.
Colour Llight to dark brown preserved.
Oscules. Small pores on margins of lobes, up to
1.5mm diameter.
Texture and surface characteristics. Surface
highly rugose, with numerous close-set
microconules, subdermal canals and grooves, between which extends a skin-like dermal
membrane; texture firm, compressible.
Ectosome and subectosome. Ectosome microscopically hispid, with plumose brushes of
choanosomal principal styles protruding through
surface, and with thin layer of subectosomal
auxiliary styles tangential to surface; subdermal
region not markedly differentiated from choansome containing bundles of diverging principal
styles embedded in peripheral fibres; peripheral
fibres heavily echinated on their exterior surfaces; acanthostyles may extend into ectosomal
skeleton.
Choanosome. Choanosomal skeletal architecture
irregularly reticulate, with well developed spongin fibres forming ovoid to elongate meshes; fibre
anastomoses relatively cavernous in axis, although smaller in peripheral skeleton; fibres
clearly differentiated into primary ascending and
secondary transverse components, although fibre
diameter is consistent throughout skeleton:
primary fibres contain pauci- or multispicular
tracts of choanosomal principal subtylostyles,
forming a radial architecture; secondary fibres
uni- or aspicular; spongin fibres echinated on
external surfaces only, with a variable density of
echinating acanthostyles, mostly relatively light
except at the periphery; choanosomal styks also
echinate fibres, particularly at fibre nodes;
mesohyl matrix moderately heavy, granular, pigmented; extra-fibre megascleres (subectosomal
subtylostyles) usually abundant.
Megascleres (Table 3). Choanosomal principal
subtylostyles thick, straight or slightly curved,
fusiform, with slightly subtylote smooth bases,
although some examples are obvious intermediates to echinating acanthostyles, bearing
rudimentary spines on the shaft.
Subectosomal auxiliary subtylostyles straight
or slightly curved, fusiform, relatively thin, evenly rounded or slightly subtylote bases, and
smooth or microspined bases.
Echinating acanthostyles variable in length,
subtylote, with evenly spined shafts on smaller
forms, or with aspinose necks on larger forms.
Microscleres (refer to Table 3 for dimensions).
Isochelae absent.
TABLE 3. Comparison between present and published
records of Clathria (Clathria) caelata (Hallmann).
All measurements are given in m, denoted as range
(and mean) of spicule length x spicule width (N=25)
Paralectotype
(AME53)
Specimen
(N=1)
Choanosomal 1484215.4)-276
principal
x 8-(12.2)-16
styles
1544222.2)-262
x 9-(11.8)-14
186-(254.5)-353
x 6-(9.4)-10
Subectosomal 128-(216.4)-294
auxiliary
x 2-(3.2)-5
styles
45-(71.9)-122
Echinating
x 245.1)-8
acanthostyles
1344234.8)-324
x 3-(4.6)-7
1664267.0)-355
x 2-(3.5)-7
64-(89.4)-131
x 4-(6.4)-8
23-(64.4)-120
x 2-(4.8)-8
SPICULE
Lectotype
(AMZ778)
Chelae
absent
absent
absent
Toxas I
1224151.7)-190
x 0.5-(1.1)-1.5
72-(101.8)-165
x 0.8-(1.4)-2
86-(121.1)-165
x 0.5-(1.4)-2
Toxas II
2I-(55.9)-83
x 1-(2.3)-4
13-(38.8)-75
24-(48.2)-92
x 1-(2.1)-3.5^, x 1-(2.2)-4
Toxas separated into two morphs: I - long, thin
accolada toxas, with straight points and slight
central curvature; II - relatively thick oxhorn
toxas, ranging from almost straight with only
slight and angular central curvature, to widely
curved at the centre with reflexed points.
REMARKS. Not all of the specimens described
by Hallmann (1912) belong to this species:
AME771, E772 and E773 are species belonging
to Axinellidae - Reniochalina (2 specimens) and
Acanthella, respectively. All three specimens
look very similar in external morphology to C.
(C.) caelata, but differences are immediately obvious upon examination of the skeleton.
Among the few known specimens of this
species there is relatively high variability in
choanosomal skeletal construction although this
is difficult to define concisely. The development
of spongin fibres, the degree to which the skeletal
meshes are compressed or elongated, and the
density and pigmentation of the mesohyl matrix
may vary between specimens. Growth form and
spicule geometry appear to be closely comparable between all specimens, but some (e.g.,
AME2273) have predominantly long thin toxas,
whereas others (e.g., AMZ952) have mostly short
and thick toxas. All specimens have both
choanosomal styles and acanthostyles echinating
fibres, and in this respect the species belongs to
Hallmann's (1912) spicata' group, together with
Clathria (C.) biclathrata, C. (C.) inanchorata, C.
(Thalysias) costifera, C. (T) coppingeri, C. (T)
lendenfeldi and others (Hooper et al., 1990).
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101
Clathria (Clathria) calopora Whitelegge, 1907
Clathria calopora Whitelegge, 1907: 499-500, p1.46,
fig.34; Hallmann, 1912: 211; Hooper & Wiedenmayer, 1994: 259.
Thalysias calopora; de Laubenfels, 1936a: 105.
MATERIAL. HOLOTYPE: AM (presently missing):
Shoalhaven Bight, NSW, 34 ° 49'S, 151°04'E, 30-90m
depth, 1.vii.1911, coll. FIV 'Endeavour' (trawl).
HABITAT DISTRIBUTION. Depth 27-32m, substrate unknown; S. coast of NSW.
DESCRIPTION. Shape. Branching, 195mm
high, with thin, bifurcate, cylindrical, mostly
non-anastomosing digits, 3-8mm diameter, with
tapering points, short cylindrical stalk.
Colour Live colouration unknown, greyish-yellow preserved.
Oscules. Small oscules, about lifun diameter,
scattered over surface.
Texture and surface characteristics. Surface uneven, minutely hispid, honeycomb-like, with
prominent bulbous conulose processes scattered
over surface; texture hard, incompressible in dry
state.
Ectosome and subectosome. Ectosome microscopically rugose, close-meshed reticulation of
peripheral spongin fibres covered by a thin
membrane; ectosomal skeleton with a thin tangential layer of subectosomal auxiliary subtylostyles, also forming plumose tufts on surface,
through which choanosomal principal styles from
peripheral fibres protrude, individually or in
plumose brushes; subdermal auxiliary
megascleres run parallel with peripheral fibres,
but not forming organised extra-fibre tracts.
Choanosome. Choanosomal skeleton irregularly
reticulate, with some axial and extra-axial differentiation; spongin fibres relatively heavy,
clearly divided into primary ascending and
secondary transverse components; primary fibres
sinuous, forming radial architecture, with a
paucispicular core of choanosomal principal
styles; primary fibres more compressed at axis
than at periphery; secondary fibres less common,
with uni- or paucispicular core of megascleres;
branching of spongin fibres produces elongate
meshes in axis and round or rectangular meshes
in periphery; spongin fibres very heavily
echinated, particularly in peripheral regions;
numerous sinuous toxodragmata dispersed in
mesohyl between fibres.
Megascleres. Choanosomal principal styles
thick, slightly curved, fusiform, with rounded
smooth bases. Length 300-500p,m, width 1522m.
Subectosomal auxiliary subtylostyles thin,
straight or slightly curved, with smooth slightly
subtylote bases. Length 150-200p,m, width 24p,m.
Acanthostyles short, stout, evenly spined,
spines large. Length 30-80p,m, width up to
12p.m.
Microscleres. Palmate isochelae. Length 810t.m.
Toxa morphology unknown, apparently long,
slender, with large central curvature. Length ?
200-300p,m, width ? up to 21.1,m.
REMARKS. This species is known only from
Whitelegge's (1907) poor description. It is not
possible to determine whether dimensions of
echinating acanthostyle (cited as 0.3-0.8mm long
by Whitelegge) is merely a typographical error.
Whitelegge gave no indication of whether C. (C.)
calopora has a special ectosomal skeleton, but
described the species as having a tangential or
paratangential layer of subdermal (auxiliary)
megascleres, through which protrude
choanosomal (principal) styles. Consequently,
there was no justification for de Laubenfels
(1936a) referring the species to Thalysias.
Clathria (Clathria) chelifera (Hentschel, 1911)
(Figs 34-35, Table 4)
Spanioplon cheliferum Hentschel, 1911: 362-363,
fig.42; Hentschel, 1912: 368-369.
Allocia chehfera; Hallmann, 1920: 768; Bergquist &
Fromont, 1988: 96, fig. 8c, pis 45e-f, 46a-c, table
73; Dawson, 1993: 44 (note).
Clathria chelifera; Dendy, 1922: 70-71, p1.14, fig.
3a-e; Hooper & Wiedenmayer, 1994: 259.
Not Microciona chelifera Lëvi,1960a: 70, fig. 12.
MATERIAL. HOLOTYPE: ZMH (not seen) (fragment ZMB4440): precise locality unknown, Perth
region, WA, 1905, coll. W. Michaelsen & R.
Hartmeyer (dredge). PARATYPE: SMF 1571 (fragment MNHNDCL2327): same locality. OTHER
MATERIAL: VIETNAM - PIBOC-05-216 (fragment
QM G300058).
HABITAT DISTRIBUTION. 10-100m depth; rock or
gravel substrates; Arafura Sea (NT) and Perth region
(S. WA) (Fig. 34F). Also Indian Ocean (Amirante),
New Zealand (Three Kings Is) and South China Sea
(Hon Trung Lon, Vinh Loi coast, S. Vietnam).
DESCRIPTION. Shape. Arborescent, foliose,
planar growth form, up to 50mm high made up of
fused porous-reticulate lamellae 10-15mm thick.
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 34. Clathria (Clathria) chelifera (Hentschel) (paratype SMF1571). A, Choanosomal principal style. B,
Subectosomal auxiliary styles (quasitylotes). C, Echinating acanthostyles. D, Palmate isochelae. E, Section
through peripheral skeleton (ectosome on far right). F, Australian distribution. G, QG300058.
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU
FIG. 35. Clathria (Clathria) chelifera (Hentschel) (QMG300058). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Palmate isochelae. G, Pattern of echinating
spicules. H, Section of peripheral skeleton.
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
TABLE 4. Comparison between present and published
records of Clathria (Clathria)chelifera (Hentschel).
All measurements are given in 1.1.m, denoted as range
(and mean) of spicule length x spicule width (N=25).
SPICULE
Paratype
(SMF1571)
(Perth, WA)
Specimen
Bergquist &
Fromont
(1988) (Three
Kings Is, NZ)
Specimen
(N=1)
(S. Vietnam)
Choanosomal
principal styles
Subectosomal
auxiliary styles
1404150.1)-161
x 4-(6.1)-7
1734193.4)-205
x 2-(3.9)-5
2704404)-550
x 15-(18)-20
280-(334)-375
x 4-(5)-6.5
150-(164.6)-174
x 4-(6.4)-10
1924204.0)-212
x 3-(4.7)-7
Echinating
acanthostyles
Chelae I
84-(93.2)-100
x 7-(7.8)-9
162-(207)-290
x 12.5-(14)-19
10341165)-132
x 5-(8.0)-10
13-(14.6)-1
20421)-23
15-(17.6)-20
Chelae H
7-(8.7)-10
absent
9-(10.3)-12
Colour Red alive (10R 5/8), beige or yellow
brown in ethanol.
Oscules. Not seen.
Texture and stuface characteristics. Surface
hispid, uneven, irregularly conulose, with
transparent ectosomal membrane stretched between adjacent conules; texture firm, compressible.
Ectosome and subectosome. Ectosomal skeleton
with a tangential layer of auxiliary tylotes and
numerous isochelae scattered between, supporting the membraneous ectosomal covering; subectosomal region with paratangential tracts of
auxiliary tylotes supporting tangential ectosomal
layer, the latter sometimes protruding through
surface, with ascending primary tracts of
choanosomal principal styles in turn supporting
these.
Choanosome. Choanosomal skeleton reticulate,
with multispicular ascending primary fibres and
paucispicular transverse connecting fibres; spongin fibres heavy, cored by principal styles and
auxiliary tylotes (the latter also scattered
throughout the mesohyl), and echinated by acanthostyles more-or-less perpendicular to fibres;
mesohyl matrix light, with numerous isochelae
scattered between fibres.
Megascleres (Table 4). Principal styles slender,
slightly curved near basal end, with abrupt hastate
points, and completely smooth.
Subectosomal auxiliary spicules tylotes or
quasitylotes, asymmetrical (and therefore
probably modified styles), usually with
microspined bases and points or sometimes completely smooth at both ends.
Echinating acanthostyles with spinose shaft,
base and point but apinose 'neck'; spines large,
recurved.
Microscleres (Table 4). Palmate isochelae of two
sizes, both with thickened and elongate alae.
Toxas are absent.
REMARKS. Dimensions of some spicules were
found to differ in type material (Table 4) from
those published by Hentschel (1911). Similarly,
two size classes of isochelae were found in the
WA population, not one as described by
Hentschel (1911). In both respects this population
is the same as the one described from Amirante
(Dendy, 1922) and the material described above
from Vietnam, whereas the specimen described
by Bergquist & Fromont (1988) from New
Zealand has substantially larger spicule dimensions than either of the Indian Ocean populations,
including only one size class of isochela. In
spicule geometry, choanosomal skeletal structure
and ectosomal characteristics (including the distribution of isochelae in the ectosomal
membrane), these 4 disjunct populations are relatively homogeneous and I follow Bergquist &
Fromont (1988) in recognising only a single
species. No intermediate populations of C. (C.)
chelifera are known, and the species is relatively
rare with only few known specimens in the Indowest Pacific.
This species is unusual to most C. (Clathria) in
possessing modified auxiliary spicules with
spines on both ends, considered by some authors
to be true tylotes typical of the Myxillidae
(Hall mann, 1920; Bergquist & Fromont, 1988) or
Iophonidae (Hajdu et al., 1994). But these
spicules are clearly asymmetrical (quasitylotes),
not true diactinal megascleres, and Dendy (1922)
correctly assigned this species (and thus the genus
Allocia) to Clathria. These modified quasidiactinal auxiliary spicules are infrequent but known
in several other microcionids (e.g., C. (C.) bulbosa, C. (Thalysias) major, C. (Dendrocia)
pyramida, C. (Wilsonella) australiensis, most
Echinochalina, some Holopsamma, and some
Echinoclathria species).
Clathria (Clathria) conectens (Hallmann, 1912)
(Figs 36-37, Plate 1B, Table 5)
Wilsonella conectens Hallmann, 1912: 245-247, p1.32,
fig.2, text-fig.50.
Clathria conectens; Hallmann, 1920: 768; Hooper &
Wiedenmayer, 1994: 259.
MATERIAL. LECTOTYPE: AMZ220: (dry) 16km E.
of Fraser I., Qld, 25°22'S, 153°07'E, 48-52m depth,
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
105
coll. FIV 'Endeavour' (dredge). PARALECTOTYPE:
AME1533 (dry; presently missing): same locality.
OTHER MATERIAL: QUEENSLAND - QMG300455, QMGL714 (fragment NTMZ1537), QMGL2757 (fragment NTMZ1564), QMGL2770 (fragment
NTMZ1581), QMG301037, QMG303190,
QMG303217, QMG304980, QMG304985,
QMG305135, QMG304005, QMG304016. NEW
SOUTH WALES - QMG301387.
HABITAT DISTRIBUTION. Acropora coral reef,
fringing rock reef, boulders, algal turf, wharf pylons;
inshore waters, 4-80m depth; Green I., S. Direction I.,
Innisfail (FNQ), Fraser I., Mudjimba I., Moreton I., N.
Stradbroke I., Moreton Bay (SEQ), and Byron Bay (N.
NSW) (16-28S) (Fig. 36G).
TABLES. Comparison between present and published
records of Clathria (Clathria) conectens (Hallmann).
All measurements are given in p,m, denoted as range
(and mean) of sp'cule length x spicule width (N=25).
SPICULE
Lectotype
(AMZ220)
Specimens (N=7)
Choanosomal
principal styles
944168.4)-268
x 2.5-(4.1)-5
1104171.4)-218
x 3-(4.3)-6
Subectosomal
auxiliary styles
92-(171.3)-219
x 1.542.9)-4
174-(230.6)-295
x 1.5-(2.4)-3.5
48-(58.3)-65
x 3-(3.7)-5
36-(64.2)-78
x 2-(3.9)-5
Echinating
acanthostyles
Chelae
Toxas
6-(7.2)-8
4.546.7)-9
22-(69.8)-111
51-(102.6)-164
x 0.5-(0.9)-1.5
x 0.5-(0.7)-1.0
DESCRIPTION. Shape. Massive, subcylindrical
mass 40-65mm high, 85-120mm broad, composed of irregularly reticulate, lamellate bulbous
branches, up to 15mm diameter, standing erect on
substrate.
Colour Live colouration bright orange-yellow to
bright orange-red (Munsell 2.5YR 7/10-10R
6/10), brown in ethanol.
Oscules. Small oscules, up to 1.5mm diameter, on
edges and tips surface bulbs.
Texture and surface characteristics. Surface uneven, porous, optically smooth.
Ectosome and subectosome. Ectosome with tangential or paratangential layer of thin subectosomal auxiliary subtylostyles, on or just below
a membraneous dermal layer; subectosomal
auxiliary styles confined entirely to peripheral
skeleton; subectosomal skeleton virtually nonexistent, with only few erect, plumose
choanosomal principal styles, arising from
peripheral choanosomal spongin fibres, projecting into tangential ectosomal layer.
Choanosome. Choanosomal skeleton irregularly
reticulate, with regular circular, oval or elongate
meshes enclosing small oval choanocyte chambers; spongin fibres relatively heavy, without size
differentiation of primary or secondary components, although ascending (primary) skeletal
fibres cored by paucispicular or multispicular
tracts of choanosomal styles, whereas connecting, transverse (secondary) fibres uni-, pauci- or
entirely aspicular; echinating acanthostyles
sparse, scattered evenly throughout skeleton, occasionally incorporated into fibres; mesohyl
matrix very light, some choanosomal styles scattered between fibres; some specimens also incorporating detritus into mesohyl, but not into fibres.
Megascleres (Table 5). Choanosomal principal
styles thin, fusiform, occasionally styloid, slight-
ly curved, with rounded or very slightly subtylote,
smooth bases.
Subectosomal auxiliary subtylostyles slightly
curved, sometimes sinuous, exceedingly thin,
hastate, almost vestigial, with very slightly subtylote, smooth bases.
Acanthostyles subtylote, with more-or-less
evenly distributed vestigial (granular) spination.
Microscleres (Table 5). Palmate isochelae small,
unmodified.
Toxas accolada rare, thin, with sharply angular
central curvature and straight arms.
REMARKS. Specimen AMZ220 is designated
lectotype (labelled `cotype of Wilsonella conectens, duplicate of E1533') as the latter specimen
is presently missing from AM collections.
Despite Hallmann's (1912) remarks to the contrary C. (C.) conectens is clearly different from
C. (C.) angulifera (see above), although both
species do fit into his (erroneous) concept of
Wilsonella. Moreover, in C. (C.) conectens the
megascleres which core fibres (choanosomal
styles) are differentiated from those occurring in
the dermal skeleton (subectosomal styles), and
although their geometry is very similar, they have
very different morphology (see Fig. 36) and
thickness (see Table 5). Each category of spicule
is localised in the choanosomal and ectosomal
regions, respectively, and they are not intermingled as supposed by Hallmann (1912). The
entire spiculation of this species is reduced, and
for that reason it is easily recognisable. This
species is a common member of the SolanderianPeronian biogeographical overlap zone centred
around Moreton Bay, Queensland.
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106zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
A
FIG. 36. Clathria (Clathria) conectens (Hallmann) (lectotype AMZ220). A, Choanosomal principal styles. B,
Subectosomal auxiliary style. C, Echinating acanthostyle. D, Accolada toxa. E, Palmate isochela. F, Section
through peripheral skeleton. G, Australian distribution. H, Lectotype.
Clathria (Clathria) crassa (Lendenfeld, 1887)
(Figs 38-39)
Antherochalina crassa Lendenfeld, 1887b: 787, p1.22,
fig.41.
Clathria crassa; Burton, 1934a: 558; Hooper &
Wiedenmayer, 1994: 259.
Microciona or Thalysias crassa; de Laubenfels, 1936a:
112.
Not Reniera crassa Carter, 1876: 312.
Not Aulena crassa; Lendenfeld, 1889a: 101.
MATERIAL. HOLOTYPE: BMNH1886.8.27.450
(fragments AMG3460, AMZ1991): Port Jackson,
NSW, 33 ° 51'S, 151 ° 16'E, other details unknown.
HABITAT DISTRIBUTION. Ecology unknown;
central coast NSW.
DESCRIPTION. Shape. Thin fan, 230mm high,
190mm maximum width, up to 8mm thick, with
short stalk, 25mm long, digitate margins.
Colour Live colouration unknown, grey-brown
in ethanol.
Oscules. Small pores seen on both faces of fan,
1-2mm diameter, with subdermal drainage canals
surrounding oscules.
Texture and surface characteristics. Surface
smooth, not optically hispid, with stellate
sculpturing on both faces of fan (associated with
aquiferous system); texture firm, flexible.
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU
107
FIG. 37. Clathria (Clathria) conectens (Hallmann) (A-B, lectotype AMZ220; C-G, QMG303217). A,
Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal fibres. D, Echinating acanthostyle. E,
Acanthostyle spines. F, Accolada toxa. G, Palmate isochelae.
108zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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0
0
FIG. 38. Clathria (Clathria) crassa (Lendenfeld) (fragment of holotype AMZ I 991). A, Choanosomal principal
style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Oxhorn toxas. E, Palmate
isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, holotype BMNH 1886.8.27.450.
^
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109
FIG. 39. Clathria (Clathria) crassa (Lendenfeld) (holotype BMNH1886.8.27.450). A, Fibre characteristics. B,
Ectosomal region. C, Choanosomal skeleton. D, Oxhorn toxas. E, Smaller toxas intermediate between oxhorn
and u-shaped. F, Palmate isochelae. G, Acanthostyle spines. H-1, Echinating acanthostyles.
110zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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Ectosome and subectosome. Ectosome microscopically hispid, with points of large
choanosomal principal styles from peripheral
fibres protruding through surface in plumose
brushes; thin tangential layer of subectosomal
auxiliary subtylostyles lying on or just below
surface, at base of protruding choanosomal
megascleres.
Choanosome. Choanosomal skeleton irregularly
reticulate, with slightly renieroid axis and
plumoreticulate extra-axis; spongin fibres in axis
flattened, very heavy, running longitudinally
through lamellae; axial fibres clearly divisible
into primary (longitudinal) and secondary (ascending, connecting) components; primary fibres
cored by paucispicular tracts of principal
choanosomal styles; secondary fibres
unispicular; extra-axial skeleton diverges from
axis at an oblique angle, with moderately heavy
fibres, divided into primary (multispicular, ascending) and secondary (unispicular, transverse)
elements; choanosomal principal styles project
from primary fibres in plumose tracts; secondary
fibres connect ascending primary lines, producing reneiroid reticulation, except at periphery
where architecture is distinctly plumose; echinating acanthostyles moderately common, evenly
distributed throughout skeleton; mesohyl matrix
abundant, containing few tnicroscleres but few
loose megascleres.
Megascleres. Choanosomal principal styles
thick, fusiform, slightly curved, with rounded or
slightly subtylote, smooth bases. Length 184(292.3)-463p,m, width 9-(17.3)-22p,m.
Subectosomal auxiliary subtylostyles thick,
straight, fusiform, with microspined subtylote
bases. Length 118-(226.7)-316p,m, width 3(4.8)-6 pun.
Acanthostyles short, thick, subtylote or
rounded, with evenly dispersed vestigial
(granular) spination. Length 51-(66.3)-82Rm,
width 5-(6.5)-9p.m.
Microscleres. Palmate isochelae large, unmodified. Length 17-(l9.5)-23p,m.
Toxas oxhorn, thick, with rounded central curvature, slightly reflexed points although the
smaller ones may lack reflexed points and are
intermediate between oxhorn and u-shaped
forms. Length 28-(68.0)-112p,m, width 0.842.6)4.511M.
REMARKS. Burton (1934a) designated A. crassa type species of Antherochalina, and subsequently declared that the genus was a synonym
of Clathria. Lendenfeld's (1887b) brief descrip-
tion of A. crassa is vague and not very useful in
distinguishing it from other Clathria, but type
material is still extant and recognisable. However, there is little agreement between characters
in the type material and as described by Lendenfeld, C. (C.) crassa is very closely related to C.
(C.) arcuophora, with similar skeletal architecture (with 2 components, renieroid and
plumoreticulate), spicule geometry, spicule sizes,
similar fibre characteristics and comparable
growth forms. It is possible that the two species
are synonyms, but their formal merger is not
warranted on the basis of the existing relatively
poor material. Similarly C. (C.) crassa shows
some similarities with C. (Isociella), particularly
to C. (I.) eccentrica. This resemblance is mostly
due to the renieroid axial skeletal architecture and
geometry of both principal and auxiliary styles.
Clathria (Clathria) decumbens Ridley, 1884
(Figs 40-41)
Cluihria decumbens Ridley, 1884a: 612, p1.53, fig.k,
p1.54, fig.g-g'; Ridley & Dendy, 1887: 148; Burton,
1938a: 29, p1.3, fig.23; Hooper & Wiedenmayer,
1994: 259.
Wilsonella decumbens; Hallmann, 1912: 239.
MATERIAL. HOLOTYPE: BMNH1882.10.17.51:
Boudouse and Etoile Is, Amirante Is Group, Indian
Ocean, 6 ° S, 53°10'E, coll. HMS 'Alert' (dredge).
PARATYPES: BMNH1882.10.17.71, 1882.10.17.76:
same locality. OTHER MATERIAL: QUEENSLAND BMNH1887.5.2.139.
HABITAT DISTRIBUTION. 6-26m depth; on sand
and coral rubble substrate; Cape York, Torres Strait
(FNQ) (Fig. 40F) (Ridley, 1884a). W. Indian Ocean
(Ridley & Dendy, 1887), Madras (Burton, 1938a).
DESCRIPTION. Shape. Small, subcylindrical,
irregularly lobate, 32-50mm long, 15-40mm
wide, up to 25mm thick.
Colour Brown to red-brown in ethanol.
Oscules. Numerous small oscules, 1-2mm
diameter, dispersed between surface conules.
Texture and surface characteristics. Surface
rugose, irregularly conulose, with canals, grooves
and ridges meandering over the surface.
Ectosome and subectosome. Ectosome
membraneous between ridges and surface projections, with spongin fibres from choanosome
producing a dermal reticulation; dermal region
lacks a mineral skeleton entirely; spongin fibres
in subectosomal region closely reticulate, with
relatively small mesh sizes.
Choanosome. Choanosomal skeletal architecture
regularly to irregularly reticulate; spongin fibres
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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111
moderately heavy, undifferentiated into primary
or secondary lines, lightly cored by paucispicular
tracts of both choanosomal principal styles and
subectosomal auxiliary styles; heavily echinated
by acanthostyles; coring spicules in peripheral
fibres ascend to surface, piercing surface processes in light brushes; subectosomal auxiliary subtylostyles also sparsely scattered within mesohyl;
fibre meshes in choanosomal skeleton cavernous;
mesohyl matrix abundant, lightly pigmented.
Megascleres. Choanosomal principal styles
straight or slightly curved near base, hastate
points, with evenly rounded bases, smooth or
with microspines on both points and bases.
Length 128-(156)-176Rm, width 4-(4.9)-6p.m.
Subectosomal auxiliary subtylostyles straight,
with hastate points, slightly subtylote bases,
bases smooth or occasionally microspined.
Length 159-(1177.8)-193Rm, width 243.7)6 Km.
Acanthostyles subtylote, with strongly formed,
recurved spines over apical end, shaft and base
but bare neck. Length 62-(83.2)-104Rm, width
Microscleres. Palmate isochelae unmodified, two
discrete size classes, the smaller showing variation in fusion of alae. I: Length 14-(20.4)-28Km,
II: length 5-(7.4)-10Km.
Toxas absent.
REMARKS. Hallmann (1912) referred this
species to Wilsonella, presumably because Ridley (1884a) did not differentiate between
choanosomal (principal) and subectosomal
(auxiliary) megascleres. Subsequently, Burton
(1938a) described additional specimens from
Madras which had differentiated principal and
auxiliary spicules (but were otherwise identical
with Ridley's (1884) description). In type
material there were consistent morphological differences between principal and auxiliary
spicules, confirming that the species should be
retained in Clathria (Clathria).
Although C. (C.) decumbens has been described
from three widely separated localities, it remains
poorly known, being only poorly differentiated
from other low growing, lobate Clathria species.
The species is similar to C. (Thalysias) major
(with spines on both the bases and points of some
of its megascleres), while at the same time being
similar to species in the juniperind species complex (i.e., Clathria in which the geometry of
choanosomal principal and subectosomal
auxiliary spicules is barely different), but the
species has little else of distinction.
Clathria (Clathria) echinonematissima
(Carter, 1881)
Wilsonella echinonematissima Carter, 1881a: 366;
Carter, 1887: 210; Hallmann, 1912: 243.
Clathria echinonematissima; Dendy, 1896: 33, 34;
Hooper & Wiedenmayer, 1994: 259.
MATERIAL. HOLOTYPE: BMNH not found (slide
containing only a desilicified section is the only type
material known to exist): Westernport Bay, 38 °26'S,
145°08'E, or Port Phillip, Vic, 38 °09'S, 144°52'E, coll.
J.B. Wilson (dredge).
HABITAT DISTRIBUTION. Ecology unknown; Victoria.
DESCRIPTION. Shape. Massive.
Colour. Unknown.
Oscules. Unknown.
Texture and surface characteristics. Unknown.
Ectosome and subectosome. Ectosomal skeletal
tracts heavily cored with detritus, megascleres
excluded.
Choanosome. Choanosomal skeleton irregularly
reticulate, with relatively heavy spongin fibres;
fibres of peripheral skeleton are solely
arenaceous, whereas within choanosome fibres
cored by subectosomal auxiliary styles; echinating acanthostyles dispersed throughout skeleton.
Megascleres. Choanosomal principal megascleres apparently absent.
Subectosomal auxiliary subtylostyles with
smooth bases. Length 21011m, width 4t.t.m.
Acanthostyles apparently divided into two size
categories. Length from 145Km, width 8.311m.
Microscleres. Isochelae arcuate. Length 251.1,m.
Toxas absent.
REMARKS. This species is barely recognisable
as Clathria from Carter's (1881) description, and
it is only poorly differentiated from other
microcionid species. One category of auxiliary
spicule and the absence of choanosomal principal
spicules places it in Wilsonella (sensu Hallmann,
1912). It also was described with arcuate
isochelae, similar to species grouped by
Hallmann (1920) in Paradoryx, but this is uncorroborated. Clathria echinonematissima is a
species inquirendum as it is only known from a
slide preparation, now desilicified, allegedly
made from the missing holotype.
Clathria (Clathria) hispidula (Ridley, 1884)
(Figs 42-43)
Amphdectus hispidulus Ridley, 1884a: 429-430, p1.40,
fig.c, p1.41, fig.y.
112zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 40. Clathria (Clathria) decumbens Ridley (paratype BMNH1882.10.17.71). A, Choanosomal principal
styles. B, Subectosomal auxiliary styles/subtylostyles. C, Echinating acanthostyles. D, Palmate isochelae. E,
Section through peripheral skeleton. F, Australian distribution. G, Holotype BMNH1882.10.17.51. H, Paratype.
Axociella hispidulus; de Laubenfels, 1936a: 114.
Esperiopsis hispidula var. ramosa; Hentschel, 1911:
313.
Not Hymeraphia hispidula Topsent, 1904a: 164-165,
p1.14, fig.2.
MATERIAL. LECTOTYPE: BMNHI881.10.21.261:
Thursday I., Torres Strait, N. Old, 10 °35'S, 142 ° 13'E,
6-10m depth. 01.vi.1881, coll. HMS 'Alert' (dredge).
PARALECTOTYPE: BMNH1881.10.21.319: same
locality (dry). OTHER MATERIAL: WESTERN
AUSTRALIA - ZMB4408.
HABITAT DISTRIBUTION. Encrusting on bivalves,
hydroids, and gorgonians and algae; 6-11rn depth;
Torres Strait (FNQ), and Shark Bay (WA) (Fig. 42G).
DESCRIPTION. Shape. Erect, irregular branch-
ing reticulate mass of clathrous digits, up to
60mm long, 55mm diameter.
Colour. Live colour unknown, light brown in
ethanol.
Oscules. Small oscules, <2mm diameter in
preserved material, scattered over lateral margin.
Texture and surface characteristics. Harsh, compressible, slightly elastic; surface with meandering irregular ridges and microconules scattered
over branches, and tips of fibres from primary
skeleton protruding.
Ectosome and subectosome. Sparse paratangential skeleton of subectosomal auxiliary subtylos-
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113
FIG. 41. Clathria (Clathria) decumbens Ridley (paratype BMNH1882.10.17.71). A, Choanosomal skeleton. B,
Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E-F, Terminations of principal styles.
G, Principal styles. H-K, Palmate isochelae.
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 42. Clathria (Clathria) hispidula (Ridley) (lectotype BMNH1881.10.21.261). A, Choanosomal principal
styles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Wing-shaped toxas. E, Palmate
isochelae. F, Section through peripheral skeleton. G, Australian distribution.
tyles forming bundles or lying more-or-less erect
on the surface, particularly on ends of surface
conules; tips of conules with choanosomal principal styles also protruding only short distance
through surface; choanosomal fibres immediately below surface skeleton with poorly
developed subectosomal region.
Choanosome. Skeleton regularly reticulate,
slightly renieroid, with thin but well developed
spongin fibres forming oval or rectangular, relatively wide meshes, 150-350Rm diameter,
generally more cavernous in axis than in
peripheral region; spongin fibres 20-701J,m
diameter, imperfectly divided into primary, ascending, multispicular tracts of 4-10 spicules per
tract, interconnected by uni-, pauci- or aspicular
secondary transverse tracts; fibres cored by
choanosomal principal styles not occupying en-
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
115
Clat hria ( Clat hria) hispidula (Ridley) (lectotype BMNH1881.10.21.261). A, Choanosomal skeleton.
FIG. 43. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spination. E, Terminations of auxiliary
spicules. F, Palmate isochelae. G, Wing-shaped toxa.
MEMOIRS OF THE QUEENSLAND MUSEUM
116zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
tire fibre diameter, and sparsely echinated by
slender acanthostyles; mesohyl matrix light containing scattered microscleres and some auxiliary
megascleres.
Megascleres. Choanosomal principal styles
slender, straight or slightly curved midway along
shaft, with evenly rounded smooth bases and
fusiform points. Length 884162.7)-21911m,
width 4-(4.8)-611m.
Subectosomal auxiliary subtylostyles similar in
geometry to principal spicules, but more slender
and with subtylote bases; thinner (younger) principal spicules frequently sinuous. Length 94(143.0)-175pAn, width 2-(3.4)-5p.m.
Echinating acanthostyles club-shaped, slender,
slightly subtylote, evenly spines or with bare
'neck' below base, fusiform or rounded points,
granular spination. Length 52-(59.4)-65Rm,
width 2-(4.1)-5p.m.
Microscleres. Palmate isochelae very abundant,
small, unmodified, with lateral alae completely
fused to shaft and front ala entire; lateral and front
alae of approximately equal length. Length 9(11.6)-13p,m.
Toxas uncommon, small, thick, wing-shaped,
with rounded central curve and slightly reflexed
arms. Length 17-(56.8)-104Rm, width 1.5-(2.1)3p.m.
REMARKS. This species has not yet been rediscovered from either of the known locations of
early collections, and it remains relatively poorly
known from museum specimens. Surprisingly,
both Ridley (1884a) and Hentschel (1911) failed
to describe several spicule types present in their
respective material, particularly echinating acanthostyles which are uncommon but certainly
present, as well as auxiliary subtylostyles and less
common toxa microscleres. Furthermore,
Hentschel's material essentially differs from
Ridley's only in the specific dimensions of
spicules and growth form (being more elongate,
branching), and it is not considered to be necessary to recognise the subspecific taxon proposed
by Hentschel (1911) for the WA population.
Although the identity of this species has never
been been clearly established from either published record, it is obviously a Clathria with relatively cavernous skeletal architecture and standard
spiculation. It is similar to C. (C.) angulifera
(Dendy) from Victoria and southern Queensland,
and C. (T) aphylla from the Houtman Abrolhos,
in having a cavernous, slightly renieroid skeletal
structure ('angulifera' species group), differing
in the protruding fibrous ectosomal skeleton,
spicule geometries (e.g., toxas, acanthostyles), a
more-or-less branching growth form (cf. lobate
lamellate and foliose lamellate, respectively), and
spicule sizes.
Clathria (Clathria) inanchorata Ridley &
Dendy, 1886 (Figs 44-45, Table 6)
Clathria inanchorata Ridley & Dendy, 1886: 475;
Ridley & Dendy, 1887: 150, p1.28, fig.4, p1.29,
fig. 13; cf. Kieschnick, 1896: 533; cf. Thiele, 1903a:
959; Whitelegge, 1907: 492-495; Hallmann, 1912:
206, 211, 214, 215; Hooper & Wiedenmayer, 1994:
259.
Pseudanchinoe inanchorata; de Laubenfels, 1936a:
109.
cf. Microciona prolifera; Vosmaer, 1935a: 610, 635,
665.
MATERIAL. HOLOTYPE: BMNH1887.5.2.99: Bass
Strait, Tas, 36°59'S, 150°20'E, 4.iv.1874, coll. HMS
'Challenger' (trawl). OTHER MATERIAL: NSW AMG5675, AMZ131, AMZ1413, AMZ1414.
HABITAT DISTRIBUTION. Depth 110-300m, substrate mud; Bass Strait (Tas) (Ridley & Dendy, 1886),
S. coast (NSW) (Whitelegge, 1907). Ternate, Mollucas, Indonesia (Kieschnick, 1896).
DESCRIPTION. Shape. Erect, irregularly
cylindrical digits, 38-120mm high, 8-12mm
diameter, encrusting on organic debris or standing free in substrate.
Colour Grey- or yellow-brown preserved.
Oscules. Numerous oscules, up to 2.5mm
diameter, scattered over branches.
Texture and surface characteristics. Surface
rugose, reticulate, minutely hispid; texture firm,
flexible.
Ectosome and subectosome. Ectosomal skeleton
prominently hispid, with choanosomal principal
megascleres from peripheral fibres protruding a
long way through surface, and with a sparse tangential layer of subectosomal auxiliary
megascleres dispersed between erect principal
spicule brushes; subectosomal skeleton plumose,
undifferentiated from choanosomal fibres which
are immediately subdermal.
Choanosome. Choanosomal skeleton irregularly
reticulate, with relatively heavy spongin fibres
incompletely divided into primary ascending and
secondary transverse components; secondary
fibres uncored, lightly echinated by small acanthostyles; primary fibres contain sparse tracts of
subectosomal auxiliary subtylostyles, identical to
those occurring in the ectosomal skeleton,
enclosed completely within spongin fibres,
together with plumose brushes of choanosomal
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
117
FIG. 44. Clathria (Clathria) inanchorata Ridley & Dendy (holotype BMNH1887.5.2.99). A, Choanosomal
principal subtylostyles. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Accolada toxa.
E, Oxhorn toxas. F, Section through peripheral skeleton. G, Australian distribution. H, Holotype.
118zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 45. Clathria (Clathria) inanchorata Ridley & Dendy (holotype BMNH1887.5.2.99). A, Choanosomal
skeleton. B, Ectosomal spicule bundles. C, Ectosomal skeleton. D, Fibre characteristics. E, Echinating
acanthostyle. F, Smaller echinating acanthostyle. G, Acanthostyle spination. H-I, Ozhorn toxas. J, Portion of
accolada toxa.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
^
119
TABLE 6.Comparison between present and published
records of Clathria (Clathria) inanchorata Ridley &
Dendy. All measurements are given in pm, denoted
as range (and mean) of spicule length x spicule width
(N=25).
SPICULE
Choanosomal
principal styles
Subectosomal
auxiliary styles
Echinating
acanthostyles I
Echinating
acanthostyles II
Chelae
Toxas I
Toxas II
Holotype
(BMNH1887.5.2.99)
Specimens (N=4)
292-(417.6)-535 x 8-
273-(384.5)-540x
(18.4)-28
12-(17.8)-25
2524328.6)-432 x 3- 290-(341.8)-410 x 3(8.2)-11
(8.5)-12
58-(66.4)-78 x 3(5.2)-7
58-(75.2)-88 x 4(5.4)-7
1184150.2)-175 x 4- 119-(169.3)-228 x 6(7.4)-11
(8.8)-12
absent
absent
35-(73.8)-121 x 1.5-
22-(66.2)-105 x 1.5-
(2.0)-3
(3.4)-5
118-(349.2)-478 x
3044408.3)-545 x
1.5-(2.1)-2.5
1.5-(2.1)-3
principal styles usually poking out of fibres;
choanosomal principal styles, protruding through
spongin fibres, together form multispicular ascending plumose tracts, also lightly echinated by
acanthostyles; fibre anastomoses form circular to
oval, cavernous meshes; mesohyl matrix very
light, with few megascleres dispersed between
fibres.
Megascleres (Table 6). Choanosomal principal
styles long, thick, fusiform, slightly curved, with
rounded or slightly subtylote, smooth or minutely
microspined bases.
Subectosomal auxiliary subtylostyles straight,
relatively thick, robust, almost hastate, with
slightly subtylote microspined bases.
Acanthostyles very variable in size, with slightly subtylote bases, incompletely separated into
two size classes with some intermediate examples. Smaller morph usually straight, often
with aspinose necks, whereas larger morphs
slightly curved, with evenly distributed large
spines.
Microscleres. (Table 6). Isochelae absent.
Toxas clearly separated into two morphs - I: most
common form are oxhom toxas, small, relatively
thick, with large, rounded or slightly angular curvature at centre, and reflexed points. II: Less
frequent are accolada toxas, long, thin, sharply
angular at centre, unreflexed arms.
REMARKS. This species has a distinctive
growth form, spicule geometry, and spongin fibre
characteristics, but otherwise it is similar to other
species included in Hallmann's (1912) spicata'
group, particularly C. (T) costifera and C. (C.)
caelata. There is no doubt that Whitelegge's
(1907) specimens from Wollongong are conspecific although this claim was disputed by
Hallmann (1912: 206). Kiescknick's (1896)
record of this species from Indonesia is dubious,
since his material was not described and could
possibly be any one of these spicata' -like sponges. Thiele (1903a) compared Kieschnick's
specimen with C. (T)coralliophila from the same
region, but that comparison is misleading: both
taxa have quite different spiculation and spongin
fibre characteristics.
Clathria (Clathria) kylista Hooper & Levi, 1993
(Figs 46-47, Plate IC, Table 7)
Clathria (Clathria) kylista Hooper & Levi, 1993a:
1265-1267, figs 21-22, table 11; Hooper & Wiedenmayer, 1994: 259.
MATERIAL. HOLOTYPE: QMG300035: Inner
Gneering Shoals, off Mooloolaba, Qld., 26°38.5'S,
153°09.5'E, 10m depth, 10.xii.I 991, coll. J.N.A.
Hooper & S.D. Cook (SCUBA). PARATYPE:
QMG300690 (ORSTOM R1338; fragment
NTMZ3876): N. entrance, Recif des Cinq Milles, SW.
New Caledonia lagoon, 22°29.3'S, 166°44.4'E, 8m
depth, 30.iv.1976, coll. G. Bargibant (SCUBA).
OTHER MATERIAL: QUEENSLAND - QMG303166.
HABITAT DISTRIBUTION. 8-20m depth; on rock
pinnacles, in caves and coral rubble substrates;
Mooloolaba and Moreton I. (SEQ) (Fig. 46F). New
Caledonia (Hooper & Levi, 1993a).
DESCRIPTION. (See Hooper & Levi, 1993a).
DIAGNOSIS. (refer to Table 7 for spicule dimensions) Simple digitate, tubular or bulbous, erect,
branching growth form; dark orange to pale
orange alive; terminal osculum on single tubes
and oscules scattered over the apical regions of
more complex lobate digitate growth forms; oscules surrounded by membraneous lip;
prominently conulose surface with large, bulbous
tubercles; ectosome membraneous, with sparse,
tangential layer of subectosomal auxiliary styles
and erect brushes of auxiliary spicules scattered
throughout both peripheral and subectosomal
regions; choanosomal skeleton plumoreticulate
with differentiated primary and secondary fibres;
primary fibres ascending, paucispicular occupying only small proportion of fibre diameter, cored
by principal spicules with fewer auxiliary
megascleres interdispersed; secondary fibres
transverse, uni- or aspicular; echinating styles
MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVUT
120^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
B
A
C
E
1.0
('.1
E
azyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO
0
0
FIG. 46. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hr ia ( Clat hr ia) ky list a Hooper & Levi (A-D, QMG303166; E, holotype QMG300035). A,
Choanosomal principal style. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada
toxas. E, Section through peripheral skeleton. F, Australian distribution.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
121
FIG. 47. Clathria (Clathria) kylista Hooper & Levi (paratype QMG300690); A, Choanosomal skeleton. B, Fibre
characteristics in peripheral skeleton. C-D, Echinating acanthostyles. E, Acanthostyle vestigial spines. F,
Subtylote base of auxiliary subtylostyles.
122zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 7. Comparison in spicule dimensions between
Clathria (Clathria) lipochela Burton, 1932
types and specimen of zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hria ( Clat hria) kylist a
(Fig. 48)
Hooper & Levi. All measurements are given in p.m,
denoted as range (and mean) of spicule length x
spicule width (N=25).
Burton, 1932a: 319, figs 6-7, textfig.29; Burton, 1940: 109, p1.4, fig.5; Koltun, 1964a:
69; Desqueyroux, 1972: 26-27, figs 87-89, 135;
Sara, 1978: 65; Hooper & Wiedenmayer, 1994: 260.
liolotype
Paratype
Specimen
Thalyseurypon lipochela; de Laubenfels, 1936a: 107.
(QMG300035) (QMG300690) (QMG303166)
SPICULEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
(SE. Qld)
Choanosomal 64484.7)-112
principal
x 2.5-(2.9)-4.0
styles
(New Cal.)
(SE. Qld)
61-(81.6)-97 x
1.6-(2.7)-3.6
55478.2)-104
x 2.043.1)-4.5
Subectosomal
auxiliary
styles
143(168.9)181 x 1.0(2.4)-4.1
138(159.4)183 x0.4(2.1)-3.5
110(142.3)166x 1.0(1.8)-3.5
Ectosomal aux
-iliary styles
absent
absent
absent
32-(35.5)-39 x
1.1-(3.1)-5.0
18-(27.8)-35 x
1.0-(2.4)-3.5
26-(32.8)-40 x
Echinating
acanthostyles 1.042.6)-4.0
Toxas
65-(129.6)231.4 x 0.5(1.3)-2.5
65-(135.6)266 x 0.2(1.1)-1.6
35-( 146.5)222 x 0.5(1.0)-2.0
Chelae
absent
absent
absent
moderately common on primary ascending
fibres, sparse on secondary connecting fibres;
choanosomal principal styles slender, straight,
relatively short, with hastate points and smooth,
slightly swollen, subtylote bases; subectosomal
auxiliary styles long, slender, straight, with hastate points and smooth, elongated, swollen subtylote bases; echinating styles entirely smooth,
short, slender, sharply pointed, with prominent
basal constriction ('neck'), subtylote base and
widest just below basal constriction; isochelae
absent; accolada toxas abundant, moderately
long, ranging from slender to raphidiform, with
straight arms and prominent central curve.
REMARKS. This species was assigned to
Clat hria ( Clat hria) by Hooper & Levi (1993a),
even though echinating spicules are smooth (cf.
Echinoclat hria) , because coring (principal) and
echinating megascleres have different geometries
(see also C. (M.) acerat oobt usa) . Clat hria ( C.)
kylist a is similar to C. ( C.) angulifera and C. (C.)
noarlungae in skeletal structure, having sparsely
cored ascending primary fibres and uncored
secondary connecting fibres, although both these
other species have acanthose echinating spicules
and palmate isochelae, and C. (C.) noarlungae
also has slightly curved toxas. The unusual
geometry of the echinating styles in C. (C.)kylista
is the strongest apomorphy for the species, not
seen elsewhere in the genus.
Clat hria lipochela
MATERIAL. HOLOTYPE: BMNH1928.2.15.352: Eddystone Rock, Falkland Is, 105-115m depth, coll. HMS
'Discovery' (trawl).
HABITAT DISTRIBUTION. 22-1I5m depth; on sand
and hard substrates; King George Land (Australian
Antarctic Territory), Antarctica (Koltun, 1964a) (Fig.
48F). South Georgia (Koltun, 1964a), Falkland Is (Burton, 1932a), Caleta Santa Marta, Chile (Desqueyroux,
1972), Mar del Plata, Argentina (Burton, 1940), Kerguelen Is (Koltun, 1964a), Magellan Straits (Burton,
1940), Cape Sebastiano, Punta Arenas, Rio Grande,
Cape Domingo, Cape Viamonte, Tierra del Fuego
(Sara, 1978).
DESCRIPTION. Shape. Stalked, irregularly
flabellate; digitate margins of fan.
Colour. Unknown.
Oscules. Not seen.
Text ure and surface charact erist ics. Firm, compressible; uneven, pitted, porous surface.
Ect osom e and subect osom e. Surface skeleton
with sparse paratangential or erect brushes of
subectosomal auxiliary styles, mainly at ends of
ascending primary spicule tracts; choanosomal
principal styles protrude through surface in
places; detritus scattered over surface but not
embedded in ectosome.
Choanosom e. Skeletal architecture regularly
reticulate with ascending primary multispicular
fibres and transverse uni-, pauci- or occasionally
aspicular fibres, interconnecting at more-or-less
regular intervals producing rectangular or elongate meshes, up to 4501.1.m diameter; fibres cored
by choanosomal principal styles and moderately
heavily echinated by acanthostyles evenly scattered over fibres in both axial and peripheral
regions of skeleton; spongin fibres well
developed; few spicules scattered between fibres;
mesohyl matrix light, smooth, virtually unpigmented.
Megascleres. Choanosomal principal styles
short, relatively slender, fusiform, rounded and
smooth bases, straight or only slightly curved
near basal end. Length 153-(206.7)-254Rm,
width 7-(11.0)-14p.m.
Subectosomal auxiliary styles long, slender,
slightly hastate pointed, slightly subtylote or
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
123
FIG. 48. Clathria (Clathria) lipochela Burton (holotype BMNH1929.2.15.352). A, Choanosomal principal style.
B, Subectosomal auxiliary subtylostyle and style. C, Echinating acanthostyle. D, Palmate isochelae. E, Section
through peripheral skeleton. F, Antarctic distribution. G, Choanosomal skeletal structure. H, Fibre characteristics.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
124zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
rounded bases, microspined or smooth bases.
Length 164-(179.4)-198p,m, width 3-(3.9)-5p.m.
Echinating acanthostyles relatively long,
slender, straight, with subtylote bases, fusiform
points, spines concentrated in basal and apical
regions and bare 'necks'. Length 79493.1)111p.m, width 6-(8.7)-1211m.
Microscleres. Palmate isochelae small, unmodified, relatively abundant. Length 7-(8.5)11p,m.
Toxas absent.
TABLE 8. Comparison between present and published
records of Clathria (Clathria)multipes Hallmann. All
measurements are given in p,m, denoted as range (and
mean) of sp'cule length x spicule width (N=25).
Lectotype
Paralectotype
Paralectotype
(BMNH1887.4.
SPICULEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP
(AMG9038)
(AMG9162)
27.9)
Citoaposomal 142-(136.6)-174
principal
styles
Subectosomal
auxiliary
styles
I45-(157.6)-178
112-(147.8)-262
x 4-(5.9)-7
x 648.2)-10
x 7-(9.3)- 12
162-(194.6)-239
x I.5-(2.8)-4
1324167.0)-222
x 2-(3.1)-4
143-(189.5)-262
x 2-(3.2)-4.5
REMARKS. Burton (1932a) named this species
52462.8)-91 x 71-(85.8)-98 x 64-(78.5)-93 x
Echinating
for the apparent absence of chelae, but these were
acanthostyles
4-(6.6)-9
4-(5.6)-8
5-(6.4)-8
found to be common in sections of the holotype.
5-(4.8)-8
4-(6.1)-8
3-(5.7)-8
Chelae
Burton's (1932a) comparison with Raspaxilla
16-(112.2)-147 19-(101.4)-141 1304124.4)-178
phakellina Topsent (Hooper, 1991: 1199), is misToxas
x 1-(4.3)-6
x 2-(4.5)-6
x I-(3.9)-5
leading as they do not resemble each other in
skeletal structure or spicule geometry. Clathria
(C.) lipochela resembles C. (T) vulpina Colour. Live colouration brick red, grey-brown
(Lamarck) in geometry of principal and echinat- dry.
ing spicules and skeletal architecture to some Oscules. Numerous large oscules, up to 3mm
extent, although they differ in many other fea- diameter, distributed over all surfaces.
Texture and surface characteristics. Surface irtures.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
regularly rugose, with small elevated conules
scattered mainly on lateral sides of branches.
Clathria (Clathria) multipes Hallmann, 1912
Ectosome and subectosome. Ectosome, rarely in(Figs 49-50, Table 8)
tact in dry type material, consists of three dimenClathria (Plectispa) arborea, in part, Whitelegge,
sional fibre reticulation, with fibre endings
1901: 88, p1.11, fig.15.
forming small surface conules, and choanosomal
Plectispa macropora,in part, Lendenfeld, 1888: 225-6.
principal styles protruding through peripheral
Not Plectispa arborea Lendenfeld, 1888: 226.
fibres in light brushes or singly; subectosomal
Clathria (Plectispa) multipes Hallmann, 1912: 204,
auxiliary subtylostyles dispersed in a tangential
211.
layer around projecting dermal fibres.
Clathria multipes; Hooper & Wiedenmayer, 1994:
Choanosome. Choanosomal skeletal architecture
260.
irregularly reticulate, consisting of relatively
MATERIAL. LECTOTYPE: AMG9038(dry): Tug- heavy spongin fibres forming incompletely difgerah Beach, Illawarra region, NSW, 34 ° 32'S, ferentiated primary (vaguely ascending, multi150°50'E (beach debris, label 'Clathria arborea Len- spicular) and secondary fibres (transverse uni-,
denfeld; ms name = Clathria plicatella'). PARALECpauci- or aspicular), and producing relatively
TOTYPES: AMG9162 (dry): Maroubra Bay, NSW,
33°455, 151°20'E (label Tlectispa arborea Lend. = tight oval to elongate meshes; fibres echinated by
Clathria arborea'). BMNH1887.4.27.9 (fragment small, sparsely distributed acanthostyles;
AMG3590): Port Jackson, NSW, 33 ° 51'S, 151°16'E mesohyl matrix light, with moderate quantities of
(label `Thalassodendron reticulata RvL, MS'). Other subectosomal subtylostyles and microscleres distype fragments ZMB2264, 6894. SYNTYPE of P. persed.
macropora: BMNH1925.11.1.555: Manly Beach,
Megascleres (Table 8). Choanosomal principal
NSW, 33°50'S, 151°17'E, other details unknown.
styles hastate or stepped, relatively thick, slightly
HABITAT DISTRIBUTION. Ecology unknown; curved, with tapering or slightly subtylote and
smooth bases.
central and S coast (NSW) (Fig. 49F).
Subectosomal auxiliary subtylostyles long,
DESCRIPTION. Shape. Branching, reticulate thin, fusiform, straight, curved or sinuous, with
branches, planar, 85-140mm long, 62-73mm slightly subtylote, smooth bases.
wide, with compressed, cylindrical and regularly
Acanthostyles subtylote, with vestigial spinaanastomosing branches,4-8mm diameter, and tion and an aspinose neck.
several small basal stalks (multiple points of at- Microscleres (refer to Table 8 for dimensions).
tachment).
Palmate isochelae minute.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
125
FIG. 49. Clathria (Clathria) multipes Hallmann (lectotype AMG9038). A, Choanosomal principal styles. B,
Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Palmate isochelae. E, U-shaped toxas. F,
Australian distribution. G, Section through peripheral skeleton. H, Paralectotype AMG9162. I, Lectotype.
Toxas u-shaped, relatively thick, oxeote, with
hastate points, typically curved at right angles at
the centre, with straight and unreflexed points.
REMARKS. All known specimens are in poor
condition, and it is not possible to accurately
determine ectosomal characteristics. The species
is most closely related to Clathria (Clathria)
rather than Clathria (Thalysias). Whitelegge
(1901) remarked that the species was frequently
washed up onto coastal beaches of S NSW after
storms, inferring that it was a relatively
prominent component of the benthos, and therefore it is surprising that it has not been collected
since that time despite intensive trawling. The
growth form of C. (C.) multipes (reminiscent of
MEMOIRS OF THE QUEENSLAND MUSEUM
126^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 50. Clathria (Clathria) multipes Hallmann (lectotype AMG9038). A, Choanosomal skeleton. B, Fibre
characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F,U-shaped toxas. G, Palmate isochelae.
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EI
tO
FIG. 51. Clathria (Clathria) murphyi sp.nov. (holotype QMG300656). A, Choanosomal principal subtylostyles.
B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada toxas. E, Palmate isochelae.
F, Section through peripheral skeleton. G, Australian distribution. H, Holotype in situ.
C. (T) coppingeri and Echinodictyum cancellatum (Raspailiidae)), the small size of isochelae,
the peculiar angular shapes of toxas, and the
vestigial acanthostyles differentiates this species
from other Clathria.
Clathria (Clathria) murphyi sp. nov.
(Figs 51-52, Plate 1D)
MATERIAL. HOLOTYPE: QMG300656 (NCIQ66C2904-N, fragment NTMZ3754): Old jetty, E. end of
Princess Royal Drive, Albany, WA, 35°02.3'S,
117°54.2'E, 9m depth, 27.ii.1989, coll. NCI (SCUBA).
HABITAT DISTRIBUTION. Wood jetty piles; 9m
depth; SW WA (Fig. 51G).
DESCRIPTION. Shape. Thickly encrusting, bulbous digitate lumps, up to 80mm diameter,
resembling the tropical Higginsia massalis (Desmoxyidae).
Colour. Orange-red alive (Munsell lOR 6/10),
pale brown preserved.
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FIG. 52. Clathria (Clathria) murphyi sp.nov. (holotype QMG300656). A, Choanosomal skeleton. B, Fibre
characteristics. C, Echinating acanthostyles. D, Base of acanthostyle. E, Acanthostyle spines. F, Palmate
isochelae. G, Accolada toxas.
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129
Oscules. Large oscules, more than 5mm diameter,
on ends of bulbs.
Texture and surface characteristics. Surface
porous, microvillose, uneven, with epi- and
zoophytes; texture soft, compressible, easily torn
off wooden piles.
Ectosome and subectosome. Ectosome
membraneous, with heavy, brown collagen and
brushes of choanosomal principal styles poking
through surface in plumose bundles, mainly on
ends of microconules; subectosomal auxiliary
styles in irregular, paratangential bundles below
surface, at base of principal style brushes, not
protruding through surface.
Choanosome. Choanosomal skeleton renieroid
reticulate, cavernous, without visible spongin
fibres, with differentiated primary and secondary
spicule tracts; primary skeletal tracts ascending,
multispicular, 20-40p,m diameter, composed of
plumose brushes of choanosomal principal styles,
3-10 spicules abreast; secondary tracts uni-, bi- or
paucispicular, transverse, 8-20p,m diameter, connecting primary tracts; echinating acanthostyles
relatively sparsely dispersed throughout
choanosome; mesohyl matrix heavy, granular,
darkly pigmented, surrounding large oval, paired
choanocyte chambers, up to 320Rm diameter,
with abundant toxa and isochelae microscleres
dispersed throughout.
Megascleres. Choanosomal principal subtylostyles short, stout, straight, fusiform points, tapering smooth bases, terminally subtylote, with
slightly swollen subterminal region. Length 87(116.5)-149pin 1, width 5-(7.1)-911m.
Subectosomal auxiliary subtylostyles short,
slender, fusiform, slightly subtylote, smooth
bases. Length 114-(138.4)-165p..m, width 1.5-
Echinating acanthostyles short, slender, slightly subtylote bases, spines recurved, evenly spined
except for bare neck. Length 42455.7)-6811m,
width 3.5-(4.2)-5.5vm.
Microscleres. Palmate isochelae, small, poorly
silicified, about 10% with twisted shafts. Length
5-(8.4)-14p.m.
Toxas accolada form, long, very slender hairlike, with straight, unreflexed arms and angular
central curvature. Length 72-( 115.6)-16411m,
width 0.5-(0.7)-0.4,m.
ETYMOLOGY. For Dr Peter Murphy, Australian Institute of Marine Science, Townsville.
REMARKS. This species is separated from other
Clathria (Clathria) by its distinctive spicule
geometries (principal styles with marked basal
constrictions and swollen 'necks'), relatively
small, poorly silicified spicules of all categories,
renieroid choanosomal skeletal structure (with
multispicular ascending tracts and uni- or
paucispicular plumose transverse connecting
spicule tracts, both sparsely echinated by acanthostyles), bulbous-digitate growth form, orangered colouration, microvillose surface with
plumose brushes of choanosomal styles protruding through the ectosome especially on the tips of
microconules. Clathria (C.)inurphyi has a skeletal architecture reminscent of C. (C.) arcuophora
and C. (C.) crassa (both of which have much
larger spicules of different geometry), and in this
respect the species is included in the `striata'
group (see remarks for C. (C.) striata).
Clathria (Clathria) nexus (Koltun, 1964)
(Figs 53-54)zyxwvutsrqponmlkjihgfedcb
Bipocillopsis nexus Koltun, 1964a: 79-80.
Clat hria nexus; Hooper & Wiedenmayer, 1994: 260
MATERIAL. HOLOTYPE: ZIL 10644 (not seen):
Clarie Coast or Wilhelm Land, Australian Antarctic
Territory, 65 °48' S, 89 °49'E, 310-400m depth (dredge).
PARATYPES: BMNH1963.7.29.56, ZIL 11525: same
locality.
HABITAT DISTRIBUTION. 310-400m depth; substrate unknown; Australian Antarctica Territory (Fig.
53F).
DESCRIPTION. Shape. Erect arborescent
growth form, 88-100mm high, 32-55mm wide,
with tightly anastomosing cylindrical branches,
up to 4mm diameter; tips of branches bifurcate,
relatively sharply pointed.
Colour. Grey-brown preserved.
Oscules. Numerous small oscules, 1-2mm
diameter, scattered over lateral margins of all
branches.
Texture and surface characteristics. Surface
hispid, raised into irregularly distributed, sharply
pointed microconules.
Ectosome and subectosome. Ectosomal skeleton
with choanosomal principal styles erect on surface, and bundles of subectosomal auxiliary
spicules surrounding protruding principal
spicules, or lying paratangential to surface.
Choanosome. Choanosomal skeleton subrenieroid-reticulate, without clearly defined
spongin fibres, but with heavy collagen enclosing
principal subtylostyles and sparse acanthostyles
echinating; mesohyl matrix heavy, with
numerous isochelae, few auxiliary styles and
some detritus dispersed throughout.
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FIG. 53. Clathria (Clathria) nexus Koltun (paratype BMNH1963.7.29.56). A, Choanosomal principal subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Unguiferous palmate isochclae.
E, Section through peripheral skeleton. F, Antarctic distribution. G, Paratype.
Megascleres. Choanosomal principal styles long,
Microscleres. Isochelae, strongly curved sig-
robust, straight or slightly curved towards the
distal end, tapering to sharp points, with slightly
swollen subtyloted bases, liberally microspined;
basal spines may partially extend up shaft near
basal end. Length 5184567.3)-62011m, width 18(22.4)-2511m.
Subectosomal auxiliary styles long, straight,
abruptly pointed,with only slight basal swelling
and pointed-hastate bases. Length 366-(394)415m, width 3-(5.6)-7[Lm.
Echinating acanthostyles relatively long,
robust, subtylote, sharply pointed, with heavy
recurved spines and heaviest concentrations of
spines on basal end. Length 214-(241.3)-278Rm,
width 9-(13.8)-17Rm.
moid, unguiferous with vestigial teeth, of
anchorate or arcuate modification. Length 14(16.6)-19iLm.
Toxas absent.
REMARKS. This species is similar to C. (T)
michaelseni in having bidentate sigmoid
isochelae, but differing in skeletal architecture
and in most other features. These unguiferous,
sigmoid-like chelae with vestigial, pointed alae,
are not unique to these austral species, also
known in W. Indian Ocean C. (C.) spongodes
Dendy (including its synonym C. madrepora
Dendy). In C. (C.) spongodes these reduced
chelae were initially thought to be sigmas
(Dendy, 1922; Burton, 1959a; Vacelet et al.,
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REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
FIG. 54. Clathria (Clathria) nexus Koltun (paratype BMNH1963.7.29.56). A, Choanosomal skeleton. B,
Peripheral skeleton C, Ectosomal structure. D, Fibre characteristics. E, Principal subtylostyles. F, Bases of
principal subtylostyles. G, Base of auxiliary style. H, Echinating acanthostyle. I, Acanthostyle spines. J,
Unguiferous palmate isochelae.
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1976), but the type material showed that these are
in fact reduced unguiferous isochelae with vestigial teeth.
Clathria (C.) nexus is also unusual in
Microcionidae in having auxiliary styles obviously associated with (surrounding) protruding
principal spicules, reminiscent (or analogous to)
Raspailiidae. Unlike raspailiids, however,
auxiliary spicules are also dispersed within the
peripheral skeleton, paratangential to the surface.
The other alternative, that the species belongs to
Raspailiidae, is rejected due to the chelae
microscleres. The bidentate-derived chelae is unusual to the Microcionidae and grounds to exclude the species from it under the phylogeny of
Poecilosclerida hypothesised by Hajdu et al.
(1994), but in all other respects the species fits in
this group supporting the present classification.
Clathria (Clathria) noarlungae sp.nov.
(Figs 55-56, Plate 1E, Table 9)
MATERIAL. HOLOTYPE: SAMTS4047 (fragment
NTMZ1632): Port Noarlunga, SA, 35 °09'S, 13 29'E,
1973, coll. SA Fisheries (trawl). OTHER MATERIAL:
S AUST - QMG300247 (NCIQ66C-2468-X, fragment
NTMZ3566).
HABITAT DISTRIBUTION. 5-30m depth; from rock
reef substrate; Port Noarlunga and Kingston, SE
S.Aust.
DESCRIPTION. Shape. Branching, 58-190mm
long, cylindrical digitate, bulbous-lobate
branches, 15-33mm diameter, with bulbous lobes
on tips of digits, single or bifurcate tips.
Colour. Orange alive (Munsell 1OR 6/10), greybrown preserved (7.5YR 5/4).
Oscules. Large oscules, 3-5mm diameter, in-line
on lateral sides of branches and on apex of terminal bulbs.
Texture and suiface characteristics. Texture rubbery, compressible, fibrous, difficult to tear; surface optically smooth, with few low rounded
bulbous projections; detachable skin-like dermis,
which is microscopically evenly porous, without
projecting spicules.
Ectosome and subectosonie. Ectosomal skeleton
consisting of relatively even, light series of
plumose brushes of ectosomal auxiliary subtylostyles, standing perpendicular or paratangential to
surface, usually surrounding inhalant pores and
forming a spiculo-fibrous reticulation on surface;
ectosomal skeleton thin, 75-149pm wide, perched on ends of peripheral, ascending choanosomal
fibres; subectosomal skeleton not differentiated
from dermal skeleton, and ectosomal spicule
brushes composed of a single category of
auxiliary megascleres only; megascleres coring
peripheral fibres sometimes project into, but not
through ectosomal skeleton, but this is exceptional.
Choanosome. Choanosomal skeletal architecture
arborescent, cavernous, consisting of well differentiated primary ascending and secondary
transverse components; primary spongin fibres
multispicular, 35-98Rm diameter, relatively
heavily invested with spongin, forming dendritic,
arborescent, radial, relatively even structure;
coring spicules in primary fibres occupy only a
small proportion of sponge diameter in axial
skeleton, becoming more heavily cored and increasingly plumose towards peripheral skeleton;
spongin fibres cored by larger subectosomal
auxiliary styles, and echinating acanthostyles
occur only sparsely throughout entire skeleton;
secondary spongin fibres entirely free of coring
megascleres, 12-57p.m diameter, forming a
plumo-reticulate structure; meshes formed by
primary and secondary spongin fibre branching
are ovoid-elongate to rectangular in shape, relatively even, and markedly cavernous (155560p,m maximum diameter); choanocyte
chambers paired, 90-311 p.m maximum diameter,
becoming larger towards periphery; mesohyl
matrix slightly granular, with toxas dispersed
singly or in dragmata; thin, rhaphidiform, vestigial auxiliary megascleres also dispersed between fibres, difficult to distinguish from toxas.
Megascleres (refer to Table 9 for dimensions).
Choanosomal principal megasclere absent, or
completely undifferentiated from subectosomal
auxiliary spicules.
Subectosomal auxiliary styles, coring fibres,
thin, mostly straight, varying from hastate to almost fusiform, with smooth tapering, rounded or
very slightly subtylote bases.
Ectosomal auxiliary styles geometrically
similar to larger auxiliary styles, with slightly
more pronounced, smooth, subtylote basal constrictions.
Acanthostyles vestigial, with subtylote bases,
fusiform points, rudimentary spination, aspinose
points and necks.
Vestigial auxiliary megascleres dispersed
within mesohyl are styloid or quasidiactinal,
often sinuous, with tapering or rounded bases,
long fusiform points.
Microscleres (refer to Table 9 for dimensions).
Palmate isochelae minute, abundant, frequently
twisted (80% of samples), two size classes observed.
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Toxas accolada, rhaphidiform, common, occurring individually or in dragmata, exceedingly
thin, rhaphidiform with very slight rounded
central curvature and straight points, or less often
with more angular central curvature and slightly
reflexed arms.
ETYMOLOGY. Named for type locality.
REMARKS. There are some minor differences in
spicule dimensions between the two known
specimens of C. (C.) noarlungae (Table 9), although there is no doubt that they are conspecific.
In having sparsely cored ascending primary
fibres and uncored secondary connecting fibres
this species is similar to C. (C.)kylista and C. (C.)
angulifera although spicule geometry and dimensions differs between all three. This species is also
similar to C. (T) cactiformis (Lamarck) in growth
form and skeletal structure but they differ in
geometry and size of spicules, number of spicule
categories and structure of ectosomal and subectosomal skeletons.
This species is enigmatic in its higher systematic placement. On the one hand C. (C.) noarlungae has a classical, albiet thin ectosomal
skeleton typical of Thalysias species, consisting
of erect plumose brushes of smaller ectosomal
auxiliary spicules. This feature is structurally discrete from the primary dendritic, and secondary
plumo-reticulate choanosomal skeleton. Conversely, there is only a single category of auxiliary
style which forms these ectosomal brushes,
whereas the larger (subectosomal) auxiliary
spicules are confined to inside the choanosomal
fibres. Consequently this species technically
belongs to C. (Clathria) (in having an undifferentiated ectosomal-subectosomal skeletal composition). Clathria (C.) noarlungae may be confused
with C. (Dendrocia) curvichela and C. (D.)
elegantula due to superficial similarities in
spiculation and fibre characteristics, but it differs
from these (and other Dendrocia) in having two
different auxiliary spicule geometries, with palmate isochelae instead of arcuate-like isochelae,
and different skeletal construction.
This species differs from all other known
Clathria (and other microcionids) in several features: orange colouration; bulbous-lobate digitate
growth form; rounded bulbous surface projections; absence of choanosomal principal styles,
where dendritic multispicular primary spongin
fibres are cored by subectosomal auxiliary styles,
and the secondary fibre system is aspicular,
plumo-reticulate, and sparsely echinated by yes-
tigial acanthostyles; and specific spicule
geometries.
Clathria (Clathria) oxyphila (Hallmann, 1912)
(Figs 57-58, Table 10)
Wilsonella oxyphila Hallmann, 1912: 249-253, p1.34,
fig.3, text-fig.52; Guiler, 1950: 9.
Paradoryx oxyphda; Hallmann, 1920: 768.
cf. Clathria elegantula; Hallmann, 1912: 253.
cf. Clathria pintformis; Carter, 1885f: 354; Hallmann,
1912: 253.
Clathria oxyphila; Hooper & Wiedenmayer, 1994:
260.
MATERIAL. HOLOTYPE: AMZ51 (dry; fragment
AME8 I 7): Off Marsden Point, Kangaroo I., SA,
35°30'S, 137°45'E, 34m depth, 19.viii.1909, coll. FIV
`Endeaour (dredge; label Wilsonella oxyphila (curvichela) Hallmann, type'). PARATYPE: AME495
(dry): Oyster Bay, Tas, 42°40'S, 148°03'E, 60-80m
depth, coll. FIV 'Endeavour', dredge; (specimen label
`Wilsonella oxyphila, Type'; AM register MS name
Taraclathria oxyphila sp. nov.').
HABITAT DISTRIBUTION. 34-80m depth; substrate
unknown; E. coast (Tas.) and Kanagaroo I. (S. Aust.)
(Fig. 57G).
DESCRIPTION. Shape. Fan, 152mm high,
155mm wide, planar or multiplanar, with a small
compressed, cylindrical basal stalk, 22mm long,
18mm diameter, thickly lobate, rounded, even
margins.
Colour. Grey-brown preserved.
Oscules. Oscules dispersed on margins of
branches.
Texture and surface characteristics. Surface relatively even, with radiating longitudinal subdermal grooves and pitted and porous ectosome.
Ectosome and subectosome. Ectosome
membraneous with relatively heavy mesohyl
matrix below surface, sparsely arenaceous, with
very few choanosomal principal megascleres
protruding through surface, and with sparse ectosomal skeleton of subectosomal auxiliary
megascleres lying paratangential to surface,
sometimes in light brushes around fibre endings.
Choanosome. Choanosomal fibres immediately
subdermal, becoming slightly plumose near
periphery; choanosomal skeleton sometimes
dendritic in places, usually plumo-reticulate.
Spongin fibres well developed, clearly divided
into primary (ascending, sinuous, multispicular)
and connecting (transverse, aspicular) elements;
coring megascleres vestigial principal styles,
barely differentiated from subectosomal
auxiliary styles; spongin fibres heavily echinated
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All
FIG. 55. Clathria (Clathria) noarlungae sp.nov. (holotype SAT54047). A, Choanosomal principal styles. B,
Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle and modified style. D, Raphidiform-accolada
toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, fragment of
holotype. I, Specimen NTMZ3566.
by acanthostyles, evenly dispersed over fibres;
mesohyl matrix heavy, with few scattered subectosomal megascleres.
Megascleres (Table 10). Choanosomal principal
styles slightly shorter and thicker, but otherwise
with similar geometry to those occurring in
peripheral skeleton; straight, hastate, with tapering or quasidiactinal, smooth bases.
Subectosomal auxiliary styles long, thin, slightly curved or sinuous, often with blackened axial
canals; geometrically similar to principal
megascleres.
Acanthostyles slightly subtylote, evenly
spined, including oxeote modifications.
Microscleres (Table 10). Palmate isochelae sig-
moid with reduced alae or vestigial teeth resem-
bling true sigmas; isochelae predominant on
membraneous ectosome.
Toxas intermediate between wing-shaped and
u-shaped, uncommon, with low, rounded central
curves, slightly reflexed points.
Larvae. Incubated parenchymella larvae ovalelongate, 120 x 210vt,m, with moderately light
mesohyl.
REMARKS. This species was included in Wilsonella by Hallmann (1912, 1920) and other
authors on the basis that principal and auxiliary
styles were allegedly undifferentiated, but re-examination of type material found them to be
similar but not identical in geometry (both relatively vestigial and modified quasidiactinal
form). In this respect the species is similar to C.
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FIG. 56. Clathria (Clathria) noarlungae sp.nov. (QMG300247). A, Choanosomal skeleton. B, Fibre characteristics. C, Echinating spicules in situ. D, Echinating acanthostyle. E, Acanthostyle spines. F, Raphidiform-accolada toxas. G, Palmate isochelae.
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FIG. 57. Clathria (Clathria) oxyphila (Hal!mann) (holotype AMZ51). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Intermediate wing shaped - u-shaped toxa.
E, Sigmoid palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution. H, Paratype
AMFA95.
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137
FIG. 58. Clathria (Clathria) oxyphila (Hallmann) (fragment of holotype AME817). A, Choanosomal skeleton.
B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spines. F, Sigmoid
palmate isochelae. G, Intermediate wing-shaped - u-shaped toxa.
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TABLE 9. Comparison between present and published
records of Clathria (Clathria)noarlungae sp.nov. All
measurements are given in p.m, denoted as range (and
mean) of spicule length x spicule width (N=25).
TABLE 10. Comparison between present and published records of Clathria (Clathria) oxyphila
(Hallmann). All measurements are given in p,m,
denoted as range (and mean) of spicule length x
spicule width (N=25).
Holotype
(SAMTS4047)
Specimen
(NC1Q66C2468X)
Holotype (AMZ51)
Paratype (AME495)
Choanosomal
principal styles
absent
absent
Choanosomal
principal styles
104-(133.0)-152 x
1.8-(2.5)-3
119-(135.4)-154 x
1.5-(2.2)-3
Subectosomal
auxiliary styles
167-(204.4)-236 x
2.5-(4.8)-7.8
232-(250.2)-278 x 3(4.3)-5
Subectosomal
auxiliary styles
142-(171.1)-199 x
0.8-(1.4)-2
143-(164.0)-186 x
1.0-(1.6)-2.0
Ectosomal
auxiliary styles
87.5-(112.8)-156 x
1.8-(3.4)-4.6
117-(146.0)-175 x 2(2.7)-3.5
Echinating
acanthostyles
49-(60.3)-72 x 2.5(4.1)-4.5
54-(63.6)-71 x 3.5(4.2)-5.5
Echinating
36-(46.6)-54 x 3.8(5.2)-6.5
48-(55.6)-61 x 4.5(5.2)-6
Chelae
17-(21.8)-25
16-(19.6)-23
acanthostyles
Vestigial
auxiliary styles
133-(186.2)-202 x
1.5-(2.1)-2.6
118-(170.1)-186 x
1.0-(1.6)-2.0
Toxas
35@20)-64x 1.5(1.7)-2.0
I9-(50.6)-84x 1.0(1.4)-2.5
SPICULE
Chelae I
3-(6.3)-8.5
5-(7.3)-I0
Chelae II
12-(13.5)-16.5
13-(15.7)-I9
36-(159.5)-216 x
0.4-(0.8)-1.1
19-(145.3)-265 x
0.5-(0.9)- 1.5
Toxas
(C.) piniformis, C. (C.) raphanus and C.
(Dendrocia) elegantula, all referred here to an
artificial species-group termed the `oxyphila'
group, loosely corresponding to Hallmann's
(1912, 1920) concept of Wilsonella (which is also
possibly artificial). The present species differs
from these by its sigmoid microscleres (virtually
intermediate between palmate and arcuate-like
geometry). Hallmann (1912) suggested that these
chelae were arcuate, and thus he considered that
the species had affinities with C. (Dendrocia)
curvichela, but they actually appear to be no more
than slightly modified, vestigial palmate forms.
Similarly, Hallmann (1912) did not record toxas
in his description, which are definitely present in
type material. The modification of echinating
acanthostyles to acanthoxeote megascleres in this
species is reminscent of Crellidae but this
modification is superficial. Clathria (C.)
oxyphila was transferred to Paradoryx on the
basis of its alleged arcuate chelae, but in all other
respects the species is clearly a Clathria
(Clathria).
Clathria (Clathria) partita Hallmann, 1912
(Figs 59-60)
Clathria partita Hallmann, 1912: 223, p1.32, fig.3,
text-fig.46; Hooper & Wiedenmayer, 1994: 260.
Clathria (Clathria) cf. partita; Rudman & Avern,
1989: 335.
Pseudanchinoe partita; de Laubenfels, 1936a: 109.
MATERIAL. HOLOTYPE: AME1024(dry): Unknown locality, South Australian coast, 60m depth,
SPICULE
12.vii.1909, coll. FIV `Endeavour' (dredge); (label
`Pseudanchinoe partita, type').
HABITAT DISTRIBUTION. 60m depth; substrate unknown; SA (Hallmann, 1912), and possibly S WA
(Rudman & Avern, 1989) (Fig. 59F).
DESCRIPTION. Shape. Arborescent branching,
340mm long, 160mm wide, with a long cylindrical stalk, 55mm long, 14mm diameter, planar,
bifurcate and occasionally anastomosing flattened branches, 12-25mm wide, approximately
8mm thick.
Colour. Orange alive (Rudman & Avern, 1989),
brown, with a slightly whitish ectosomal crust.
Oscules. Small oscules, up to 2mm diameter, on
lateral sides of branches.
Texture and surface characteristics. Surface
rugose, laterally striated, resembling Ectyoplasia
tabula (Raspailiidae; Hooper, 1991: Fig. 50), and
margins of branches slightly wider and more
spatuliferous than basal portions.
Ectosome and subectosome. Ectosomal skeleton
almost halichondroid, well differentiated from
choanosomal region, consisting of paratangential
multispicular bundles of subectosomal auxiliary
styles, forming criss-cross tracts on surface.
Choanosome. Choanosomal architecture basically isodictyal, consisting of differentiated axial
and extra-axial sections of skeleton; axial region
composed of very heavy and sinuous spongin
fibres, forming relatively tight, oval or elongated
triangular meshes; axial fibres divided into larger
primary and smaller secondary elements, both
containing paucispicular core of choanosomal
principal styles; extra-axial skeletal architecture
an irregular isodictyal reticulation of sinuous
spongin fibres of approximately same diameter as
axial secondary elements, relatively lightly invested with spongin, containing uni-, bi-, or more
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139
rarely paucispicular tracts of choanosomal styles;
extra-axial fibres incompletely divided into
primary (ascending) and secondary (vaguely
transverse) components, differentiated mainly by
number of coring spicules; echinating acanthostyles sparse, evenly distributed throughout
skeleton, rarely incorporated into fibres (cf.
Hallmann, 1912); some choanosomal principal
styles may protrude through (echinate) fibres,
particularly in extra-axial skeleton; mesohyl
matrix very light containing few subectosomal
auxiliary megascleres.
Megascleres. Choanosomal principal styles
short, slightly curved or straight, almost hastate,
with smooth rounded or very slightly subtylote
bases. Length 1124172.6)-23911m, width 5(8.7)-1211m.
Subectosomal auxiliary styles long, slightly
curved or straight, fusiform, with rounded or
slightly subtylote bases. Length 2224267.7)315p.m, width 3.5-(5.1)-7p.m.
Acanthostyles slightly subtylote, with vestigial
spines and an aspinous basal region. Length 56(60.4)-8811m, width 3-(5.0)-61.m.
Microscleres. Isochelae absent.
Toxas sinuous rhaphidiform, usually symmetrical with slight angular central curvature and
straight points, but asymmetrical and reflexed
examples also occur. Length 724115.6)-2121m,
width 0.5-(0.7)-1.2Rm.
REMARKS. This species was referred to
Pseudanchinoe by de Laubenfels (1936a) because it lacked chelae microscleres, but that feature has since been shown to be of little
systematic value (e.g. Hooper, 1991: 288). As
noted by Hallmann (1912), C. (C.) partita is
known only from a single dry specimen, and it is
possible that isochelae may be eventually discovered in other better preserved specimens
should additional material become available.
Clathria (C.) partita has an unusual skeletal
construction, combining features of myxillid-like
microcionids (e.g. a basically isodictyal extraaxial skeleton closely resembling C. (Isociella)
eccentrica) and raspailiid-like microcionids (e.g.
compressed axis of C. (Axociella) canaliculata),
and features of the compressed axial skeleton are
also reminiscent of the skeletal plan seen in certain Axinellidae such as Cymbastela. On the basis
of these characteristics C. (C.) partita is similar
to C. (C.) rubens (Lendenfeld).
The record of C. (C.)partita from WA (Rudman
& Avern, 1989), is based on a photograph
without accompanying sample. The photograph
was taken of a nudibranch (Rostanga calumnus
Rudman & Avern) feeding on an orange sponge
from Esperance Bay (33°51'S, 121°57'E).
Clathria (Clathria) paucispicula (Burton, 1932)
(Fig. 61)
Rhaphidophlus paucispiculus Burton, 1932a:
320, p1.56, fig.1, text-fig.30; Burton, 1940: 111;
Desqueyroux, 1975: 68; Koltun, 1964a: 75;
Desqueyroux-Faundez & Moyano, 1987: 49.
Clathria paucispicula; Hooper & Wiedenmayer,
1994: 260
MATERIAL. HOLOTYPE: BMNH1928.2.15.243a: Near
Shag Rocks, South Georgia, 53°43.4'S, 40°57.0'W,
177m depth, coll. R.R.S. 'Discovery' (dredge).
HABITAT DISTRIBUTION 74-198m depth; on mud,
sand and rock substrates; Australian Antarctic Territory: Queen Mary Land (Koltun, 1964a) (Fig. 61D).
Also Falkland Is, South Georgia and S. Shetland Is
(Burton, 1932a), Mar del Plata, Argentina (Burton,
1940), Low Is, Chilean Antarctic Territory
(Desqueyroux, 1975), Tierra del Fuego (DesqueyrouxFaundez & Moyano, 1987).
DESCRIPTION. Shape. Massive, flabellatedigitate with irregular ridges and convoluted
branches.
Colour Live colouration unknown, light brown
preserved.
Oscules. Large oscules 3-5mm diameter sparsely
scattered on apex of ridges.
Texture and surface characteristics. Texture firm,
tough, compressible; surface uneven, minutely
conulose.
Ectosome and subectosome. Surface skeleton
tangential confused crust of smaller ectosomal
auxiliary styles, choanosomal principal styles
and foreign spicule fragments together forming a
dense crust 0.5-1.0mm thick; immediately below
tangential ectosomal skeleton are erect bundles
of both auxiliary styles (producing an irregular
radial palisade of spicules), and principal styles
(more sparsely dispersed).
Choanosome. Skeletal architecture reticulate,
slightly plumo-reticulate near periphery, less organised towards axis; spongin fibres moderately
well developed cored by pauci- or multispicular
ascending tracts and interconnected by uni- or
paucispicular tracts of choanosomal principal
styles, producing rectangular or triangular
meshes up to 250Rm diameter; echinating
megascleres absent; mesohyl matrix light,
smooth, unpigmented.
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FIG. 59. Clathria (Clathria) partita Hallmann (holotype AME1024). A, Choanosomal principal styles. B,
Subectosomal auxiliary style. C, Echinating acanthostyle. D, Sinuous-raphidiform toxas. E, Section through
peripheral skeleton. F, Australian distribution. G, Holotype.
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141
FIG. 60. Clathria (Clathria) part ita Hallmann (holotype AME1024). A, Choanosomal skeleton. B, Peripheral
skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Vestigial acanthostyle spines. F, Fragments of
sinuous and raphidiform toxas.
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FIG. 61. Clathria(Clathria)paucispicula (Burton) (holotype BMNH1928.2.15.243a). A, Choanosomal principal
styles. B, Subectosomal auxiliary styles. C, Section through peripheral skeleton. D, Antarctic distribution. E,
Choanosomal skeletal structure.
Megascleres. Choanosomal principal styles long,
thick, slightly hastate pointed or occasionally
slightly telescoped pointed, curved at centre, with
smooth rounded or slightly subtylote bases.
Length 535-(663.4)-754Rm, width 18423.3)31pm.
Subectosomal auxiliary styles relatively long,
thick, straight or slightly curved near basal end
with evenly rounded smooth bases and tapering
fusiform points. Length 255-(318.6)-402Rm,
width 6410.4)-1411m.
Echinating spicules absent.
Microscleres. Absent.
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143
REMARKS. This species resembles a Suberitidae (Hadromerida) in skeletal structure, with
the main clue as to its affinities with
Microcionidae being the possession of
megascleres that are not truly tylote in geometry,
two distinct categories of principal and auxiliary
megascleres, and slight compression of the
skeleton in the axial region not generally seen in
suberitids. Certain allocation of this species is
difficult given that spicule diversity and skeletal
structure are the main diagnostic characters for
microcionids, and the species is aptly named for
the reduction in these characters. It is well known
and possibly widely distributed in Antarctic
waters.
Clathria (Clathria) pauper Brondsted, 1926
(Figs 62-63)
Clathria pauper Brondsted, 1926: 3, text-fig.3; Burton,
1929a: 398; Burton, 1940: 109; Koltun, 1964a: 69,
p1.12, figs 4-6; Sara, 1978: 66-67, text-fig.40;
Desqueyroux-Faundez & Moyano, 1987: 50.
Ramoses pauper; de Laubenfels, 1936a: 109.
MATERIAL. HOLOTYPE: NRHM (fragments
BMNH 1930.11.5.2, AMZ2239): N. of Discovery
Inlet, Victoria Land, Antarctica, 640m depth,
10.ii.I924, coll. Sten Warren (dredge).
HABITAT DISTRIBUTION. 10-640m depth; hard
and soft substrates; Australian Antarctic Territory: Victoria Land, Banzare Coast, Wilkes Land (Brondsted,
1926; Koltun, 1964a) (Fig. 62G). Also SW. Maldonado, Mar del Plata, Argentina, S. Brazil (Burton,
1940), Cape Sebastiano, Cape Domingo, Rio Grande,
Tierra del Fuego (Sara, 1978).
DESCRIPTION. Shape. Erect branching digitate
sponge, up to 80mm high.
Colour. Grey-brown preserved.
Oscules. Unknown.
Texture and surface characteristics. Texture firm,
compressible, elastic; surface conulose with irregularly digitate processes, up to 20mm long and
4mm thick, and with convoluted distinct ectosomal membrane visible between conules.
Ectosome and subectosome. With irregular
plumose brushes of subectosomal auxiliary styles
protruding from end of choanosomal primary
skeletal columns.
Choanosome. Skeletal architecture plumosereticulate, with differentiated primary and secondary fibre systems; primary fibres ascending,
multispicular, diverging near surface, cored by
erect choanosomal principal styles with points of
spicules directed upwards and outwards; skeletal
columns dominated by spicules with poor spon-
gin; secondary fibres uni- or paucispicular, interconnecting adjacent primary tracts, forming irregular or triangular meshes up to 350Rm
diameter; echinating acanthostyles protruding at
acute angles from fibres, or clumped in plumose
brushes, scattered over the whole length of primary
tracts although more sparse near surface; mesohyl
moderately heavy, lightly pigmented, granular.
Megascleres. Choanosomal principal styles long,
thick, straight or slightly curved at midsection,
tapering to long fusiform points, most commonly
with rounded smooth bases or occasionally with
microspined subtylote bases. Length 372(606.1)-810p,m, width 11-(15.8)-21Rm.
Subectosomal auxiliary styles shorter, more
slender than principals, straight, with fusiforrn
points and microspined subtylote bases. Length
352-(480.8)-590Rm, width 3-(7.6)-10p.m.
Echinating acanthostyles in two size
categories. Larger form intermediate between the
smaller and the principal styles, slightly curved at
midsection, fusiform pointed, subtylote, with
heavily spined bases, lightly spined shafts and
often with bare base. Length 2194293.0)384iim, width 10-(12.3)-15Rm. Smaller acanthostyles straight, slender, subtylote, fusiform,
more-or-less evenly spined except usually for an
aspinose point. Length 92-(148.4)-183p.m, width
Microscleres. Chelae absent.
Toxas of two forms: Accolada toxas, long, thin,
with long straight arms, sharply angular central
curve or spirally twisted centre, and non-reflexed
points. Length 934139.5)-18511m, width 0.8(0.9)-1.511m. Wing-shaped toxas, short, thick,
gradually curved at centre, curved arms, slightly
reflexed points. Length 31-(45.5)-52Rm, width
REMARKS. Koltun (1976) suggested that C.
pauper is synonymous with Stylostichon
toxiferum Topsent and Microciona basispinosa
Burton, but respective types indicate that they
differ substantially in a number of significant
features, considered here to warrant recognition
of C. pauper as a distinct species (see remarks for
C. (Microciona) antarctica below). Clathria
pauper differs from other species by a combination of characters (plumose-reticulate skeletal
structure with differentiated primary and secondary fibre systems; principal spicules protruding
through fibres in `spicate' arrangement; two sizes
of acanthostyles echinating fibres; two toxa morphologies; and absence of chelae).
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FIG. 62. Clathria (Clathria) pauper Brondsted (fragment of holotype BMNH1930.11.5.2). A, Choanosomal
principal style. B, Subectosomal auxiliary style. C, Young auxiliary style. D, Larger category of echinating
acanthostyles. E, Smaller form of echinating acanthostyles. F, Short wing-shaped toxas and larger accolada
toxas. G, Known Antarctic distribution. H, Section through peripheral skeleton. I, Fragment of holotype.
^
145
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
FIG. 63. Clathria (Clathria) pauper Brondsted (fragment of holotype BMNH1930.11.5.2). A, Choanosomal
skeleton. B, Ectosomal skeleton. C, Peripheral skeletal structure. D, Fibre characteristics. E, Choanosomal
principal subtylostyle. F, Base of principal subtylostyle. G, 2 sizes of echinating acanthostyles. H, Spined base
of acanthostyle. I, Acanthostyle spine morphology. J, Accolada toxa. K, Wing-shaped toxa.
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A
FIG. 64. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hr ia ( Clat hr ia) pet for at a (Lendenfeld) (lectotype AMMG9010). A, Choanosomal principal
subtylostyle. B, Subectosomal auxiliary style. C, Echinating acanthostyle. D, Palmate isochelae. E, Section
through peripheral skeleton. F, Distribution. G, Skeletal structure. H. Surviving portion of type specimen.
Clathria (Clathria) perforata (Lendenfeld,
1887) (Fig. 64)
in part, Lendenfeld, 1887b:
788; in part, Lendenfeld, 1888: 89-90. (Not Lendenfeld, 1887b: p1.22, fig.44.)
Ant herochalina perforat a,
Whitelegge, 1902b: 275, 279, 287;
Hooper & Wiedenmayer, 1994: 260.
Clat hria perforat a;
MATERIAL. LECTOTYPE: AMG9010: Port Phillip,
Vic, 38°09'S, 144°52'E, other details unknown
(specimen label reads `Antherochalina perforata
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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147
Lend.', whereas AM register reads `Suberapsamma
philippi Lend., Port Phillip'). (Not SYNTYPE BMNH1886.8.27.459: Broughton I., Port Stephens,
NSW, 32°36'S, 152°19'E).
forata could also be assigned to Pandaros or
Echinoclathria, whereas the few vestigial acanthostyles seen in the lectotype suggest that
Clathria (Clathria) is more appropriate.
HABITAT DISTRIBUTION. Ecology unknown; SE.
coast of Australia (Vic, NSW) (Fig. 64F).
Clathria (Clathria) piniformis (Carter, 1885)
DESCRIPTION. Shape. Irregularly flabello-
digitate, thin, with a small stalk.
Colour. Live colouration unknown, dark brown
preserved.
Oscules. Unknown.
Texture and surface characteristics. Surface
arenaceaous, porous; texture harsh..
Ectosome and subectosome. Ectosome fibrous,
arenacious, little remaining of original dermal
skeleton.
Choanosome. Choanosomal skeleton irregularly
isodictyal reticulate, with relatively heavy spon-
gin fibres forming oval or hexagonal meshes,
cored by one or few vestigial choanosomal principal styles; echinating acanthostyles very scarse,
scattered evenly throughout skeleton; mesohyl
matrix contains heavy deposits of siliceous
detritus, especially large sand grains, with heavily
pigmented and granular mesohyl matrix.
Megascleres. Choanosomal principal subtylostyles thin, fusiform, straight, with slightly subtylote, smooth bases. Length 96-(105.0)-11811m,
width 4-(4.8)-6p.m.
Subectosomal auxiliary styles barely differentiated from principal megascleres, usually much
thinner, only seen scattered between fibres, few
at surface. Length 854103.3)-14611m, width I
-
(2.7) 411m.
-
Acanthostyles small, slightly subtylote, with
vestigial or granular spination. Length 46457.7)72p.m, width 2.5-(3.1)-4.5p.m.
Microscleres. Palmate isochelae rare. Length 8(9.7)-11[Lm.
Toxas absent.
REMARKS. Lendenfeld (1887) originally
nominated two `syntypes', but one (BMNH)
(Lendenfeld, 1887b: p1.22, fig.44) is Antho (see
A. (Isopenectya)chartacea), whereas the description was based on the other AM specimen (Lendenfeld, 1887b: 788, 1888: 89). This latter
specimen is designated lectotype, but is in poor
condition and the species is poorly known: ectosomal features cannot be clearly discerned;
most megascleres within fibres are broken; and
microscleres are extremely difficult to pick out
between the abundant detritus within the
mesohyl. In having flattened fibres which form
irregular (isodictyal) anastomoses, C. (C.) per-
(Figs 65-66, Table 11)
Dictyocylindrus piniformis Carter, 1885f: 354.
Clathria piniformis; Dendy, 1896: 34; Hooper &
Wiedenmayer, 1994: 261.
Wilsonella piniformis; Hallmann, 1912: 241.
Paradoryx piniformis; Hallmann, 1920: 768.
MATERIAL. HOLOTYPE: BMNH1886.12.15.62
(fragments MNHNDCL60, AMG2803): Port Phillip
Heads, Vic, 38°17'S, 144°39'E, 38-40m depth, coll.
J.B. Wilson (dredge). OTHER MATERIAL: VIC NMVRN412-508.
HABITAT DISTRIBUTION. 38-40m depth; substrate
unknown; Port Phillip Bay(Fig. 65G).
DESCRIPTION. Shape. Massive, subspherical
lobate or lobo-digitate growth form, 110-130mm
long, 55-65mm wide, with thick cylindrical stalk,
45mm long, 18-22mm diameter.
Colour Bright orange-brown alive, olive-brown
preserved.
Oscules. Large oscules, 2-4mm diameter dispersed over both faces of digitate lobes.
Texture and surface characteristics. Surface
rugose, with irregularly distributed conules, and
a skin-like dermis.
Ectosoine and subectosome. Ectosome
membraneous, with tangential and paratangential
bundles of subectosomal auxiliary styles, sometimes forming quite thick, sinuous tracts.
Choanosome. Choanosomal skeleton reticulate;
spongin fibres well developed, thin, forming
sinuous tracts, branching and splitting, and
producing oval or elongate meshes; spongin
fibres without clear differentiation between
primary or secondary elements, although thicker
fibres, usually ascending and cored by principal
styles, whereas connecting fibres usually
aspiculose; spongin fibres cored with
paucispicular tracts of vestigial choanosomal
principal styles, and echinated by small acanthostyles in heavy concentrations and relatively evenly dispersed; mesohyl matrix heavy, with
numerous extra-fibre spicules, especially acanthostyles and auxiliary styles in the peripheral
skeleton.
Megascleres (Table 11). Choanosomal principal
subtylostyles very thin, vestigial, with blackened
axial canals, fusiform, usually straight, some-
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C
FIG. 65. Clathria (Clathria) piniformis (Carter) (holotype BMNH1886.12.15.62). A, Choanosomal principal
subtylostyle. B, Subectosomal auxiliary style. C, Echinating acanthostyle and modified form. D, Raphidiform
toxa. E, Palmate isochela. F, Section through peripheral skeleton. G, Australian distribution. G, Holotype. I,
Specimen NMVRN412. J, Choanosomal skeleton.
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149
FIG. 66. Clathria (Clathria) piniformis (Carter) (holotype BMNH1886.12.15.62). A, Choanosomal skeleton. B,
Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Modified
acanthostyle. G, Modified acanthostyle spination. H, Base of principal style. 1, Principal style.
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times slightly curved, with rounded, tapering or
more frequently quasidiactinal, smooth bases.
Subectosomal auxiliary styles almost identical
to principal styles, longer, thin, slightly curved or
sinuous, fusiform, with basal endings similar to
principal megascleres.
Acanthostyles slightly subtylote with vestigial,
evenly dispersed spines, include quasidiactinal
modifications to base.
Microscleres (Table 11). Palmate isochelae rare,
unmodified.
Toxas rhaphidiform, uncommon, with slight
central curvature and reflexed points, sometimes
sinuous.
REMARKS. In vestigial spicule geometry and
generally reduced skeletal structures C. (C.)
pintformis is most similar to C. (C.) oxyphila,
both of which have most of their megascleres
modified to quasidiactinal forms (see remarks for
C. oxyphila), but the present species has quite
different microsclere geometry, skeletal construction and growth form.
Clathria (Clathria) raphanus (Lamarck,
1813) (Figs 67-68)
Spongia raphanus Lamarck, 1813: 444; 1814: 373.
Clathria raphanus; Topsent, 1932: 100, p1.4, fig.9;
Hooper & Wiedenmayer, 1994: 261.
Clathria raphana; Van Soest, 1984b: 108.
Thalyseurypon raphanus; de Laubenfels, 1936a: 107.
Panda ros raphanus; Wiedenmayer, 1977: 143-4.
MATERIAL. HOLOTYPE: MNHNDT572(dry):
Australian seas, Peron & Lesueur collection, no other
details known.
HABITAT DISTRIBUTION. Ecology unknown;
Australia.
DESCRIPTION. Shape. Arborescent fan, 90mm
high, 95mm wide, with flattened multiplanar
branches 48-80mm long, 5-10mm thick, which
anastomose tightly to form several lobes.
Colour Grey-brown dry.
Oscules. Large oscules, up to 3.5rnm diameter,
scattered over both faces of flattened branches.
Texture and surface characteristics. Surface
microconulose with tapering processes; texture
harsh in dry state.
Ectosome and subectosome. Ectosomal skeleton
a sparse tangential or paratangential layer of subectosomal auxiliary styles.
Choanosome. Choanosomal skeleton almost
regularly reticulate, with heavy spongin fibres
forming oval or elongate meshes, without differentiation of primary or secondary elements;
TABLE 11. Comparison between present and published records of Clathria (Clathria) piniformis (Carter).
Measurements in pin, denoted as range (and mean)
of spicule length x spicule width (N=25).
SPICULE
Holotype (BWINH-
1886.12.15.62)
Specimen (N=1)
Choanosomal
principal styles
124-(137.6)-156 x
1.5-(2.2.)-3
Subectosomal
auxiliary styles
147-(164.4)-182 x
2.5-(3.1)-4
128-(144.4)-154 x 2(2.6)4
149-(178.3)-197 x 2(3.9)-6
Echinating
acanthostyles
51-(57.1)-64 x 2.5(3.2)-4
49-(54.2)-58 x 2(3.3)-6
Chelae
9-(10.2)-12
10-(10.8)-12
Toxas
92-(117.2)-143 x
0.5-(0.7)-1
1104122.0)-135 x
0.540.7)-0.8
spongin fibres almost totally uncored, and where
present, choanosomal principal styles form unior paucispicular tracts within axis of fibres; spongin fibres sparsely echinated by acanthostyles;
mesohyl light (dried material).
Megascleres. Choanosomal principal subtylostyles fusiform, slightly curved, with rounded or
slightly subtylote, smooth bases. Length 108(148.6)-230p.m, width 3.5-(5.6)-9p.m.
Subectosomal auxiliary subtylostyles straight
or slightly curved, thin, fusiform, with subtylote,
smooth bases. Length 168-(225.7)-315p,m, width
1.5-(2.6)-4tim.
Acanthostyles curved, slightly subtylote, with
vestigial, granular, evenly dispersed spines.
Length 47-(65.2)-82p,m, width 2.5-(4.1)-51.tm.
Microscleres. Absent.
REMARKS. This species is known only from an
antiquated dried specimen collected from 'somewhere in the vicinity of Australia' (Topsent,
1932). It was made the type species of
Thalyseurypon de Laubenfels and has since been
the subject of lengthy discussions by de Laubenfels (1936a), Hechtel (1965), Wiedenmayer
(1977), Van Soest (1984b) and Fromont &
Bergquist (1990). These discussions have centred
mostly on the value of the loss of meniscoid
(chelae) microscleres used as a valid taxonomic
character. These arguments have been addressed
in the synopsis of Thalyseurypon (see Included
Genera, above). Clathria raphanus bears little
resemblance to Pandaros as supposed by
Wiedenmayer (1977) which has flattened fibres,
or Echinoclathria (as interpreted by Wiedenmayer, 1989) which has smooth echinating
megascleres. This species is most appropriately
placed in Clathria (Clathria) based on the distribution and geometry of its megascleres.
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Clathria (C.) raphanus is depauperate in morphological features, with a reduced spicule diversity and vestigial spicule development, and a
relatively heavy evenly reticulate fibre skeleton,
and in this respect the species is associated with
a species-group (the oxyphila' group; also containing C. (C.) pinifonnis, C. (C.) oxyphila and C.
(Dendrocia) elegantula).
Clathria (Clathria) rubens (Lendenfeld,
1888) (Figs 69-70, Table 12)
Thalassodendron rubens Lendenfeld, 1888: 223.
Not Thalassodendron rubens var. dura, in part, Lendenfeld, 1888: 223-224.
Not Thalassodendron rubens var. lamella, in part, Lendenfeld, 1888: 224, p1.7.
Clathria rubens, in part; Whitelegge, 1901: 85-86,
p1.11, fig.13.
Clathria rubens; Hallmann, 1912: 218-223, p1.32,
fig.1, text-fig.45; Burton, 1934a: 558; Burton,
1938b: 20; Shaw, 1927: 425-426; Guiler, 1950: 7;
Hooper & Wiedenmayer, 1994: 261.
Clathria tenuifibra Whitelegge, 1901: 82-83, p1.11,
fig.10; Whitelegge, 1902b: 274, 279, 287;
Hallmann, 1912: 211.
Thalysias tenuifibra; de Laubenfels, 1936a: 105.
Placochalina pedunculata var. monis, in part; Lendenfeld, 1888: 91-92; Whitelegge, 1902b: 274.
cf. Microciona prolifera; Vosmaer, 1935a: 611, 644,
668.
Not Microciona rubens Bergquist, 1961a: 38.
MATERIAL. LECTOTYPE: AMG9119 (dry): Port
Jackson, NSW, 33°51'S, 151°16'E, no other details
known (label `Thalassodendron rubens RvL, type').
PARALECTOTYPES: AMZ455 (slide AMG3585:
same locality (dry, label 'Clathria rubens, type or
cotype, Lendenfeld's duplicate A50'). AMZ634: same
locality; (dry, label `Thalassodendron rubens RvL,
type or cotype, Lendenfeld's no.240').
BMNH1887.1.24.35 (fragment AMG3586): same
locality, Ramsay collection (listed as 'var. intermedia
MS name'). HOLOTYPE of C. tenuifibra:
AMG3045(dry): Lake Illawarra region, NSW,
34°32'S, 150°50'E (label `Thalysias tenuifibra
Whitelegge; type'). OTHER MATERIAL: NSW QMG300452.
HABITAT DISTRIBUTION. Shallower than 56m
depth; on shell, gravel and Halimeda substrates; Port
Jackson (Lendenfeld, 1888), Broken Bay, Lake IIlawarra, and Woolongong (NSW) (Whitelegge, 1901;
Hallmann, 1912); Direction and Eagle Is, Great Barrier
Reef (FNQ) (Burton, 1934a); Maria I. (Tas) (Shaw,
1927; Guiler, 1950); St Vincent Gulf (SA) (Hallmann,
1912) (Fig. 69G).
DESCRIPTION. Shape. Arboresecent digitate
sponge,100-190mm high, 60-110mm wide, with
numerous anastomosing, repeatedly bifurcate,
stoloniferous branches, 30-75mm long, up to
19mm diameter, without definite basal stalk;
branches subcylindrical, slightly flattened at
ends.
Colour Grey-brown in ethanol.
Oscules. Large oscules, 2-4mm diameter, scattered over surface of branches, often raised on a
small lip.
Texture and surface characteristics. Surface
even, optically smooth, but with prominent subdermal grooves and stellate drainage canals
radiating from a central osculum (not seen in dry
material).
Ectosome and subectosome. Ectosomal region
membraneous, with a sparse tangential layer of
subectosomal auxiliary styles, through which
protrude sparse brushes of choanosomal principal
styles; subectosomal skeleton has auxiliary styles
dispersed paratangentially, and peripheral choansomal fibres diverging slightly just below surface.
Choanosome. Choanosomal skeletal architecture
subisodictyal, with triangular or irregular meshes
and thin spongin fibres, cored by uni-, bi- or
paucispicular tracts of choanosomal principal
styles; spongin fibres imperfectly divided into
axial and extra-axial components, differing only
in fibre diameter; axial fibres comparatively
thick, paucispicular, forming cavernous and irregular ovoid meshes; extra-axial skeletal meshes
more obviously triangular, subisodictyal, with
thin, paucispicular ascending primary spongin
fibres interconnected by uni- or bispicular secondary fibres; echinating acanthostyles lightly and
evenly dispersed over fibres; mesohyl matrix
heavier in peripheral skeleton than at core, containing few principal and auxiliary styles and
microscleres.
Megascleres (Table 12). Choanosomal principal
styles thick, mostly straight, hastate-fusiform,
with rounded, tapering or subtylote bases, usually
smooth, occasionally microspined.
Subectosomal auxiliary subtylostyles longer
than principal styles, thin, straight, fusiform, with
slightly subtylote, microspined bases.
Acanthostyles subtylote, with mostly granular,
evenly dispersed spines, sometimes with
aspinose necks, and some clearly intermediate
forms of choanosomal styles.
Microscleres (refer to Table 12 for dimensions).
Palmate isochelae rare, small, unmodified.
Toxas accolada, thin, long, usually with very
large, angular, sometimes coiled, central curves,
slightly reflexed points, or they may be nearly
straight with coiled central flexion.
MEMOIRS OF THE QUEENSLAND MUSEUM
152zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
(C.) partita), geometry of
choanosomal styles and toxas
(Figs 59, 69), and having
similar growth forms (although
branches are cylindrical in C.
(C.) rubens but flattened in C.
(C.) part ita). Nevertheless,
spicule geometries, fibre characteristics and spicule dimensions indicate that they are
probably separate sibling
species.
There are 3 other AM
specimens which have been associated with C. (C.) rubens.
One, AMZ4810 from Port
Jackson (with label `Thalassodendron rubens, donated by
A. Dendy'), is probably a fragment of the BMNH paralectotype, but this has not been
verified. AMZ2241 from Tasmania (apparently identified
by M. Burton, with an old label
Rhaphidophlus typicus, dried
up in 1917-18'), and AMZ2246
(locality unknown, also identified by M. Burton, with an old
label which reads `Crella
incrustans'), were both obtained from the Antarctic Publications Committee (donated
by Prof. Haswell). All three are
in too poor condition to identify reliably, but they appear
related to C. (Thalysias) cactifonnis. The origin of the QM
specimen, obviously of great
antiquity, is not known (L.
FIG. 67. Clathria (Clathria) raphanus (Lamarck) (holotype MNHNDT572). Cannon, pers. comm.), but it
A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary sub- may have been an exchange
tylostyle. C, Echinating acanthostyle. D, Section through peripheral specimen from the AM during
skeleton. E, Holotype.
Whitelegge's era. That
specimen certainly belongs to
REMARKS. Clathria (C.) rubens is very similar
C. (C.) rubens.
to C. (C.)partita in skeletal architecture and fibre
The holotype of C. tenuifibra is also in poor
characteristics. Both species have a more-or-less
condition,
and irrespective of skeletal reconstitudifferentiated axial and extra-axial choanosomal
skeleton. Whereas the present species has an ir- tion following rehydration during sectioning only
regularly subisodictyal reticulate skeleton with few features were discernible. What little can be
ovoid meshes and heavy fibres, C. (C.)partita has seen of the skeletal structure is identical to C. (C.)
a nearly regularlsociella-like isodictyal skeleton.
These two species are also similar in the extent to rubens, particularly the fibre characteristics and
which axial and extra-axial skeletons are spiculation, and on this basis the two species are
separated (with more marked separation in C. merged here.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
153
FIG. 68. Clathria (Clathria) raphanus (Lamarck) (holotype MNHNDT572). A, Choanosomal skeleton. B, Fibre
characteristics. C, Membraneous ectosomal region. D, Echinating acanthostyles. E, Vestigial acanthostyle
spines.
Clathria (Clathria) squalorum Wiedenmayer
in Hooper & Wiedenmayer, 1994
(Figs 71-72)
Clathria squalorum Wiedenmayer, in Hooper &
Wiedenmayer, 1994: 261.
Clathria dura var. moths Hentschel, 1911: 370-372,
text-fig.45; Hallmann, 1912: 242.
Not Clathria dura Whitelegge, 1901: 83.
Not Clathria mollis Kirkpatrick, 1903: 249.
MATERIAL. LECTOTYPE: ZMB4444: Freycinet
Reach, Shark Bay region, WA, 26°05'S, 113°30'E,
3.5-11m depth, 5.ix.1905, coll. W. Michaelsen & R.
Hartmeyer (dredge). PA RA LECTOTYPES: HM:
same locality (9 dry specimens). HM: Geographe Bay,
Bunbury region, WA, 33°35'S, 115°26'E (5 dry
specimens).
HABITAT DISTRIBUTION. 3.5-11m depth; on sand,
stone, mud and algal bed substrates; Bunbury-Shark
Bay region (WA) (Fig. 71G).
^ MEMOIRS OF THE QUEENSLAND MUSEUM
154zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 69. Clathria (Clathria) rubens (Lendenfeld) (lectotype AMZ9119). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Palmate isochela. E, Accolada toxas. F,
Section through peripheral skeleton. G, Australian distribution. H, Lectotype.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
155
Clat hria ( Clat hria) rubens (Lendenfeld) (A-B, NTMZ1527; C-G, QMG300452). A, Choanosomal
FIG. 70. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
skeleton. B, Fibre characteristics (x313). C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of
auxiliary subtylostyle. F, Palmate isochelae. G, Accolada toxas.
MEMOIRS OF THE QUEENSLAND MUSEUM
156zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
TABLE 12. Comparison between present and published records of bases. Length 854148.5)Clathria (Clathria) rubens (Lendenfeld). All measurements are given in 184p.m, width 242.8)-411,m.
p.m, denoted as range (and mean) of spicule length x spicule width
Acanthostyles slightly sub(N=25).
tylote, with evenly dispersed
Lectotype
(AMG9119)
Paralectotypes
(N=3)
relatively large spines, slightly
Holotype of C.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Specimen (N=I) less spinose below basal region.
t enuij ibra
Length 48-(62.3)-7411m, width
(AMG3045)
Choanosomal
principal
styles
96-(135.3)-162
x 5-(9.0)-I I
128-(135.6)-147
x 5.5-(7.1)-9
1124139.2)-159
x 8-(9.4)-11
1184140.6)-155
x 6-(7.8)-10
Subectosomal
auxiliary
styles
106-(157.2)-212
x 1.5-(2.9)-4
1324146.2)-168
x 2.5-(2.9)-4
112-(153.7)-185
x 2.543.7)-4.5
105-(139.2)-165
x 2.5-(3.3)-4.5
Echinating
acanthostyles
55-(71.8)-111 x
3-(5.6)-7
58-(77.3)-112 x
4.5-(5.6)-6.5
51-(68.6)-118 x
5-(6.4)-8
48-(69.4)-I08 x
4-(5.8)-7.5
SPICULE
Chelae
6-(9.6)-I3
7-(9.8)-13
8-(9.7)-11
6-(9.0)-12
Toxas
101-(153.4)-215
x 0.8-(1.3)-1.5
115-(161.6)-208
x I-(1.4)-2
94-(126.3)-153
x I-(1.2)-1.5
114-(138.6)-194
x 1-(1.4)-2
DESCRIPTION. Shape. Variable growth form,
ranging from thickly encrusting with small
stoloniferous digits, to digitate planar arborescent
with anastomosing branches.
Colour Live colouration unknown, beige
preserved.
Oscules. Large oscules on apex and lateral margins of stoloniferous branches
Texture and surface characteristics. Surface
microconulose.
Ectosome and subectosome. Ectosomal skeleton
hispid, with points of choanosomal principal
styles protruding through surface individually or
in small bundles, piercing a sparse paratangential
layer of subectosomal auxiliary subtylostyles.
Choanosome. Choanosomal skeletal architecture
wide-meshed nearly renieroid reticulate, more
regular in peripheral skeleton than in axis, with
differentiated primary (ascending, multispicular)
and secondary (transverse, pauci- or unispicular)
spongin fibres; fibre diameter generally small
(primaries 30-501..im; secondaries 10-22p,m),
fibre anastomoses rectangular, and fibres cored
by choanosomal principal styles; coring spicules
in axial region heavier, more plumose (protruding
through fibres) than in peripheral skeleton;
echinating acanthostyles relatively sparse, evenly
distributed throughout skeleton; mesohyl matrix
light, usually containing choanosomal and/or
subectosomal megascleres dispersed between fibres.
Megascleres. Choanosomal principal subtylostyles slightly curved, fusiform, entirely smooth,
very slightly subtylote. Length 124-(152.4)165Rm, width 4-(8.8)-12Rm.
Subectosomal auxiliary subtylostyles straight,
thin, fusiform, with smooth slightly subtylote
2-(4.2)-61.m.
Microscleres. Palmate isochelae
unmodified. Length 448.6)12Rm long.
Accolada toxas very thin,
straight arms, sharply angular
central curvature, occasionally
raphidiform. Length 105220v,m, width 1-(2.2)-311,m.
REMARKS. Hallmann (1912)
noted that C. dura var. mollis
Hentschel was distinctly different from C. dura
Whitelegge and he considered that Hentschel's
(1911) species required a new name. The subspecific name 'moths' is preoccupied by C. monis
Kirkpatrick, 1903, and so a new name C.
squalorwn was proposed by Wiedenmayer (in
Hooper & Wiedenmayer, 1994). Some of the
characters and measurements of the lectotype
differ from those published by Hentschel (1911).
His description was presumably based on a series
of syntypes, but these have not been located or
re-examined.
Comparison between the type material of C.
(C.) squalorum and C. (Dendrocia) dura show
that both species are different in several significant respects: (1) differences in the geometry
of structural spicules where C. (C.) squalorum
has differentiated principal and auxiliary
spicules (and hence is referrable to C.
(Clathria)), and C. (D.) dura has completely undifferentiated structural megascleres (and hence
is referrable to C. (Dendrocia)); (2) palmate
isochelae versus arcuate-like isochelae; (3) long
thick accolada toxas versus no toxas; (4) nearly
renieroid reticulation of thin fibres versus a
regularly reticulate skeleton with heavy fibres,
respectively. In having a slightly renieroid skeletal architecture C. (C.) squalorum is reminiscent
of C. (lsociella) eccentrica, although spicule
geometry and spicule dimensions are otherwise
different.
Clathria (Clathria) striata Whitelegge, 1907
(Figs 73-74, Plate 1F, Table 13)
Clathria striata Whitelegge, 1907: 495-496, p1.45,
fig.27; Hooper,
& Wiedenmayer, 1994: 261.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
157
Thalysias striata; de Laubenfels, 1936a: 105.
Rhaphidophlus tenebratus Whitelegge, 1907: 501-
503, p1.45, fig.I9.
Clathria tenebrata; Hallmann, 1912: 211.
cf. Spongia sartaginula Lamarck, 1813: 383; 1814:
362.
MATERIAL. LECTOTYPE: AMG4344 (in part):
Near Coogee, Sydney, NSW, 34°05'S, 151°10'E, coll.
Hy 'Thetis' (dredge; label 'Clathria striata; type').
PARALECTOTYPES: AMG4344 (in part), Z823 (in
part): Off Botany Bay, NSW, 34°00'S, 151°11'E, 4046m depth, coll. FIV 'Thetis' (dredge; dry, two
specimens, label `Thalysias striata; cotypes').
HOLOTYPE of R. tenebratus: AMG4336: Off
Woolongong, NSW, 34°30'S, 150°50'E, coll. FIV
'Thetis' (dredge; dry, label 'Clathria tenebrata
Whitelegge; type'). OTHER MATERIAL: NSW QMG303755.
HABITAT DISTRIBUTION. 40-50m depth; on rock
substrate; central and S. coasts (NSW) (Fig. 73G).
DESCRIPTION. Shape. Flabellate, digitate or a
combination of both, 170-285mm long, 30110mm wide, with short, tapering, cylindrical
stalk, 35-55mm long, 6-13mm diameter, cylindrical or flattened branches and even, slightly undulating, ragged margins.
Colour. Live colouration unknown, dark greybrown preserved.
Oscules. Not observed on flabellate specimens,
but small oscules, up to 2mm diameter, seen on
lateral margins of branches for digitate specimen.
Texture and surface characteristics. Surface optically smooth, with prominent longitudinally
radiating subdermal grooves and low ridges.
Ectosome and subectosome. Ectosome strongly
hispid, with discrete, plumose, or paratangential
tufts of subectosomal auxiliary spicules surrounding protruding choanosomal principal
styles; principal styles in peripheral skeleton
usually larger than principal spicules within
fibres; ectosomal skeleton relatively dense in
places, merely paratangential or sometimes tangential to the surface in other places; thick tracts
of palmate isochelae mostly confined to dermal
and subdermal regions; subectosomal region
plumose, with diverging tracts of choanosomal
principal styles intermingled with extra-fibre
tracts of auxilairy styles, together producing relatively dense peripheral skeleton.
Choanosome. Choanosomal skeleton
predominantly renieroid but with differentiated
plumose and renieroid-reticulate components,
and clearly differentiated axial and extra-axial
regions but no well-marked separation of
primary or secondary fibre elements (cf.
Whitelegge, 1907); axial skeleton with very
heavy, thick spongin fibres forming relatively
tight ovoid meshes, with paucispicular core of
choanosomal styles forming vaguely ascending
skeletal tracts; extra-axial region with much
heavier, rectangular, almost renieroid spicule
skeleton, with a criss-cross of longitudinal and
transversely orientated spicules, and spongin
fibres lighter than in axis; longitudinal spongin
fibres in periphery cored by paucispicular tracts
of principal styles; transverse/ascending tracts
with multispicular tracts of spicules, many
protruding through fibres and forming plumose
brushes at right angles to surface; tendency for
some larger principal styles to form ascending
tracts, and small styles to occur mainly in longitudinal tracts; ultimate choanosomal spicule
tracts diverge into subectosomal region; echinating acanthostyles relatively sparse in axis, only
marginally more abundant in peripheral
skeleton; mesohyl matrix heavy, granular, with
few megascleres between fibres.
Megascleres (Table 13). Choanosomal principal
styles thick, curved, relatively variable in length,
fusiform, mostly sharply-pointed, less frequently
with rounded points, with smooth rounded, or
occasionally very slightly subtylote bases.
Subectosomal auxiliary subtylostyles thin,
fusiform, straight or slightly curved, occasionally
sinuous, usually with smooth, sometimes
microspined, slightly subtylote bases, occasionally polytylote bases.
Acanthostyles subtylote, with small and evenly
distributed spines.
Microscleres (Table 13). Palmate isochelae abundant, unmodified.
Toxas accolada, long, moderately thick, with
sharply angular or slightly angular central curvature, straight unreflexed points.
REMARKS. Rhaphidophlus tenebratus is a
synonym of Clathria striata, both species having
identical skeletal architecture, fibre characteristics and spicule geometry, although there is
some variation in spicule dimensions between
these two nominal species (Table 13). They differ
only in growth form (C. striata being flabellate,
R. tenebratus being digitate, with flattened and
anastomosing branches; Fig. 731). In his original
description Whitelegge (1907) omitted to mention toxas, abundant in the preserved specimens
but relatively rare in the dry holotype, and also.
that echinating acanthostyles were 0.6-0.8mm
long (typographical error for 0.06-0.08mm).
158zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 71. Clathria (Clathria) squalorum Wiedenmayer (holotype ZMB4444). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada toxa. E, Palmate
isochela. F, Section through peripheral skeleton. G, Australian distribution. H, Skeletal structure. I, Subrenieroid
reticulate fibres.
Clathria (C.) striata is similar to C. (C.) sartaginula (Lamarck) in shape, slightly renieroid
skeletal architecture, and to some extent spiculation. However, fibre characteristics differ
markedly between the two species, particularly
the density of coring and echinating spicules on
fibres, the degree to which peripheral fibres
radiate from the skeletal axis, and the very small
but thick, stumpy choanosomal styles in C. (C.)
sartaginula. This species is also compared with
C. (C.) arcuophora and C. (C.) biclathrata in
spicule geometry, spicule dimensions, and to
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
159
FIG. 72. Clathria (Clathria) squalorum Wiedenmayer (holotype ZMB4444). A, Choanosomal skeleton. B,
Ectosomal skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Base of
auxiliary subtylostyle. G, Accolada toxa. H, Palmate isochelae.
some extent fibre characteristics. Clathria (C.)
striata also has a similar skeletal architecture as
C. (C.) arcuophora, differing substantially only
in toxa geometry. All these species (C. (C.)
striata, C. (C.) arruophora, C. (C.) sartaginula,
C. (C.) crassa and C. (C.) biclathrata), are characterised by their slightly renieroid or subrenieroid choanosomal skeletons and spongin
fibres cored by few, thick principal spicules. They
are grouped together here in the striata' species
160zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 73. Clathria (Clathria) striata Whitelegge (lectotype AMZ4344). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada toxa. E, Palmate isochela. F,
Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Holotype of R. tenebratus
AMG4336.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
161
FIG. 74. Clathria (Clathria) striata Whitelegge (paralectotype AMZ823). A, Choanosomal skeleton. B,
Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines. F, Palmate
isochelae. G, Accolada toxas. H, Base of auxiliary subtylostyle.
162zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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TABLE 13. Comparison between present and published records of Clathria bundles superficially resem(Clathria) striata Whitelegge. All measurements are given in pm, denoted bling a Pseudaxinella (Axinelas range (and mean) of spicule length x spicule width (N=25).
lidae).
Colour. Pale yellow-brown
Lectotype
(AMG4344)
Paralectotypes
(N=2)
Subectosomal
auxiliary styles
1684268.3)-523
x 12-(17.7)-28
142-(245.6)-363
x 3-(4. 1)-6
184-(285.2)-496
x 14-(19)-24
153-(229.2)-294
x 3-(4.2)-6
Echinating
acanthostyles
71-(81.8)-93 x
6-(7.3)-9
75-(86)-94 x
7-(7.6)-9
SPICULE
Choanosomal
principal styles
Holotype of R.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB
preserved.
QMG303755
t enebrat us
Oscules. Not seen.
(AMG4336)
Texture and surface charac193-(282.2)-546 2094299.7)-462
Texture harsh, comteristics.
x
14-(16.5)-22
x 15-(19.1)-23
pressible; surface prominently
175-(274.4)-387 132-(181.4)-273
conulose, hispid.
x 2-(2.9)-5
x 3.5-(5.6)-8
Ectosome and subectosome.
Surface skeleton with protruding single principal styles em13-(15.1)-17
14-(16.4)-19
I4-(16.4)-20
14-(16.5)-21
Chelae
bedded
in peripheral skeleton
165-(224.3)-265 110-(211.0)-255 I 08-(202.4)-305 163-(221.8)-325
Toxas
and forming a plumose hispid
x 0.841.7)-3.0
x 0.8-(1.4)-2
x 1-(1.3)-2
x 0.8-(1.4)-2
surface; thick bundles of both
subectosomal auxiliary styles
group. Clathria (C.) striata differs from other
and
toxa
microscleres
surrounding protruding
striata' species in growth form, spicule
principal spicules, and also scattered paratangengeometry and spicule dimensions.
De Laubenfels (1936a) referred C. striata to tially across surface.
Thalysias but his decision is not supported. Al- Choanosonze. Skeletal architecture plumo-reticuthough there are relatively dense plumose ec- late, with slightly differentiated axial and extratosomal brushes composed of auxiliary axial regions; in peripheral region skeleton more
megascleres (structurally similar to Clathria plumose than reticulate, with skeletal columns
(Thalysias)) there is only one undifferentiated diverging but without connecting elements;
size category (142-38711m long) (composition of skeletal columns composed of erect multispicular
bundles of light spongin fibres fully cored by
Clathria (Clathria)).
choanosomal principal styles; spicules mostly
contained within fibres but also slightly protrudClathria (Clathria) toxipraedita Topsent, 1913
ing through fibres producing the plumose
(Figs 75-76)
('spicate) arrangement; towards base skeleton
more disorganised, predominantly reticulate, and
Clathria toxipraedita Topsent, 1913a: 620-621, p1.5, spicules more-or-less completely contained
fig.4, p1.6, fig.12; Burton, 1932a: 319; Burton,
1934b: 32, p1.4, figs 2-3, text-fig.3; Koltun, 1964a: within fibres; echinating acanthostyles moderate68-69, p1.12, figs 15-24; Koltun, 1976: 187; Hooper ly abundant; mesohyl matrix heavy, smooth, containing abundant microscleres.
& Wiedenmayer, 1994: 262.
Thalysias toxipraedita; de Laubenfels, 1936a: 105.
Megascleres. Choanosomal principal styles long,
Rhaphidophlus toxipraedita; Van Soest, 1984b: 115.
robust, straight or slightly curved near base, with
cf. Clathria toxipraedita; Sim & Byeon, 1989: 38 fusiform points and rounded or slightly subtylote
(Korea; possible misidentification).
smooth bases. Length 5184616.2)-894 width
19-(27.7)-33Rm.
MATERIAL. HOLOTYPE: RSME1921.143.1400: BurSubectosomal auxiliary subtylostyles short,
wood Bank, off Tierra del Fuego, S. Atlantic, 54°25'S,
57°32'W, 112m depth, 1.xii.1903, coll. R.R.V. very slender, straight, hastate pointed, subtylote,
with pointed (mucronate) or minutely spined
'Scotia' (dredge).
bases. Length 278-(310.2)-343p.m, width 4HABITAT DISTRIBUTION. 93-540m depth; on mud,
(4.8)-61.1,m.
sand and hard substrates; Australian Antarctic TerEchinating acanthostyles small, straight, long,
ritory: MacRobertson Land (Koltun, 1976) (Fig. 7511). tapering fusiform pointed, subtylote or tylote
Also Tierra del Fuego (Topsent, 1913a), South Georgia, Shag Rock (Burton, 1932a, 1934b), South bases, with small spines and aspinose bases and
Sandwich Is, Falkland Is (Koltun, 1964a), Palmer Ar- points. Length 102499.2)-1541m, width 4-(7.8)11 vm.
chipelago, Antarctica (Burton, 1932a).
Microscleres. Palmate isochelae in two size
DESCRIPTION. Shape. Massive encrusting, categories, both modified. Smaller chelae with
subspherical, 70mm long, 60mtn wide, up to relatively small alae and a small plate/ridge on
25mm thick, composed of fused irregular fibre central inner margin of shaft protruding between
84-(97.8)-112 x
6-(9.2)-11
58-(78.0)-92 x
4-(8.1)-11
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
163
the alae. Length 12-(15.7)-18Rm. Larger chelae
often cleistochelate, with teeth partially or completely fused and a large central plate/ridge on
inner margin of shaft between alae. Length 20(22.3)-24Rm.
Two categories of toxas. Longer toxas accolada, very long, thick, with sharply angular but
only slight central curvature, straight arms and
smooth fusiform points. Length 63541022.6)147011m, width 4-(5.2)-6Rm. Smaller toxas
wing-shaped, more widely curved at centre, occasionally looped/twisted at centre, with slightly
reflexed arms and points. Length 56485.3)132p,m, width 0.5-(1.1)-2.0pLm.
REMARKS. Clathria (C.) toxipraedita has
peculiar, large palmate cleistochelae resembling
sigmancistras (in some Mycalidae), with alae
varying from partially to completely fused, and
with the addition of a well formed central platelike ridge protruding between the alae (Fig. 76G).
Smaller chelae have smaller diffuse alae and a
smaller central plate-like ridge, indicating that
they are ontogenetic stages of the larger forms.
Cleistochelae are known in a number of other
poecilosclerids including a several microcionids
(e.g. C. (Microciona)cleistochela Topsent, C. (T)
ramosa (Dendy),Antho(Plocamia)signata (Topsent)), whereas the central plate-like ridge appears to be unique to the family and only
previously recorded from Mycalidae (E. Hajdu,
pers.comm.). Clathria toxipraedita is unusual
amongst the antiboreal species in having a great
diversity of spicule geometries; generally
microcionids from Antarctic waters appear to
have reduced spicule diversity in comparison to
the tropical fauna.
Clathria (Clathria) transiens Hallmann, 1912
(Figs 77-78, Plate 2A, Table 14)
Clathriatransiens Hallmann, 1912: 205, 226-234, 253,
254, p1.33, figs 1-3, p1.34, fig.2, text-figs 47-48;
Shaw, 1927:426; Burton, 1934a: 599; Guiler, 1950:
7; Burton, 1959a: 244; Wiedenmayer, 1989: 57,
p1.5, fig.8, p1.23, fig.6, text-fig.38; Hooper &
Wiedenmayer, 1994: 262.
Thalysias transiens; de Laubenfels, 1936a: 105.
cf. Microciona prolifera, tropus stylota and tropus
senta, Vosmaer, 1935a: 611, 649-650, 666.
MATERIAL. LECTOTYPE: AME302: Off Devonport, N. coast Tas, 41°11'S, 146°21'E, coll. FIV
'Endeavour' (dredge) (label 'Clathria transiens; typical
form a'). PARALECTOTYPES: AMZ49 + E819:
64km W. of Kingston, SA, 36°50'S, 139°05'E, 60m
depth, coll. FIV 'Endeavour' (dredge) (specimen cut in
half, form b). AME779: same locality; (form c).
AMZ744: Port Phillip Bay, Vic, 38°09'S, 144°52'E,
coll. J.B. Wilson (dredge) (label 'Clathria transita
Hallmann', AM register ref. 342/85, JBW sp.3, cotype,
form d). (uncertain type status - AMZ743: Port Phillip
Bay, Vic., 38°09'S, 144°52'E, coll. J.B. Wilson;
(dredge; label 'Clathria transita Hallmann', dry,
`type')). OTHER MATERIAL: VIC. - NCIQ66C3231-C (fragment NTMZ3694). TAS. - QMG300268
(NCIQ66C-3638-U, fragment NTMZ3802).
HABITAT DISTRIBUTION. Shallow subtida1-60m
depth; on rock substrates; Port Phillip (Vic) (Hallmann,
1912; present study), Furneaux Is and Devonport (Tas)
(Shaw, 1927; Guiler, 1950; Wiedenmayer, 1989;
present study), Kingston (SA) (Hallmann, 1912), Qld.
(Burton, 1934a) (Fig. 77G). Burton (1959a) recorded
the species from the Red Sea but his material was not
described, and his record is questionable.
DESCRIPTION. Shape. Usually arborescent
with short, bulbous branches, 75-185mm long,
40-160mm wide; stipitate with short cylindrical
stalk, 25-55mm long, 8-22mm diameter;
branches simply bifurcate, or repeatedly bifurcate and arborescent, flabello-digitate, or they
may anastomose to form thickly clathrous, bushy
lobes.
Colour. Live colouration vermilion red (Munsell
5R 3/10), beige-brown or dark brown preserved.
Oscules. Small oscules, up to 3mm diameter, in
ridges and grooves of branches.
Texture and surface characteristics. Surface
markedly conulose, with conules actually being
rounded vestigial branches; with or without irregular ridges striating surface; surface covered
by distinct membraneous skin-like ectosome.
Ectosome and subectosome. Ectosome minutely
hispid, with the points of larger choanosomal
principal styles protruding through membraneous
surface, with a paucispicular, usually tangential
layer of subectosomal auxiliary subtylostyles,
lying on or just below surface; occasionally
auxiliary styles form plumose brushes, surrounding 1 or more principal style; subectosomal skeletal structure vaguely plumose, formed by
diverging peripheral spongin fibres and spicule
tracts of smaller choanosomal principal styles.
Choanosome. Choanosomal skeletal architecture
almost regular, radial-renieroid, with poorly differentiated axial and extra-axial skeletons; axis
formed by evenly anastomosing spongin fibres,
forming relatively cavernous, rectangular or circular meshes; spongin fibres cored by pauci- or
multispicular tracts of smaller choanosomal principal styles; primary and secondary skeletal tracts
in axis vaguely differentiated, but not
pronounced, and spongin fibre elements only dif-
^ MEMOIRS OF THE QUEENSLAND MUSEUM
164zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
A
EzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
0
0
CV
FIG. 75. Clathria(Clathria)toxaepraedita Topsent (holotype RSME1921.143.1400). A, Choanosomal principal
styles and modifications. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Accolada
toxas. E, Wing-shaped toxas. F, Modified palmate isochela. G, Section through peripheral skeleton. H,
Antarctica distribution. I, Holotype. J, Choanosomal structure.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
165
FIG. 76. Clathria(Clathria)toxaepraedita Topsent (holotype RSME1921.143.1400). A, Choanosomal skeleton.
B, Peripheral skeleton. C, Fibre characteristics. D, Base of auxiliary subtylostyle. E, Echinating acanthostyle.
F, Acanthostyle spines. G, Transitional series from cleistochelae to modified palmate isochelae. H, Accolada
and wing-shaped toxas. I, Accolada toxas.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
166zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
A
Clat hr ia ( Clat hr ia) t r ansiens Hallmann (lectotype AME302). A, Choanosomal principal styles. B,
FIG. 77. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Subectosomal auxiliary subtylostyle. C, Echinating acanthostyles. D, Oxhorn toxas. E, Palmate isochelae. F,
Section through peripheral skeleton. G, Australian distribution. H, lectotype.
ferentiated by density of coring spicules; extraaxial skeletal architecture regularly renieroid,
with ascending plumo-reticulate primary and
secondary fibre elements which diverge increasingly towards surface; fibre anastomoses much
closer toward periphery than at core; primary
ascending fibres heavily multispicular, secondary
transverse fibres more irregular, pauci- or multispicular; all fibres relatively heavily invested
with spongin; echinating acanthostyles abundant
on all fibres, only slightly more dense at fibre
nodes; mesohyl matrix light, containing
numerous subectosomal styles and microscleres
scattered between fibres.
Megascleres (Table 14). Choanosomal principal
styles divided into 2 (overlapping) size classes:
larger found in brushes protruding from surface;
smaller coring choanosomal spongin fibres; both
similar in geometry, fusiform, straight or slightly
curved, with smooth, rounded or very slightly
subtylote bases.
Subectosomal auxiliary subtylostyles straight,
thin, fusiform, with smooth subtylote bases, variable length but only 1 size class.
Acanthostyles slightly subtylote, very sharply
pointed, with granular, vestigial, evenly dispersed spines, sometimes completely smooth.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
167
FIG. 78. Clathria (Clathria) transiens Hallmann (paralectotype AMZ743). A, Choanosomal skeleton. B, Fibre
characteristics (x306). C, Echinating acanthostyles. D, Vestigial acanthostyle spines. E, Echinating spicules in
situ. F, Accolada toxa. G, Palmate isochelae.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
168zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Microscleres (Table 14). Palmate isochelae large,
unmodified, rare in some material.
Toxas oxhorn, relatively thick, with rounded,
large or small central curvature, slightly reflexed
or straight arms.
Larvae. Larvae viviparous, ovoid or elongate
parenchymella, up to 210p,m diameter, with larval megascleres dispersed throughout axis.
REMARKS. The status of specimen AMZ743
(from Port Phillip Bay) is not completely certain.
The label states that it is a 'dry, type', but the four
type specimens described by Hallmann (1912)
are accounted for by other AM register numbers
(cited above). It is possible that the specimen is a
fragment of the paralectotype (AMZ744), as both
come from the same locality.
Clathria (C.) transiens is relatively well known
from SE. Australian waters (e.g. Wiedenmayer,
1989). The species appears to be the temperate
water equivalent of the widely distributed tropical species C. (Thalysias) lendenfeldi (see
Hooper et al., 1990). Both species have a large
range of growth forms and comparable surface
features, although this resemblance is superficial,
in the field, and the two species differ markedly
in most other features. The ectosomal features of
C. (C.) transiens (with protruding choanosomal
styles), and fibre characteristics (almost a subrenieroid peripheral skeleton), are also reminiscent of C. (T) vulpina, although growth form and
spicule diversity differ significantly between
these two species. Clathria (T) darwinensis
sp.nov. from northwest Australia is similar to C.
(T) transiens in ectosomal features, vaguely subrenieroid skeletal architecture, and toxa morphology, but differs in acanthostyle geometry,
possession of 2 size classes of auxiliary styles,
gross morphology and spicule dimensions.
Despite a relatively variable growth form the
skeletal architecture of this species is consistent.
There are only minor differences between
specimens in the degree to which the axial skelton
is compressed and differentiated from extra-axial
structures (e.g. compare Hallmann's (1912)
'form a' and 'form c'). Similarly, in some places
on the ectosomal skeleton are distinct plumose
brushes of auxiliary spicules, radiating from
bases of protruding choanosomal principal styles,
whereas in other places ectosomal structure is
simply tangential. This plumose structure is
generally associated with the ectosomal skeleton
characteristic of Clathria (Thalysias) species, but
only a single size class of auxiliary styles is
TABLE 14. Comparison between present and published records of Clathria (Clathria) transiens Hallmann.
All measurements are given in p.m, denoted as range
(and mean) of spicule length x spicule width (N=25).
SPICULE
Lectotype
(AM E302)
Paralectotypes
(N=4)
Specimens
(N=2)
Large
choanosomal
principal
styles
273-(339.4)409 x 12(16.8)-19
264-(359.7)528 x 13(16.1)-19
2694408.2)632 x 9-(13.8)19
Small
choanosomal
principal
styles
133-(l 88.4)24 x 3 8(13.2)-18
113-(170.3)247 x 6-(9.5)13
1 32-(191.8)275 x 8411.2)13
Subectosomal
auxiliary
styles
162-(272.4)357 x 2.5(4.0)-5.5
96-(249.2)416x 1.5(3.6)-6
142(277.0)402x 1.5(4.2)-6
42-(71.1)-98 x
2-(4.9)-7.5)
63-(76.2)-92 x
3.5-(5.1)-7
15-(17.8)-20
14-(18.4)-22
I4-(19.2)-22
17-(84.6)-131
x 0.8-(1.5)-3
18-(109.1)192 x 0.8(1.8)-3.5
7I-(108.4)134 x 0.8(1.1)-1.8
58-(70.7)-88 x
Echinating
acanthostyles
3-(4.9)-6.5
Chelae
Toxas
present in C. (C.) transiens and it technically
belongs to Clathria (Clathria).
Clathria (Clathria) wilsoni Wiedenmayer,
1989
(Figs 79-80)
Clathria wilsoni Wiedenmayer, 1989: 57-58, p1.5,
fig.9, p1.24, fig.1, text-fig.39; Hooper & Wiedenmayer, 1994: 262.
Rhaphidophlus wilsoni; Carpay, 1986: 27.
MATERIAL. HOLOTYPE - NMVF51967: Garden
Cove, N. side of Deal I., Kent Is Group, Bass Strait,
Tas, 39°29'S, 147°20'E, 10m depth, 25.ii.1981, coll.
F. Wiedenmayer et al. (SCUBA).
HABITAT DISTRIBUTION. 10m depth; on algal
covered boulders; Bass Strait and E. coast (Tas) (Fig.
79H).
DESCRIPTION. Shape. Small sponge, thickly
encrusting with irregular low-formed branches,
5mm high, up to 20mm diameter.
Colour. Live colouration dull vermilion, beige
preserved.
Oscules. Oscules small, up to 2mm diameter,
scattered on base and sides of branches.
Texture and surface characteristics. Surface uneven, microconulose, hispid, particularly on
points of branches. Texture easily compressible,
spongy.
Ectosome and subectosome. Ectosome skeleton a
layer of subectosomal auxiliary styles, in small
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
169
D
FIG. 79. Clathria (Clathria) wilsoni Wiedenmayer (holotype NMVF51967). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Accolada toxa. E, Oxhorn toxa. F, Palmate
isochela. G, Section through peripheral skeleton. H, Australian distribution. I, Choanosomal skeletal structure.
J, Holotype.
bundles, erect on surface, with acanthostyles styles, embedded in peripheral fibres, form
echinating peripheral fibres also contributing to plumose brushes, which approach but usually do
ectosomal skeleton; choanosomal principal not pierce surface.
MEMOIRS OF THE QUEENSLAND MUSEUM
170^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 80. Clathria (Clathria) wilsoni Wiedenmayer (holotype NMVF51967). A, Choanosomal skeleton. B,D,
Echinating acanthostyles. C,E, Acanthostyle spines. F, Base of principal style., G, Base of auxiliary styles. H,
Palmate isochelae. I, Larger accolada toxa. J, Smaller oxhorn toxas.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVU
171
Choanosome. Choanosomal skeleton irregularly
reticulate, consisting of moderately light, slender
spongin fibres, up to 35pm diameter, forming
irregular rectangular or polygonal meshes; spongin fibres cored by multispicular tracts of
choanosomal styles, in rows of 2-10 spicules
abreast, echinated by regularly dispersed acanthostyles; mesohyl matrix light, many vestigial
and fully formed subectosomal styles, and
numerous rhaphidiform toxa microscleres.
Megascleres. Choanosomal principal styles
slightly curved, with evenly rounded or slightly
subtylote, smooth or rarely microspined bases.
Length 164-(191.8)-290p,m, width 2.445.1)Subectosomal auxiliary subtylostyles straight,
occasionally slightly curved, with tapering or
rounded, smooth bases. Length 834153.6)282p,m, width 1-(2.3)-3.5p,m.
Acanthostyles short, thick, subtylote, with relatively evenly dispersed spines. Length 31.6(62.6)-71 p.m, width 2.6-(6.6)-10Rm.
Microscleres. Palmate isochelae very rare, small,
poorly silicified. Length 4-(5.5)-9p,m.
Toxas accolada, straight, very thin
rhaphidiform, occasionally with roughened
points; some smaller examples slightly curved,
with reflexed arms. Length 71-(121.6)-168p,m,
width 0.4-(0.7)-2p.m.
REMARKS. This species is transitional between
the ectosomal structures of Clathria and
Thalysias, and it is not surprising that Carpay
(1987) referred it to the latter. However, although
there is a large size range of auxiliary styles,
ectosomal spicule brushes are composed only of
a single size of spicule, and the species is technically correctly included in Clathria (Clathria).
Nevertheless, this example further illustrates the
inappropriateness of recognising a strict
(phylogenetic) boundary between these two
genera.
Clathria (C.) wilsoni is peculiar in having almost straight raphide-like toxas, which are
known to occur in only a few microcionids (e.g.
C. (Thalysias) costifera and C. (Wilsonella)
reticulata, both from Australia, C. (T.) juniperina
from the Caribbean, C. (C.) microxea from
Madagascar, and Artemisina archegona from the
Pacific coast of America). Wiedenmayer (1989)
notes some similarities between C. (C.) wilsoni
and other species of Hallmann's (1912) spicata'
group, but this is not upheld here.
OTHER SPECIES OF CLATHRIA
(CLATHRIA)
Clathria (Clathria)acanthostyli (Hoshino, 1981)zyxwvutsrqpo
Thalysias acant host yli Hoshino, 1981: 156-157, text-fig.68,
p1.7, fig.2 [Uchinoura, Japan].
MATERIAL. HOLO'TYPE: MMBSSAT-020. Japan.
Clathria (Clathria) acanthotoxa (Stephens, 1916)
Eurypon acant hot oxa Stephens, 1916: 238-239 [SW coast,
Ireland].
Microciona acant hot oxa; Stephens, 1921: 50, p1.5, fig.!:
Alander, 1942:62 [Sweden]; Burton, 1959b: 43 [Iceland];
Levi, 1960a: 75 [Isle of Man, W. Ireland].
Not Microciona acant hot oxa; Lilly et al., 1953: 97.
Dict yociona acant hot oxa; de Laubenfels, 1936a: 110 [note];
de Laubenfels, 1953: 527.
Clat hr ia acant hot oxa; Van Soest, 1984b: 7 [generic
synonymy for Dict yociona) .
Not Microciona acant hot oxa Levi & Levi, 1989: 81, fig. 49
[see C. claudei, nom.nov.].
MATERIAL. HOLOTYPE: INM5R253.8.1916. NE. Atlantic,
North Sea.
Clathria (Clathria) anchorata (Carter, 1874)
Dict yocylindrus anchorat us Carter, 1874: 251 [Gulf of
Manaar, Ceylon].
Clat hria anchorat a; Vosmaer, 1880: 153 [Atlantic]; [?]
Stephens, 1916:242 [W coast, Ireland]; Burton, 1959b: 42
[Iceland]; Levi, 1960a: 63-64 [English Channel, Atlantic];
[7] Vacelet, 1969: 206, text-fig.45 [Mediterranean].
Microciona anchorat a; Alander, 1942: 62 [Sweden].
Echinonem a anchorat um ; Dendy, 1889a: 44.
Wilsonella anchorat um ; Hallmann, 1912: 152, 185, 189, 210,
211, 243, 296, 298, 299.
Cionanchora anchorat a; de Laubenfels, 1936a: 108 [note].
Not Echinonem a anchorat um Carter, 1881a: 362, 379, 380 [S.
coast of Australia].
Not Echinonem a anchorat um , varr.; Lendenfeld, 1888: 219220 [Port Jackson, New South Wales].
Not Echinonem a anchorat um var. ram osa; Whitelegge, 1901:
81.
Clat hria t ypica, in part; Dendy, 1896: 32; Kirkpatrick, 1903:
248-249.
Clat hria longichela Topsent, 1928a: 300, p1.10, fig.9.
cf. Microciona prolifera; Vosmaer, 1935a: 610, 632, 668.
MATERIAL. HOLOTYPE: LFM destroyed, fragmentsBMNH1953.11.11.63-69. NE. Atlantic, Mediterranean, Gulf
of Manaar.
Clathria (Clathria) antyaja (Burton & Rao,
1932)
Dendrocia ant yaj a Burton & Rao, 1932: 348-350, p1.28,
fig.12, text-figs 13-14 [Indian Ocean]
MATERIAL. HOLOTYPE: IMP791/1. Indian Ocean.
Clathria (Clathria) anthoides Levi, 1994
Clat hria ant hoides Levi, 1994: 36-37, fig. 21A, p1.7, fig.5
[Norfolk Rise, 610m depth].
MATERIAL. HOLOTYPE: MNHNDCL3637. SW. Pacific.
Clathria (Clathria) arbuscula (Row, 1911)
Row, 1911:347-349, p1.39, fig.22,
p1.40, fig.25, text-fig.22 [Red Seat
Ophlit aspongia arbuscula
Ophlit aspongia horrida Row, 1911: 349-351, p1.40, fig.26,
text-fig.23 [Red Seat
172zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Litaspongia arbuscula; de Laubenfels, 1954: 162 [note].
Kerasemna arbuscula; Pulitzer-Finali, 1982: 105.
Kerasemna horrida; Pulitzer-Finali, 1982: 105.
MATERIAL. HOLOTYPE: BMNH1912.2.1.63 (0. arbuscula); BMNH1912.2.1.65 (0. horrida). Red Sea.
Clathria (Clathria) asodes (de Laubenfels,
1930)
Eutypon asodes de Laubenfels, 1930: 27; de Laubenfels,
1932: 92-93, text-fig.54 [Carmel,California]; Lee &
Gilchrist, 1985: 24-32 [biochemistry].
Dictyociona asodes; de Laubenfels, 1936a: 110 [note].
Clathria asodes; Van Soest, 1984b: 7 [generic synonymy].
Leptoclathria asodes; Sim & Bakus, 1986: 10 [California];
Bakus & Green, 1987: 72 [S. California].
MATERIAL. HOLOTYPE: USNM21442. NE. Pacific.
Clathria (Clathria) atoxa (Bergquist &
Fromont, 1988)
Dictyociona atoxa Bergquist & Fromont, 1988: 105-106,
p1.49, figs a-c [Tokatu]; Dawson, 1993: 38 [note].
MATERIAL. HOLOTYPE: NMNZPOR116. New Zealand.
Clathria (Clathria) axociona Levi, 1963
Clathria axociona Levi, 1963: 49, text-fig.56, p1.8A [Cape of
Good Hope, South Africa]; Uriz, 1988a: 83-84, pls 20-21a,
text-fig. 59 [Namibia].
MATERIAL. HOLOTYPE: MNHNDCL632. SW. and S.
Africa.
Clathria (Clathria) barleei (Bowerbank, 1866)
Isodiciya barleei Bowerbank, 1866: 333; Bowerbank, 1874:
p1.57.
Tragosia barleei; Topsent, 1894a: 25.
Axinella barleei; Arndt, 1935: 88, fig.186.
Clathria barleei; Van Soest & Stone, 1986: 45-46, fig.4
[Norway]; Ackers, Moss & Picton, 1992: 138 [Ireland].
Halichondria foliata Bowerbank, 1874: 198, p1.73, figs 1-5;
Carter, 1876: 310, p1.12, fig.10, p1.29, fig.29; Bowerbank,
1882: 106.
Esperia foliata; Fristedt, 1885: 41.
Homeodictya foliata; Topsent, 1894a: 12.
Echinoclathria foliata; Topsent, 19 13b: 38; Stephens, 1916:
234 [Ireland]; Stephens, 1921: 57; Hentschel, 1929: 894,
971 [Arctic].
Artemisina foliata; Burton, 1930a: 501, 529, p1.2, figs 1-2
[Norway]; Burton, 1959b: 42-43 [Iceland].
Amphilectus foliatus; Vosmaer, 1880: 118.
Antho foliata; Alander, 1942: 63 [Sweden; defined in
Esperiopsidae].
? Halichondria mutula Bowerbank, 1874: 209, p1.74,
figs 4-8; Bowerbank, 1882: 96 [as H. mutulus].
? Amphilectus mutulus; Vosmaer, 1880: 118.
Isodiciya laciniosa Bowerbank & Norman, 1869: 333.
Amphilectus laciniosus; Vosmaer, 1880: 116.
Clathria laciniosa; Arndt, 1935: 81, fig.167; Burton,
1959b: 42 [Iceland].
Art emisina laciniosa; Burton, 1950: 891 [revision].
MATERIAL. HOLOTYPE: BMNI-11930.7.3.338. Arctic, NE.
Atlantic.
Clathria (Clathria) basilana Levi, 1961
Clathria basilana Levi, 1961b: 520, fig.11 [Zamboanga,
Philippines]; Van Soest, 1989: 1-2, fig.6 [Lesser Sumba Is,
Indonesia].
MATERIAL. HOLOTYPE: MNHNDCL722. S. Philippines
- E. Indonesia.
Clathria (Clathria) borealis sp.nov.
Clathria robusta Koltun, 1959: 186, p1.25, fig.5, text-fig. 147
[Arctic, USSR]; Van Soest & Stone, 1986: 47 [note].
Not Microciona strepsitoxa var. robusta Dendy, 1922: 60.
Not Tenacia robusta; Burton & Rao, 1932: 339-340.
MATERIAL. HOLOTYPE: ZIL, no fragment in BMNH.
Note: C. (C.) robusta (Dendy, 1922) has priority. Arctic.
Clathria (Clathria) bulbosa Hooper & Levi,
1993
Clathria (Clathria) bulbosa Hooper & Levi, 1993a: 12681270, figs 23-24 [Chesterfield Is, Coral Sea].
MATERIAL. HOLOTYPE: QMG300021. SW. Pacific.
Clathria (Clathria) burtoni sp. nov
Clathria proltfera Burton, 1940: 109, p1.4, figs 3-4, text-fig.2
[Argentina].
Not Spongia prolifera Ellis & Solander, 1786: 189.
MATERIAL. HOLOTYPE: MA BA 15582, fragmentsBMNH1934.1.17.13, 116, 117. Note: C. prolifera (Ellis &
Solander, 1786) has seniority. Province: SW. Atlantic.
Clathria (Clathria) caespes (Ehlers, 1797), unrecognizable.
Spongia tubulosa, in part, Esper, 1797: 196, p1.44 [Cape of
Good Hope, South Africa].
Scopalina caespes; Ehlers, 1870: 19-20, 31.
MATERIAL. HOLOTYPE: unknown. S. Africa.
Clathria (Clathria) calypso Boury-Esnault,
1973
Clathria calypso Boury-Esnault, 1973: 286, text-fig.47
[Brazilian Basin]
MATERIAL. HOLOTYPE: MNHNDNBE1035. tropical SW.
Atlantic.
Clathria (Clathria) cantabrica (Orueta, 1901)
Rhaphidophlus filifer var. cantabrica Orueta, 1901: 331-335,
pls 3,4 [Bay of Biscay].
Tenacia filifer var. cantabrica; Hallmann, 1920: 771.
Clathria cantabrica; Levi, 1960a: 55-56, 63.
MATERIAL. HOLOTYPE: Madrid. NE. Atlantic.
Clathria (Clathria) compressa Schmidt, 1862
Clathria compressa Schmidt, 1862: 8, 10, 58-59, 86, p1.6,
fig.1; Schmidt, 1864: 35, p1.4, fig.3; Crivelli, 1863: 299;
Gray, 1867: 513; Claus, 1868: 23; Carter, 1875: 195;
Vosmaer, 1880: 150 [Triest]; Graeff, 1882: 318; Vosmaer,
1884: 119; Ridley, 1884a: 443-449, 612-615; Schmidt,
1880: 34-35, 45, p1.4, fig.3; Ridley & Dendy, 1887: 147;
Dendy, 1889c: 8; Hope, 1889: 337; Norman, 1892: 13;
Topsent, 1892a: 17; Topsent, 1894a: 18; Heider, 1895:
280; Topsent, 1896: 123; Thiele, 1899: 13; Topsent, 1911:
10, 13; Dendy, 1922:64; Dendy, 1924a: 352-354; Wilson,
1925: 439; Topsent, 1925: 647-650, fig.8 [Banyuls, Gulf
of Naples]; Topsent, 1928a: 62,299; Burton & Rao, 1932:
334-337; Burton, 1932a: 319; Burton, 1934a: 558; Topsent
& Olivier, 1943: 1 [Monaco]; Koltun, 1959: 184; Levi,
1960a: 50, 52, 61, 62 [Mediterranean, Atlantic]; Levi,
1960b: 761, fig. 14 [Kayar, W. Africa]; Melone, 1963: 1-8;
Sara & Melone, 1963: 362; Sara, 1964: 229; Simpson,
1968a: 102, 104-106, p1.17, table 49; Boury-Esnault, 1971:
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
173
323 [Banyuls]; Pulitzer-Finali, 1983: 610; DesqueyrouxFaundez & Stone, 1992: 9, 35, 103 [list].
Thalysias compressa; de Laubenfels, 1936a: 105-106.
? Spongia clathrus Esper, 1797; Vosmaer, 1880: 150.
Not Citalina compressa; Carter, 1882b: 112-113.
Not Halichondria compressa Carter, 1886g: 450
[Westemport Bay, Victoria].
Not Pseudoclathria compressa; Dendy, 1897: 259.
MATERIAL. HOLOTYPE: LMJG15509, fragments
BMNH1867.7.26.78, BMNH1910.1.1.2362, 2363. Mediterranean, NE. Atlantic, W. Africa.
Clathria (Clathria) conica Levi, 1963
Clathria conica Levi, 1963: 50-51, text-fig.57, p1.8E [Cape of
Good Hope].
MATERIAL. HOLOTYPE: MNHNDCL617. S. Africa.
Clathria (Clathria) contorta (Bergquist &
Fromont, 1988)
Dictyociona contorta Bergquist & Fromont, 1988: 105, p1.48,
figs d-f [Manukau]; Dawson, 1993: 38 [note].
MATERIAL. HOLOTYPE: NMNZPOR115. New Zealand.
Clathria (Clathria) coralloides (Olivi, 1792)
Spongia coralloides Olivi, 1792: 264; Bertoloni, 1819: 228;
Blainville, 1819: 125; Lamouroux, 1824: 369; Martens,
1824: 535.
Grantia coralloides; Nardo, 1833: 522.
Halichondria corona Lieberkiihn, 1859: 521, 529, p1.11, fig.3.
Clathria coralloides , Schmidt, 1862:7, 10, 11, 58, 85, p1.5,
figs 10-11; Kiilliker, 1864: 52, 71; Schmidt, 1864: 34;
Schmidt, 1868: 9,41; Schmidt, 1870: 56, 60; Gray, 1867:
513, 533, 552; Dybowsky, 1880: 5, 70, p1.4, figs 7,9;
Krukenberg, 1880: 70, 72; Vosmaer, 1880: 149-150
[Mediterranean]; Ridley, 1881: 485; Vosmaer, 1881: 4;
Vosmaer, 1882-6: 119, 121-122; Vosmaer, 1884b: 492;
Vosmaer, 1885a: 186, 356; Carter, I882a: 281; Graeffe,
1882: 318; Carter, 1884a: 204; Ridley & Dendy, 1887:
147; Lendenfeld, 1889a: 410-415, 498-505, 586, 592, 594,
598, 602, 604, 608, 612, 614, 618, 622, 624, 628, 634, 640,
644, 646, 650, 669, p1.34, figs 193-205; Lendenfeld,
1890a: 72; Maas, 1892:427-428; Norman, 1892: 13; Maas,
1893: 331, 334-335, 414, 441, 444, p1.20, fig.19, p1.21,
fig.32; Topsent, 1893d: 445; Topsent, 1894a: 18,24; Topsent, 1894b: 35; Heider, 1895: 280; Dendy, 1896: 36;
Loisel, 1898: 38; Zimmermann, 1907: 308; Korschelt &
Heider, 1910: 321; Walther, 1910: 21; Babic, 1921: 84
[Adriatic]; Babic, 1922: 244; Topsent, 1925: 646-647,
fig.7 [Gulf of Naples]; Topsent & Olivier, 1943: 1
[Banyuls and Naples]; de Laubenfels, 1954: 139; Levi,
1960a: 61-62 [Mediterranean, Adriatic,Naples, Banyuls];
Laubier, 1966 [Banyuls]; Vidal, 1967 [Mediterranean];
Boury-Esnault, 1971: 323 [Banyuls]; Pulitzer-Finali,
1983: 568-569, 610 [Mediterranean]; Boury-Esnault &
Lopes, 1985: 194-195, fig.43 [Azores]; Pansini, 1987
[Adriatic Sea]; Desqueyroux-Faundez & Stone, 1992: 9,
35, 103 [list].
Ophlitaspongia coralloides; Bowerbank, 1874: 10.
Ophlitaspongia corona; Levi, 1960a: 61
MATERIAL. HOLOTYPE: unknown; fragments LMJG
15356, BMNH1867.7.26.74, 1910.1.1.2364, 2365. Mediterranean, NE. Atlantic.
-
Clathria (Clathria) curvispiculifera (Carter,
1880)
Microciona curvispiculifera Carter, 1880a: 43,44,151,153,
p1.4, fig.6a-d [Gulf of Manaar]; Vosmaer, 1935a: 608.
? Rhabdoploca curvispiculifera; Topsent, 1904a: 157-159.
MATERIAL. HOLOTYPE: LFM missing, no fragment in
BMNH. Gulf of Manaar. Note: virtually unrecognisable.
Clathria (Clathria) dayi Levi, 1963
Clathria dayi Levi, 1963:51, p1.8B, text-fig.58 [Cape of Good
Hope]; Sim & Byeon, 1989: 39, p1.4, figs 1-2 [Korea;
possible misidentification].
MATERIAL. HOLOTYPE: MNHNDCL611. S. Africa.
Clathria (Clathria) depressa Sara & Melone,
1966
Clathria depressa Sara & Melone, 1966: 2-4, text-figs 1-2,
p1.1 [Olivetta, Portofino, Levante coast, Italy]; PulitzerFinali, 1983: 610.
MATERIAL. HOLOTYPE: IMZUB. E. Mediterranean.
Clathria (Clathria) discreta (Thiele, 1905)
Microciona discreta Thiele, 1905: 447-449, 494, p1.3 I , fig.
65a-e [Calbuco, Chile].
Diciyociona discreta; Topsent, 1913a: 580, 583, 614, 618620, 642, p1.3, fig.5 [Gough I., S. Atlantic Ocean]; Burton,
1932: 324, p1.56, figs 3-4 [Falklands]; Burton, 1940: 112,
p1.4, figs 1-2, p1.6, fig.2 [Argentina and Chile]; de Laubenfels, 1953a: 527; Desqueyroux, 1972: 29-30, figs 95-102,
136-137 [Chile]; Desqueyroux & Moyana, 1987: 49
[Chile, Tierra del Fuego, Argentina, Falkland Is, Kerguelen Is].
cf. Microciona prolifera; Vosmaer, 1935a: 608,646.
MATERIAL. HOLOTYPE: ZMB3302, fragment:
BMNH1908.9.24.159. SW. Atlantic, SE. Pacific, Subantarctic.
Clathria (Clathria) elastica Levi, 1963
Clathria elastica Levi, 1963: 52, text-fig.59, p1.8C [Cape of
Good Hope].
Not Clathria elastica Sara, 1978: 70 [see C. sarai, nom.novi.
MATERIAL. HOLOTYPE: MNHNDCL604. S. Africa.
Clathria (Clathria) elegans Vosmaer, 1880
Clathria elegans Vosmaer, 1880: 152 [North America]; Vosmaer, 1884b: 492.
? Anaata elegans; de Laubenfels, 1936a: 109.
Not Plectispa elegans Lendenfeld, 1888: 226.
Not Antherochalina elegans Lendenfeld, 18876: 787.
cf. Microciona prolifera; Vosmaer, 1935a: 630.
MATERIAL. HOLOTYPE: unknown. NE. Pacific. Note:
virtually unrecognisable.
Clathria (Clathria) foliacea Topsent, 1889
Clathria foliacea Topsent, 1889: 39-40, text-fig.5 [Banc de
Campeche]; Van Soest, 1984b: 107, table 4; Van Soest &
Stentoft, 1988: 126 [table].
Thalyseurypon foliacea; de Laubenfels, 1936a: 107-108
[Florida].
Pandaros foliaceum; Wiedenmayer, 1977: 144 [note].
Clathria carteri Topsent, 1889: 38-39, text-fig.4B [Banc de
Campeche]; Van Soest, 1984b: 108.
Thalyseurypon carteri; de Laubenfels, 1936a: 107.
Pandaros carteri; Wiedenmayer, 1977: 144 [note].
MATERIAL. HOLOTYPE: MNHN missing, reference
specimen USNM22516. NE. Atlantic, Caribbean.
Clathria (Clathria) foliascens Vacelet & Vasseur, 1971
MEMOIRS OF THE QUEENSLAND MUSEUMzyxwvutsrqponmlkjihgfedcbaZYXWVUT
174^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
fig.49, p1.4,figs 5-6 [Tulear Madagascar]; Vacelet & Vasseur, 1977: 114; Vacelet et al., 1976: 71 [Tulear,
Madagascar]
MATERIAL. HOLOTYPE: MNHNDJV30. W. Indian
Ocean.
MATERIAL. HOLOTYPE: MNHN missing (fragment
BMNH1954.2.20.85, unconfirmed); 'representative
specimen': BMNH1926.2.19.2. Note: Burton's species is
probably the same as Lamarck's but this cannot be verified
and the species is taken in the sense of Burton (1931a). South
Africa.
Clathria (Clathria) frondiculata (Schmidt,
1864)
Clathria (Clathria) labyrinthica (Schmidt,
1864)
Clat hria foliascens Vacelet & Vasseur, 1971: 95-96, text-
Reniera ( ?) frondiculat a Schmidt, 1864: 39, 40, 45, p1.4,
fig. 10 [Adriatic].
Pit alia frondiculat a; Gray, 1867: 524.
Clat hriafrondiculat a; Vosmaer, 1880:153 [Triest]; Vosmaer,
1885:236; Norman, 1892: 13; Topsent, 1892b: 23; Heider,
1895: 282; Burton, 1930a: 514.
cf. Clat hria com pressa and C. coralloides; Vosmaer, 1935a:
625 [intermediate form].
MATERIAL. HOLOTYPE: LMJG, fragment: BMNH1910.1.1.542. E. Mediterranean.
Clathria (Clathria) gorgonoides (Dendy, 1916)
Echinodict yum gorgonoides Dendy, 1916a: 129 [Kattiawar,
Indian Ocean]; Hooper, 1991: 1348.
MATERIAL. HOLOTYPE: BMNH1920.12.9.38. W. India.
Clathria (Clathria) hexagonopora Levi, 1963
Clat hria hexagonopora Levi, 1963: 53-54, text-fig.60, p1.8D
[Cape of Good Hope].
MATERIAL. HOLOTYPE: MNHNDCL620. S. Africa.
Clathria (Clathria) indica Dendy, 1889
Clat hria indica Dendy, 1889b: 73, 84, p1.4, fig. 10 [Gulf of
Manaar, Ceylon]; Burton 8c Rao, 1932: 336-337, p1.18,
figs 8-9 [Tuticorin, India]; Thomas, 1979a: 58, p1.3, fig. 10
[Mozambique].
Wilsonella indica; Dendy, 1905: 171 [Ceylon]; Hallmann,
1912: 242; Burton, 1931a: 346 [Natal Coast].
MATERIAL. HOLOTYPE: BMNH1887.8.4.19, fragment:
MNHNDCL2528. E. Africa, SE. India, Gulf of Manaar.
Clathria (Clathria) inhacensis Thomas, 1979
Clat hria inhacensis Thomas, 1979: 27-28, p1.2, fig.2 [Inhaca
I., Mozambique].
MATERIAL. HOLOTYPE: MRAC511. E. Africa.
Clathria (Clathria) intermedia Kirk, 1911
Clat hria int erm edia Kirk, 1911: 579, text-fig.5 [Kermadec
Is]; Fell, 1950: 11, text-fig.2; Berquist & Fromont, 1988:
109-110; Dawson, 1993:37 [note].
Thalysias int erm edia; de Laubenfels, 1936a: 105.
MATERIAL. HOLOTYPE: NMNZ unregistered. N. New
Zealand.
Clathria (Clathria) irregularis (Burton, 1931)
Mar leyia ir r egular is Burton, 1931a: 346, p1.23, fig.6, text-
fig.5 [Durban, Natal]; de Laubenfels, 1936a: 109 [note];
Van Soest, 1984b: 130 [note].
MATERIAL. HOLOTYPE: NM1279; paratype: BMNH1934.10.1.17. S. Africa.
Clathria (Clathria) juncea Burton, 1931
Clat hria j uncea Burton, 1931a: 343, p1.23, fig.5, text-fig.3
[Natal coast]; Levi, 1960a: 85; Levi, 1963: 66 [note].
Labacea j uncea; de Laubenfels, 1936a: 66 [note].
cf. Alcyonium j unceum Lamarck, 1816: 77.
cf. Anom oclat hria opunt ioides var; Topsent, 1933: 26 [note].
Reniera labyrint hica Schmidt, 1864:39, p1.4, fig.9 [Adriatic].
Clat hr ia labyr int hica; Burton, 1930a: 514 [imperfectly
known].
MATERIAL. HOLOTYPE: LMJG. Mediterranean.
Clathria (Clathria) laevigata Lambe, 1893
Clat hria laevigat a Lambe, 1893:31-32, p1.2, figs 9-9f [Pacific
coast, Canada]; Lambe, 1900: 160; Koltun, 1959: 185,
text-fig. 146 [Kurile Is].
Thalysias laevigat a; de Laubenfels, 1936a: 105.
cf. Microciona prolifera, tropus sent a; Vosmaer, 1935a: 641.
MATERIAL. HOLOTYPE: NMCIC1900-2892. NW. and
NE. Pacific.
Clathria (Clathria) lissosclera Bergquist &
Fromont, 1988
Clat hria lissosclera Bergquist & Fromont, 1988: 106-107,
p1.49, figs d-f [Mayor I.]; Dawson, 1993: 37 [note].
MATERIAL. HOLOTYPE: NMNZPOR112. New Zealand.
Clathria (Clathria) lobata Vosmaer, 1880
Clat hria lobat a Vosmaer, 1880: 151 [Cape of Good Hope];
Vosmaer, 1882: 45; Carter & Hope, 1889: 101, 105;
Stephens, 1915: 444-445 [Cape of Good Hope]; Levi,
1963: 54, text-fig.61, p1.8F [Cape of Good Hope].
Rhaphidophlus lobat us, var. horrida; Ridley & Dendy, 1887:
153-155, p1.28, fig.1, p1.29, fig.4a-c [Simon's Bay, Cape
of Good Hope].
Ligrot a lobat a; de Laubenfels, 1936a: 125.
A rt em isina lobat a; Van Soest, 1984b: 130 [generic synonymy
for Ligrot a] .
cf. Microciona prolifera; Vosmaer, 1935a: 631-632.
MATERIAL. HOLOTYPE: unknown, fragments
RMNH276-277, BMNH1887.5.2.100. S Africa.
Clathria (Clathria) macroisochela Levi, 1994
Clat hria ( Clat hria) m acroisochela Levi, 1994: 37, fig.21b,
p1.7, figs 7-9 [Norfolk Rise, 680m depth].
MATERIAL. HOLOTYPE: MNHNDCL3638. SW Pacific.
Clathria (Clathria) maeandrina Ridley, 1884
Clat hria m aeandrina Ridley, 1884a: 588, 614, p1.53, fig.1,
p1.54, fig.H [Amirante Is]; Burton, 1930c: 668 [Gulf of
Manaar]; Burton, 1959: 244 [Red Sea].
MATERIAL. HOLOTYPE: BMNH1882.10.17.55. Red Sea, W
Indian Ocean, Gulf of Manaar.
Clathria (Clathria) marissuperi PulitzerFinali, 1983
Clat hria m arissuperi Pulitzer-Finali, 1983: 569-571, text-
fig.67 [Italy].
MATERIAL. HOLOTYPE: IZUG(MSNG 47179). Mediterranean.
Clathria (Clathria) menoui Hooper & Levi,
1993
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
175
Clathria (Clathria) menoui Hooper & Levi, 1993a: 12701273, figs 25-26 [New Caledonia].
MATERIAL. HOLOTYPE: QMG301267. SW. Pacific.
Clathria (Clathria) meyeri (Bowerbank, 1877)
Ophlitaspongia meyeri Bowerbank, 1877: 456 [Selat Japen,
Irian Jaya, E. Indonesia].
Clathria meyeri; Vosmaer, 1880: 154.
? Anaata meyeri; de Laubenfels, 1936a: 109.
MATERIAL. HOLOTYPE: Dresden Musuem destroyed
(fragments BMNH1877.5.21.1306-1309). Indonesia.
Clathria (Clathria) microchela (Stephens,
1916)
Eurypon microchela Stephens, 1916: 240-241 [SW coast,
Ireland]; Lilly et al., 1953 [Lough me, Ireland]; Burton,
1959b: 44 [Iceland]; Van Soest & Weinberg, 1980: 10
[Lough Inc. Ireland].
Dictyociona microchela; de Laubenfels, 1936a: 110 [note]; de
Laubenfels, 1953: 528.
Clathria microchela; Van Soest, 1984b: 7 [generic
synonymy].
Not Microciona microchela Hechtel, 1965: 41.
MATERIAL. HOLOTYPE: INMSR353.10.1916. NE. Atlantic.
Clathria (Clathria) microxa Desqueyroux,
1972
Clathria microxa Desqueyroux, 1972: 27-28, text-figs 7680,134 [Gulf of Corcovado, Chile].
MATERIAL. HOLOTYPE: ICBUC. SE Pacific.
Clathria (Clathria) mortensenii Brondsted,
1923
Clathria mortensenii Brondsted, 1923: 143-144, text-fig.22
[Campbell Is]; Burton, 1940: 111 [Argentina]; Bergquist
& Fromont, 1988: 107-108, p1.50, figs a-c [N. New
Zealand]; Dawson, 1993: 37 [note].
Microciona mortensenii; de Laubenfels,I936a: 111; Burton,
1940: 111 [Argentina].
Microcionaheterospiculata; Bergquist, 1961a: 39 [N of New
Zealand].
Not Microciona heterospiculata Brondsted, 1924: 465, textfig.20.
MATERIAL. HOLOTYPE: UZM, fragment:
BMNH1930.8.11.10. SW. Atlantic, Subantarctic, New
Zealand.
Clathria (Clathria) mosulpia Sim & Bakus,
1989
Clathria mosulpia Sim & Byeon, 1989: 38-39, p1.3, figs 1-5.
[South Korea].
MATERIAL. HOLOTYPE: HNUKP0r9. S. China Sea.
Clathria (Clathria) obliqua (George & Wilson,
1919)
Esperiopsis obliqua George & Wilson, 1919: 148-150, p1.60,
figs 20-23, p1.66, fig.58 [North Carolina].
Microciona obliqua; de Laubenfels, 1936a: 111.
cf. Microciona prolifera; de Laubenfels, 1947; de Laubenfels,
1949a.
Haliclona oculata; Pearse & Williams, 1951 [North
Carolina].
Chalina arbuscula; Coues & Yarrow, 1879: 312; Verrill &
Smith, 1873: 743 [North Carolina].
Tenaciella obliqua; Wells et al., 1960: 218-219, text-figs
16,25 [North Carolina].
Not Tenaciella obliqua; Alcolado, 1976: 5.
Clathriaobliqua; Van Soest, 1984b: 104, 108, table 4 [affinity
with Clathria proliferat
Not Dictyociona adioristica de Laubenfels, 1953a: 526; Van
Soest, I984b: 108 [possible synonym].
MATERIAL. HOLOTYPE: USNM23612, paratypeUSNM23613. NW. Atlantic-Caribbean.
Clathria (Clathria) oculata Burton, 1933
Clathria oculata Burton, 1933: 250-251, text-fig.2 [Natal];
Levi, 1963: 67 [Natal].
Thalysias oculata; de Laubenfels, 1936a: 105.
MATERIAL. HOLOTYPE: BMNH1933.7.4.65. S. Africa.
Clathria (Clathria) oxneri (Topsent, 1928)
Hymedesmia oxneri Topsent, 1928a: 256 [Azores].
Dictyociona oxneri; de Laubenfels, 1936a: 110 [note]; de
Laubenfels, 1953a: 528.
Clathria oxneri; Van Soest, 1984b: 7 [generic synonymy for
Dictyociona].
MATERIAL. HOLOTYPE: MOM (fragment MNHNDT963). NE. Atlantic.
Clathria (Clathria) pachystyla Levi, 1963
Clathria pachystyla Levi, 1963: 56, text-fig.63, p1.8G [Cape
of Good Hope].
MATERIAL. HOLOTYPE: MNHNDCL609. S. Africa.
Clathria (Clathria) papillosa Thiele, 1905
Clathria papillosa Thiele, 1905: 449-450, text-fig.66 [Calbuco, Chile]; Burton, 1932a: 319 [Falkland Is];
Desqueyroux-Faundez & Moyana, 1987: 49 [Chile, Argentina, Falkland Is, Kerguelen Is].
Pseudanchinoepapillosa; de Laubenfels, 1936a: 109; Burton,
1940: 115, p1.5, figs 1-7 [Argentina].
MATERIAL. HOLOTYPE: ZMB not located, fragment:
BMNH1908.9.24.162. SE. Pacific, SW. Atlantic.
Clathria (Clathria) pellicula Whitelegge, 1897
Clathria pellicula Whitelegge, 1897: 327-328 [Funafuti (Ellice Is)].
Hymeraphia pellicula; Hallmann, 1912: 208 209.
cf. Microciona prolifera tropus tegens; Vosmaer, 1935a: 641-2.
-
MATERIAL. HOLOTYPE: AMG1660. SW central Pacific.
Clathria (Clathria) plurityla Pulitzer-Finali,
1983
Clathria plurityla Pulitzer-Finali, 1983: 571-572, 610, textfig.68 [Italy].
MATERIAL. HOLOTYPE: IZUG(MSNG47180). Mediterranean.
Clathria (Clathria) productitoxa (Hoshino,
1981)
Thalysias productitoxa Hoshino, 1981: 157-159, text-fig.69,
p1.7, fig.3 [Uchinoura].
MATERIAL. HOLOTYPE: MMBSSAT-018. Japan.
Clathria (Clathria) prolifera (Ellis & Solander,
1786)
Spongia prolifera Ellis & Solander, 1786: 189-190, p1.58,
fig.5; Linnaeus, 1791: 3822; Esper, 1794: 178,281; Bose,
1802:143; Lamarck, 1814: 372; Lamouroux, 1816: 81-82;
MEMOIRS OF THE QUEENSLAND MUSEUM
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Montagu, 1818: 86; Blainville, 1819: 106; Lamouroux,
1821: 31, 109, p1.58, fig.5; Lamouroux, 1824: 368;
Templeton, 1836: 472; Johnston, 1842: 170-171; Gray,
1848: 19, 23.
Not Spongia prolifera; Grant, 1826: 115-116, 123, 135, 138,
347.
Microciona prolifera; Verrill, 1873: 741-742, pls 1-5 [USA];
Verrill, 1880: 232; Coues & Yarrow, 1879: 312 [North
Carolina]; Hyatt, 1885: 131; Norman, 1892: 14; Wilson,
1900: 350 [Beaufort, N.Carolina]; Wilson, 1902:396-397;
Wilson, 1907: 246; Wilson, 1910: 1269; Wilson, 1911:
3-11, 14, 29-30, p1.1, figs 1-6, p1.2, figs 7-12, p1.3, figs
13-20, p1.4, figs 21-25, p1.5, figs 26-32; George & Wilson,
1919: 157-158, p1.62, figs 31, 33, p1.63, figs 35-36, p1.66,
fig.57 [North Carolina]; Allee, 1923: 175; Galtsoff &
Pertzoff, 1926: 239-254 [physiology]; Burton & Rao,
1932: 344 [Arabian Sea; probable misidentification]; Proctor, 1933: 104; de Laubenfels, 1936a: 111; McDougall,
1943: 331-332; Bergmann, Schedl & Low, 1945: 580; de
Laubenfels, 1947:35; Hopkins, 1956a: 44; Hopkins, 1962:
124; Hartmann, 1958a: 36-41, text-fig.10, table 11 [New
England, USA]; Levi, 1960a: 52; Wells eta!., 1960: 213216, text-figs 18,22 [North Carolina]; Little, 1963: 49
[Honda]; Wells et al., 1964: 757-758 [North Carolina];
Bagby, 1966: 167-181, pls 3-5 [cytology]; Simpson,
1968a: 18, pls 1-5 [Connecticut, USA]; Simpson, 1968b:
252-277 [reproduction]; Wendt, 1970: 3500-B [cytology];
Sindelar, 1970: 3771-B [cytology]; Bagby, 1970: 579-594
[ultrastructure]; Kunen et al., 1970:565-576 [physiology];
Madri et al., 1970: 1-5 [biochemistry]; Swartz, 1972: 17
[ecology]; Bagby, 1972: 217-244 [ultrastructure]; Bito,
1972: 65 [biochemistry]; Bose et al., 1972: 217-222
[biochemistry]; Turner & Burger, 1973: 509-510, textfig.! [cytology]; Weinbaum & Burger, 1973: 510-512,
[biochemistry]; Stempien, 1973: 363; Sutherland, 1974:
859-873 [ecology]; Turner et al., 1974: 35 [cytology];
Bose, 1974: 476-490 [chemistry]; Jefferts et al., 1974:
244-247 [biochemistry]; Reiswig, 1975:493-502 [physiology]; Reed et al., 1976: 153-169 [cytology]; Jumblatt et
al., 1976: 73-86 [cytology]; Morales & Litchfield, 1976:
206-216 [biochemistry]; Burger, 1977: 357-376 [cytology]; Greenberg et al., 1977: 95-102 [cytochemistry];
Morales, 1977: 5043 [biochemistry]; Morales &
Litchfield, 1977: 570-576 [biochemistry]; Simpson, 1978:
31-42 [morphology]; Burkart eta!., 1979: 239-246 [cytology]; Leith, 1979: 212-223 [cytology]; Litchfield &
Liaaen-Jensen, 1980: 359-365 [biochemistry]; Jumblatt et
al., 1980: 1038-1042 [biochemistry-cytology]; Saxegaard
eta!., 1981: 325-327 [biochemistry]; Lee & Nicol, 1981:
445 [chemistry]; Biernbaum, 1981: 85-96 [ecology];
Liaaen-Jensen et al., 1982: 170-171 [biochemistry];
Misevic & Burger, 1982: 200 [biochemistry-cytology];
Misevic eta!., 1982: 6931-6936 [biochemistry-cytology];
Kuhns et al., 1980: 73-79 [cell reaggregation]; Dunham et
al., 1983: 4756 [cell reaggregation]; Rice & Humphreys,
1983: 6394-9 [biochemistry]; Akiyama & Johnson, 1983:
687-694 [biochemistry]; Collier, 1983: 428-432
[biochemistry]; Lee & Gilchrist, 1985: 24-32
[biochemistry]; Dunham et al., 1985: 2914 [cytology];
Sharma et al., 1985: 241 [biochemistry]; Knight & Fell,
1987: 263 [physiology]; Misevic eta!., 1987: 5870 [cytology]; Stanley-Samuelson, 1987:92 [chemistry]; Sliwka et
al., 1987: 245 [chemistry]; Knight & Fell, 1987: 253
[cytology]; Misevic & Burger, 1988: 134-152; Fell, Knight
& Rieders, 1989: 195; Misevic & Burger, 1988: 134;
1990a: 307; 1990b: 20577 [cytology]; Kuhns, Misevic &
Burger, 1990: 358 [biochemistry]; Leamon & Fell, 1990:
265 [cytology]; Ayanoglu et al., 1990: 597; Lam et al.,
1991: 372 [biochemistry]; Misevic & Burger, 1990c: 81
[chemistry]; Fell, 1990: 497 [ecology]; Misevic et al.,
1990: 182 [ontogeny]; Spillmann et al., 1993: 13378
[chemistry]; Aho eta!., 1993: 7288 [genetics].
Not Microcionaprolifera; de Laubenfels, 1949a: 12, text-figs
8-10 (Pearse & Williams, 1951: 135); Johnson, 1971:
110-111, text-fig.14 (Van Soest, 19846: 93).
cf. Microciona prolifera, in part; Vosmaer, 1935a: 612-613
[excessive lumping of species].
Clathria (Clathria) prolifera; Van Soest, 19846: 91-93, textfig.35, table 4 [North Carolina].
Not Clathria prolifera Burton, 1940: 109 [see C. burtoni,
nom.nov.].
Spongia ostacina Rafinesque, 1819: 150.
Spongia urceolata Desor, 1851: 67.
Clathria delicata Lambe, 1896: 12, 160, 192, p1.2, fig.2
[St.Lawrence Gulf]; Lambe, 1900: 160; Whiteaves, 1901: 18
[Canada]; Hentschel, 1912: 365; Hentschel, 1929: 971;
Hartman, 1958: 37.
Thalysias delicata; de Laubenfels, 1936a: 105.
Esperiopsis obliqua, in part; de Laubenfels, 1947: 5.
MATERIAL. HOLOTYPE: BMNH missing; representative
specimens: USNM23562, ZMAPOR38, MCZH6907,
PMNH754, PMNH1912E, NMCIC1900-2874. NW. Atlantic.
Clathria (Clathria) pyramidalis (Brondsted,
1924)
MicrocionapyramidalisBrondsted, 1924:466, text-figs 21a-e
[Slipper Is]; Dawson, 1993: 37 [note].
Dictyociona pyramidalis; de Laubenfels, 1936a: 110 [note];
de Laubenfels, 1953a: 528; Bergquist & Fromont, 1988: 104.
MATERIAL. HOLOTYPE: UZM (not located). New
Zealand.
Clathria (Clathria) rectangulosa Schmidt,
1870
Clathria rectangulosa Schmidt, 1870: 60 [Tortugas, Florida];
Vosmaer, 1880: 149; Ridley & Duncan, 1881: 485; Wiedenmayer, 1977: 261, table 52 [imperfectly known];
Desqueyroux-Faundez & Stone, 1992: 36, 103.
cf. Clathria coralloides; Vosmaer, 1880: 149.
MATERIAL. HOLOTYPE: BMNH1870.5.3.68. NW. Atlantic - Caribbean.
Clathria (Clathria) rhaphidotoxa Stephens,
1915
Clathria rhaphidotoxa Stephens, 1915: 445-447, p1.38, fig.2,
p1.40, fig. 15 [Saldanha Bay]; Levi, 1963: 57-58, text-fig. 65,
p1.8H [St. Helena and Saldanha Bays].
Thalysias raphidotoxa; de Laubenfels, 1936a: 105.
MATERIAL. HOLOTYPE: RSME 1921.143.1451 (fragment
BMNH 1953.11.11.144). S Africa.
Clathria (Clathria) sarai sp.nov.
Clathria elastica Sara, 1978: 70-73,text-figs 44-46 [Cape San
Sebastiono, Tierra del Fuego]; Desqueyroux-Faundez &
Moyano, 1987: 50 [Tierra del Fuego, Argentina].
Not Clathria elastica Levi, 1963: 52.
MATERIAL. HOLOTYPE: IZUG116, fragment:
MNHNDCL604. Note: C. elastica Levi, 1963 has seniority.
SW Atlantic.
Clathria (Clathria) saraspinifera sp. nov.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWV
^
177
Clathria spinifera Sara, 1978: 67-70, text-figs 41-43 [Rio
Grande, Tierra del Fuego]; Desqueyroux-Faundez &
Moyano, 1987: 50 [Tierra del Fuego, Argentina].
Not Rhaphidophlus filifer var. spinifera Lindgren, 1897: 483.
Not Rhaphidophlus spinifer Thiele, 1903a: 958, p1.28, fig. 23;
Hallmann, 1912: 177.
MATERIAL. HOLOTYPE: IZUG147, fragment:
MNHNDCL2755. SW. Atlantic. Note: C. spinifera (Lindgren,
1897) has seniority.
Clathria (Clathria) sartaginula (Lamarck, 1814)
Spongia sartaginula Lamarck, 1814: 383,362 [locality unknown].
Pseudanchinoe sartaginula; de Laubenfels, 1936a: 109
[note].
Clathria sartaginula; Topsent, 1930:45, p1.4, fig. 1 .
MATERIAL. HOLOTYPE: MNHNDT527. Unknown.
Clathria (Clathria) shirahama Tanita, 1977
Clathria shirahama Tanita, 1977: 38, p1.2, fig.9, text-fig.6
[Kii-Shirahama]; Hoshino, 1981: 161.
MATERIAL. HOLOTYPE: MMBS. Japan.
Clathria (Clathria) spinispicula Tanita, 1968
Clathria spinispicula Tanita, 1968: 48-49, p1.1, fig.6, textfig.8 [Ariake Sea]; Rho et al., 1972: 5, p1.4, figs 9-10 [South
Korea]; Hoshino, 1981: 161 [Ariake Sea]; Sim & Bakus,
1988: 25 [Korea]; Sim & Byeon, 1989: 38 [Korea].
MATERIAL. HOLOTYPE: MMBS. Japan, S. China Sea.
Clathria (Clathria) spongodes Dendy, 1922
Clathria spongodesDendy, 1922: 69, p1.6, fig.1, p1.14, fig.2
[Amirante]; Vacelet et al., 1976: 70-71, text-fig. 48, p1.3, fig.e
[Madagascar].
Damoseni spongodes; de Laubenfels, 1936a: 110 [note].
Clathria madrepora Dendy, 1922: 68-69, p1.5, fig.3, p1.14,
fig.1 [Seychelles]; Sim & Kim, 1988: 25, p1.2, figs 1-2
[Korea]; Sim & Byeon, 1989: 38 [Korea; possible misidentification].
Thalysias madrepora; de Laubenfels, 1936a: 105.
Clathria spongiosa Burton, I959a: 245, text-fig. 26 [Red
Sea]; Vacelet et al., 1976: 70 [with question].
MATERIAL. HOLOTYPE: BMNH1921.11.7.58. W. Indian
Ocean, ? S. China Sea.
Clathria (Clathria) surculosa (Esper, 1797)
Spongia surculosa Esper, 1797: 39, p1.65 A3 ['East Indies'].
Clathria surculosa; Ehlers, 1870: 23,31.
MATERIAL. HOLOTYPE: Unknown. Indonesia.
Clathria (Clathria) terranovae Dendy, 1924
Clathria terrae-novae Dendy, 1924a: 353-354, p1.12, fig.5,
p1.14, figs 9-13 [North Cape, New Zealand]; Dawson,
1993: 37 [note].
Dictyociona terrae-novae; Burton, 1932a: 324 [Falkland Is];
Burton, 1940: 112-114, p1.5, figs 3-4 [Argentina]; de
Laubenfels, 1953a: 528; Koltun, 1964a: 72-73 [Antarctica].
Clathria terranova; Koltun, 1976: 188; Rho & Sim, 1976:74,
p1.6, figs 3-4 [Seogwipo, South Korea; possible misidentification]; Boury-Esnault & van Beveren, 1982: 107-108,
p1.18, fig.69, text-fig.31 [Kerguelen Is]; Bergquist &
Fromont, 1988: 109, p1.50, figs d-f, p1.51, fig.a [N. New
Zealand]; Sim & Byeon, 1989: 38 [Korea].
MATERIAL. HOLOTYPE: BMNH1923.10.1.132, paratypes
BMNH1923.10.1.133, 134. Antarctica, Subantactic, New
Zealand, SW Atlantic.
Clathria (Clathria) textile (Carter, 1876)
Cornulum textile Carter, 1876: 309.
Clathria textile; Vosmaer, 1880: 154 [Shetland Is; imperfectly
known].
MATERIAL. HOLOTYPE: BMNH1882.7.28.75. NE Atlantic.
Clathria (Clathria) tortuosa Uriz, 1988
Clathria tortuosa Uriz, 1988a: 86-87, pls 22a, 42a-b, textfig.62 [Namibia].
MATERIAL. HOLOTYPE: ABIP6B-11d. SW Africa.
Clathria (Clathria) toxistricta Topsent, 1925
Clathria toxistricta Topsent, 1925:656-658, text-fig. 13 [Gulf
of Naples]; Sara, 1960a: 462 [Ischia]; Levi, 1960a: 62-63
[Naples]; Boury-Esnault, 1971: 324 [Banyuls]; PulitzerFinali, 1983: 610 [list].
MATERIAL. HOLOTYPE: MOM, fragments-MNHNDT327,
1244. Mediterranean.
Clathria (Clathria) toxistyla (Sara, 1959)
Microciona toxistyla Sara, 1959: 17, text-fig.6 [Naples];
Siribelli, 1960:9-10, text-fig. 3B [Naples]; Sara, 1961:47
[Adriatic Sea]; Cimino et al., 1979: 3619-3622
[biochemistry].
Clathria toxistyla; Melone, 1963: 5-7, p1.1, fig.2, text-fig.2
[Adriatic Sea; erect form of Microciona toxistyla]; Sara,
1964: 229 [Ligurian Sea]; Sara & Melone, 1963: 20
[Adriatic Sea]; Labate, 1964: 334 [Adriatic Sea]; PulitzerFinali, 1983: 569, 610, fig.66 [Mediterranean].
MATERIAL. HOLOTYPE: ZSN1000. Mediterranean.
Clathria (Clathria) toxivaria (Sara, 1959)
Microciona toxivaria Sara, 1959: 14, p1.1C, text-fig.5
[Naples]; Siribelli, 1960: 8, text-fig.3A [Naples].
Clathria toxivaria; Melone, 1963: 2-3, p1.1, fig.1, text-tig.1
[Adriatic; erect forms of Microciona toxivaria]; PulitzerFinali, 1977: 61 [Bay of Naples]; Sara, 1964: 229-230
[Ligurian Sea]; Sara & Melone, 1963: 20-21 [Adriatic];
Labate, 1964: 334 [Adriatic]; Boury-Esnault, 1971: 323
[Banyuls]; Pulitzer-Finali, 1983: 568, 610 [Mediterranean].
MATERIAL. HOLOTYPE: ZSNGG920, paratype ZSNGG923. Mediterranean.
Clathria (Clathria) toxotes (Schmidt, 1868)
Scopalina toxotes Schmidt, 1868: 12, 26, 39, 40, 44, p1.5, fi g.5
[Canal of Zara, Adriatic]; Schmidt, 1870: 2, 56;
Pagenstecher, 1872:43; Hyatt, 1877: 500; Schmidt, 1880:
81; Vosmaer, 1880: 118-119; Fristedt, 1885:37; Vosmaer,
1885b: 353; Carter & Hope, 1889: 101; Schulze & Lendenfeld, 1889: 9; Heider, 1895: 281; Kieschnick, 1896:
533; Thiele, 1903a: 959; Svarcevskij, 1906: 342; Levi,
1960a: 55.
? Microciona toxotes; de Laubenfels, 1936a: 111 [imperfectly
known].
cf. Microciona armata; Vosmaer, 1935a: 627.
MATERIAL. HOLOTYPE: unknown, possibly LMJG.
Mediterranean.
MEMOIRS OF THE QUEENSLAND MUSEUM
178^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathria (Clathria) typica Kirkpatrick, 1904
Clathria typica Kirkpatrick, 1904: 148 [Natal]; Hallmann,
1912: 208 [anomalous species].
? Tenacia clathrata; Carter, 1875: 195 [nomen nudum];
Carter, 1878: 160,163; sensuV osmaer, 1935a: 628.
Not Echinonema typicum Carter, 1881a: 362.
MATERIAL. HOLOTYPE: BMNH1902.11.16.31. Note: imperfectly known; listed in BMNH register as C. typica Carter. S
Africa.
Clathria (Clathria) ulmus Vosmaer, 1880
Clathria ulmus Vosmaer, 1880: 151 [locality unknown]; Ridley, 1884a: 444.
Thalysias ulmus; de Laubenfels, 1936a: 105 [note].
MATERIAL. HOLOTYPE: possibly RNHL. Unknown
locality.
Clathria (Clathria) unica Cuartas, 1993
Clathria unica Cuartas, 1993: 112 [Argentina].
MATERIAL. HOLOTYPE: Division Invertebrados del Museo
de Ciencias Naturales de La Plata, Argentina. SW Atlantic.
Clathria (Clathria) vasiformis (de Laubenfels,
1953)
Thalyseutypon vasiformis de Laubenfels, 1953a: 525, textfig.4 [Gulf of Mexico]; Little, 1963:50 [note].
Clathria vasiformis, Van Soest, 1984b: 109, table 4.
Myxilla beani; Topsent, 1892b: 23; Topsent, 1894a: 8, 9, 25;
Hanitsch, 1894: 179.
cf. Microciona prolifera; Vosmaer, 1933: 610.
MATERIAL. HOLOTYPE: BMNH1930.7.3.339. Note:
Synonym of Antho involvens (Levi, 1960a: 76).
Echinonema caespitosa Carter, 1885f: 352 [Port Phillip,
Victoria].
Plumohalichondria caespitosa; Dendy, 1896: 41.
MATERIAL. HOLOTYPE: BMNH1886.12.15.453. Note:
referred to Anchinoidae, Plumohalichondria.
Antherochalina concentrica Lendenfeld, 1887b: 788, p1.22,
fig.42 [Port Molle (Airlee Beach), Q1d.].
Cymbastela concentrica; Hooper & Bergquist, 1992: 114.
MATERIAL. HOLOTYPE: AMZ1993 (lectotype), fragment:
BMNH1886.8.27.451, 460 (paralectotypes). Note: referred to
Axinellidae, Cymbastela.
Clathria corallorhizoides Fristedt, 1887: 460, p1.25, figs 7377, p1.29, fig.23.
MATERIAL. HOLOTYPE: unknown, fragment:
BMNH1910.1.1.1445. Note: referred to Myxillidae, synonym
of Lissodendoryx complicata (Lundbeck, 1905: 166).
Antherochalina elegans Lendenfeld, 1887b: 787, p1.22,
ML4-232. E Pacific.
fig.40.
Syringella elegans; Burton, 1934a: 558.
Raspailia (Syringe/la) elegans; Hooper, 1991: 1262.
MATERIAL. HOLOTYPE: BMNHI886.8.27.452. Note:
referred to Raspailiidae, Raspailia.
Clathria (Clathria) zoanthifera Levi, 1963
Echinonema incrustans Carter, 1885f: 353 [Port Phillip,
.
Pandaros vasiformis; Wiedenmayer, 1977: 144 [note].
MATERIAL. HOLOTYPE: USNM23403, paratype MLUM-
Clathria zoanthifera Levi, 1963: 58, text-fig.66, p1.9D [Cape
of Good Hope].
MATERIAL. HOLOTYPE: UCT (fragment IVINHNDCL607).
S Africa.
TRANSFERS
List of other species described in Clathria
(Clathria), or one of its synonyms, but now transferred to another genus.
ClathrM australis Lendenfeld, 1888: 222 [Port Phillip, Victoria].
Wilsonella australis; Hallmann, 1912: 239.
Not Clathria australis Whitelegge, 1901: 90.
Not Plumohalichondria australis; Whitelegge, 1901: 90,
p1.11, fig.14.
MATERIAL. LECTOTYPE: AMZ957. Note: referred Crellithe, synonym of Crella incrustans var. arenacea (Hallmann,
1914c: 411).
Plumohalichondria australis Whitelegge, 1901: 90-92,
p1.11, fig.14a,b [New South Wales coast]; Whitelegge,
1907: 492 [New South Wales coast].
MATERIAL. HOLOTYPE: AMG9042. Note: referred Crellidae, synonym of Crella incrustans (Hallmann, 1912: 160).
Isodictya beanii Bowerbank, 1866: 274, 334, 335 [Britain];
Gray, 1868: 164; Schmidt, 1870: 77; Bowerbank, 1874:
147, p1.58, figs 1-6.
Amphilectus beanii; Vosmaer, 1880: 115.
Clathria beanii; Ridley, 1881: 485, 486; Bowerbank, 1882:
13, 23, 150; Topsent, 1890c: 203.
Victoria].
Plumohalichondria incrustans; Dendy, 1896: 42.
Plumohalichondria mammillata; Carter, 1885f: 355; Ridley
& Dendy, 1887: 156, p1.30, fig.4, p1.47, fig.4.
Crella incrustans-, Hallmann, 1912: 152-156, p1.23, figs 2-3,
p1.24, text-figs 28-34.
MATERIAL. HOLOTYPE: BMNH1886.12.15.123, paratypes
1886.12.15.249, AME651, AMZ957. Note: referred to CrelCrella.
Cornulum johnsoni de Laubenfels, 1934: 15.
Clathria johnsoni; Van Soest, 1984b: 104 [possible synonym
of Clathria].
MATERIAL. HOLOTYPE: USNM Note: referred to Coelosphaeridae, possibly Cornulum (imperfectly known).
Echinonema levis Lendenfeld, 1888:220 [Port Jackson, New
South Wales].
Plumohalichondria australis, in part; Whitelegge, 1901: 65,
91,92; Whitelegge, 1902a: 212.
Crella incrustans var. levis; Hallmann, 1912: 164-167, textfigs 33-34.
MATERIAL. LECTOTYPE: AMZ959, paralectotype
AMG9708. Note: referred to Crellidae, synonym of Crella
incrustans (Hallmann, 1912: 164).
Clathria loveni Fristedt, 1887: 458, p1.25, figs 70-72, p1.30,
fig.24 [Cape Jakan, Siberian Arctic Ocean]; Lambe, 1900:
160.
[?] Esperella loveni; Lambe, 1895: 123, p1.4, fig.1 (W coast,
North America).
Esperia loveni; de Laubenfels, 1936a: 120 [note].
MATERIAL. HOLOTYPE: unknown, fragments
BMNH1927.2.22.2. Note: referred to Mycalidae, Mycale.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
179zyxwvutsrqponm
Plumohalichondria microcionides Carter, 1876: 236, p1.12,
fig.!!, p1.15, fig.30 [between Scotland and Faroe Is].
Clathria microcionides; Vosmaer, 1880: 154.
Plocamionida microcionides; Alander, 1942: 53 [Sweden].
Not Hymeraphia microcionides Carter, 1876: 390.
Not Plocamia microcionides; Topsent, 1891a: 529, 544-545.
MATERIAL. HOLOTYPE: BMNH1954.3.9.173. Note:
referred to Anchinoidae, Plocamionida.
Clathria mollis Kirkpatrick, 1903: 249-250, p1.5, fig. 15, p1.6,
fig. 16 [East London Coast, Natal].
Pronax mollis; Levi, 1963: 66.
Not Wilsonella mollis; Hallmann, 1912: 243.
Not Clathria dura var. molls; Hentschel, 1911: 370.
MATERIAL. HOLOTYPE: BMNH1902.11.16.32. Note:
referred to Anchinoidae, Pronax.
MATERIAL. HOLOTYPE: LMJG 15517; fragment BMNI-1
1867.3.11.29. Note: referred to Dictyoriellidae, Dictyonella.
Clathriaprocumbens Brondsted, 1923; Brondsted, 1926:329
[probably a typographical error for Clathrina (Calcarea),
and possibly synonym of Ascetta procumbens Lendenfeld,
1885a].
Antherochalina quercifolia Keller, 1889: 383-384, p1.23,
fig.34 [Red Sea].
Querciclona quercifolia; de Laubenfels, 1936a: 46 [note].
MATERIAL. HOLOTYPE: ZMB429, fragment
BMNH1908.9.24.179. Note: referred to Axinellidae, Phakellia.
Clathria raphida Hechtel, 1976:244; Van Soest, 1984b: 153.
Note: cited in a list of Brazilian endemic sponges; attributed to Boury-Esnault (1973); probably a misprint for
Cliona rhaphida Boury-Esnault, 1973.
Clathria morisca Schmidt, 1868: 9, 41, 43, p1.2, fig.7 [Algiers, Mediterranean]; Vosmaer, 1880: 150-151 [Algiers];
Antherochalina renieroides Lendenfeld, 1887b: 788, p1.28,
Topsent, 1902: 329; Topsent, 1938: 11; Desqueyrouxfigs 18, 23 [New Zealand].
Faundez & Stone, 1992: 10, 35.
MATERIAL. HOLOTYPE: BMN1-11886.8.27.449. Referred to
Diciyoclathria morisca; Topsent, 1920b: 18-21 [re-examinaAxinellidae, synonym of Phakellia flabellata (Carter) (see
tion of holotype]; Topsent, 1925: 660-661, p1.8, fig.! [Gulf of
Hooper, 1991).
Naples]; Topsent, 1928a: 301-302, p1.3, fig.3 [Porto Santo,
Azores]; Levi, 1959: 134, text-fig.27, p1.5, fig.! [Rio de Oro,
Hymeraphia tuberosocapitata Topsent, 1890b: 68 [Azores,
Gulf of Guinea]; Levi, 1960b: 761-762, text-fig.15 [var.
Atlantic]; Topsent, 1892a:113, p1.11, fig.6 [Atlantic].
anisotyla; SW. Cape of Naze, W. Africa]; Sara, 1960a: 462
Cionanchora tuberosocapitata; de Laubenfels, 1936a: 108
[Ischia, Mediterranean].
[note].
Artemisina mediterranea Babic, 1921: 87 [Adriatic]; Babic,
Clathria tuberosocapitata; Van Soest, 1984b: 7 [generic
synonymy for Cionanchora].
1922: 258, text-fig.3; Burton, 1930a: 528.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Myxilla banyulensis, in part, Topsent, 1892b: 23; Topsent,
Discorhabdella tuberosocapitata; Boury-Esnault, Pansini &
1902: 351, 363, 366; Cotte, 1903:423.
Uriz, 1992: 2-6.
cf. Clathria coralloides and C. compressa; Vosmaer, 1935a:
MATERIAL. HOLOTYPE: MOM040323, fragment
626 [intermediate form].
MNHNDT938. referred to Hymedesmiidae, Discorhabdella.
MATERIAL. HOLOTYPE: MNHN DT2170, fragment
BMNH 1868.3.2.21. Note: Synonym of Antho involvens (Levi,
Clathria (Wilsonella) Carter, 1885
1960a: 57).
Hooper & Wiedenmayer, 1994
Clathriaoroides Schmidt, 1864:35, p1.4, figs 1-2 [Quarnerno,
Adriatic]; Carter, 1875: 195; Vosmaer, 1880: 155; Dendy,
1889a: 41; Desqueyroux-Faundez & Stone, 1992: 10, 35,
103.
? Oroidea adriatica; Gray, 1867: 520.
Chalinopsis oroides; Schmidt, 1870: 60.
Ophlitaspongia oroides; Bowerbank, 1874: 10.
Age/as oroides; Riitzler, 1965: 34 [Banyuls]; Boury-Esnault,
1971: 322 [Banyuls]; Laubier, 1966 [Banyuls]; PulitzerFinali, 1983: 534 [Mediterranean].
MATERIAL. HOLOTYPE: LMJG 15957, fragment
BMNH1868.3.2.22. Note: referred to Agelasidae, Agelas.
Artemisina paradoxa Babic, 1921:87; Babic, 1922:258-261,
p1.8, fig.6, text-fig.c [Adriatic]; Levi, 1960a: 85-86
[Adriatic; with question]
Clathria paradoxa; Burton, 1930a: 528.
Dictyoclathria morisca; Topsent, 1925: 660; Ristau, 1978:
585-586 [note on affinities].
MATERIAL. HOLOTYPE: unknown. Note: synonym ofAntho
involvens (Topsent, 1925: 660).
Clathria pelligera Schmidt, 1864: 34, p1.3, fig.13 [Lesina,
Adriatic]; Desqueyroux-Faundez & Stone, 1992: 10, 36,
103.
Rhaphidostyla pelligera; Burton, 1935b: 652; Sara, 1958:
246-247, fig.15 [Gulf of Naples].
Stylotella pelligera; Topsent, 1925: 638; Boury-Esnault,
1971: 328 [Banyuls].
Dictyonella pelligera; Pulitzer-Finali, 1983: 545.
Wilsonella Carter, 1885f: 366 (not Hallmann, 1912:242).
Clathriopsamma Lendenfeld, 1888: 227.
Aulenella Burton & Rao, 1932: 345.
TYPE SPECIES. Wilsonella australiensis Carter,
1885f: 366 (by monotypy).
DEFINITION. Sand grains and foreign spicules
partially or completely replacing coring spicules
inside fibres; coring spicules same or very similar
geometry to auxiliary spicules located outside
fibres; skeletal architecture reticulate.
REMARKS. Microcionids with sand and foreign
debris incorporated into fibres ('arenaceous
sponges') are a predominant feature of S
Australian waters. Of the 17 species in Wilsonella
or a synonym, 14 are valid and 6 are known to
live in this region including 2 new species.
This group of arenaceous microcionids is well
known as Clathriopsamma (Hooper, 1990a;
Hooper & Levi, 1993a). However, the inclusion
of C. (W) australiensis in this group means that
Wilsonella (1885) takes precedence over Clathriopsamma (1888).
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A
FIG. 81. Clathria (Wilsonella) abrolholensis sp.nov. (holotype NTMZ3218). A, Auxiliary style. B, Echinating
acanthostyles. C, Palmate isochelae. D, Wing-shaped toxas. E, Section through peripheral skeleton (hatches
foreign spicules). F, Australian distribution. G, Preserved holotype. H, Holotype.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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181
Clathria (VVilsonella) abrolhosensis sp. nov.
(Figs 81-82, Plate 2C)
MATERIAL. HOLOTYPE: NTMZ3218 (fragment
QMG300584): N. edge of Pelsart Is, Houtman Abrolhos, WA, 28°47.2'S, 113°58.5'E, 10.vii.I987, 22m
depth, coll. J.N.A. Hooper (SCUBA).
HABITAT DISTRIBUTION. 22m depth; on an
Acropora reef; Houtman Abrolhos, WA (Fig. 81F).
DESCRIPTION. Shape. Fistulose, with multiple
fistules composed of erect, bulbous-cylindrical
digits, single or fused together with adjacent
digits, each tapering at base and apex, thickest in
apical portion, attached directly to substrate
without stalk; each digit between 75-140mm
long, up to 45mm maximum diameter.
Colour Pale beige-yellow alive (Munsell 2.5Y
8/4), dark brown in ethanol.
Oscules. Single, large, apical oscule on apex of
each digit, 10-15mm diameter in life, with slightly raised membraneous lip surrounding oscule;
oscules nearly completely contractile upon
preservation; pores not observed in life or
preserved.
Texture and surface characteristics. Compressible, spongy, relatively easily torn; surface
glabrous, even, without any ornamentation.
Ectosome and subectosome. Discrete, erect
bundles of auxiliary styles, relatively closely
packed together, arising from ends of peripheral
fibres, with tangential layer of auxiliary styles
connecting adjacent bundles; erect spicules
protrude only a short distance through surface;
mesohyl matrix light in ectosomal region; subectosomal region slightly cavernous, reticulate.
Choanosome. Almost regular, ovoid reticulation
of fibres and spicule tracts forming ovoid, square
or rectangular meshes, 220-360vm diameter;
spongin fibres moderately light, relatively
homogenous in size, 40-70Rm diameter, but with
differentiated primary and secondary tracts;
primary ascending fibres cored by multispicular
tracts of auxiliary styles, interconnected by
secondary, pauci- or multispicular transverse
tracts of auxiliary styles; fibres relatively heavily
echinated by acanthostyles; sparse core of
detritus in primary ascending fibres only, mostly
foreign spicules; mesohyl matrix moderately
heavy, lightly pigmented; choanocyte chambers
small, oval, 40-901im diameter, usually lined by
isochelae.
Megascleres. Principal spicules absent.
Auxiliary spicules coring fibres and forming
dermal skeleton moderately thin, straight or
slightly curved near base, with slightly subtylote
bases, hastate or slightly telescoped points, and
apical and basal spination on most spicules.
Length 1244151.7)-164,m, width 3-(4.1)-6p.m.
Echinating acanthostyles short, thin,
claviform, sharply pointed, slightly subtylote
bases, lightly and evenly spined, with relatively
large recurved spines. Length 71-(76.3)-86Rm,
width 3-(4.7)-7p,m.
Microscleres. Isochelae of 'typical' palmate
form, with straight shaft, lateral alae completely
fused to shaft, very long, broad front ala. Length
14-(15.7)-18Rm.
Toxas wing-shaped, slender, with pronounced
central curvature, slightly reflexed arms. Length
48488.4)-11211m, width 0.841.6)-2.0p.m.
ETYMOLOGY. For Houtman Abrolhos.
REMARKS. This species is a sibling species of
C. (Wilsonella) australiensis having basal and
apical spines on auxiliary styles. It was first assigned to the Western Australian subspecies C.
australiensis spinulata Hentschel (1911) but subsequent re-examination of Hentschel's (1911)
syntype (ZMB4446) found that he omitted
several crucial characters from his description
(and that his species was not substantially different from typical C. australiensis). Conversely,
C. (W) abrolhosensis differs from that species in
several respects, warranting its recognition as a
distinct taxon. Growth form in C. (W) abrolhosensis is endolithic, consisting of discrete bulbous-cylindrical digits arising from a partially
burrowing, encrusting massive base, and with
large terminal oscules (superficially resembling
syconoid calcarean growth forms) (whereas C.
(W) australiensis has lobate or clavulate morphologies); there is only one class of auxiliary
spicule (as compared with two); fibre reticulation
is small, close-set, nearly regular, and
predominantly ovoid (whereas in C. australiensis
meshes are elongate and cavernous); spongin
fibres are poorly developed (compared with well
developed fibres); there is very little detritus in
fibres, and these are completely confined to
primary ascending fibres and more-or-less
restricted to foreign spicules without sand (C.
(W) australiensis has abundant detritus in all
fibres, including sand grains, and this is a
prominent feature of the skeleton); and spicule
dimensions differ between both species. Further
comparisons are given below under remarks for
C. (W) australiensis.
This species strictly belongs to Clathria
(Dendrocia) in having only a single undifferentiated category of structural megasclere within
MEMOIRS OF THE QUEENSLAND MUSEUM
182zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
FIG. 82. Clathria (Wilsonella) abrolholensis sp.nov. (holotype NTMZ3218). A, Choanosomal skeleton. B, Fibre
characteristics. C-D, Spined base and point of auxiliary style. E, Echinating acanthostyles. F, Acanthosty le
spines. G, Wing-shaped toxas. H, Palmate isochelae.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
183
TABLE 15 Comparison between present and published records of Clathria (Wilsonella)australiensis Carter. All
measurements are given in rim, denoted as range (and mean) of spicule length x spicule width (N=25).
SPICULE
choanosomai
auxiliary
styles
Lectotype
Paralectotype
(BMNH1886. (BMNH1886.
12.15.43)
12.15.284)
92(119.8)152 x 2.5(3.6)4
105-(125.6)152 x 2.5(3.2)4
Holotype of
C. lobosa
(AMG9053)
Holotype of
T. ramsayi
(AMG8820)
Specimen
Specimen
Specimen
Paratype of
(N=1)
(Abrolhos Is)
(I-evi,1967)
var. spinulata
(NWAustralia)
(New
(NCIQ66
(ZMB4446)
C4266C)
Caledonia)
92(122.6)146 x 1.5(2.7)-4
106-(127.4)173 x 2.5(3.7)-5
112(118.8)135 x 2-(3.3)5
121-(149.0)163 x 2.5(3.7)4.5
89(111.4)175 x 2.5(4.1)-8
Subect^mal 92(122.6)148 x 2-(2.9)auxiliary
styles
4
101(118.3)155 x 1.5(2.9)4
94(119.6)98(117.2)141 x 2-(2.7)- 139 x 2-(2.8)3.5
4
108(120.1)134 x 2.5(3.7)-6
118(154.8)95(123.1)172 x 3-(4.1)- 164 x 2-(2.8)4.5
5
Echinatini
49-(59.8)-68
acanthosty es
x 2-(3.6)-4.5
45-(64.0)-77
x 2-(3.8)-4.5
54-(62.6)-78
x 2-(3.5)-4.5
46(58.4)-70
x 2.5-(3.4)4.5
46-(57.7)-66
x 4-(6.1)-9
59-(67.6)-74
x 3-(3.9)-6
11-(14.7)-18
12-(13.2)-16
12-(13.8)-16
12-(13.4)-15
12-(13.8)-16
72-(89.4)105 x 1-(1.6)2.5
0.8
Chelae
Toxas
35-(61 .2)-89
28-(53.4)-75
32-(52.3)-74
24-(45.4)-75
x 0.5-(0.7)-1
x 0.5-(0.8)-1
x 0.5-(0.8)-1
x 0.5-(0.8)-1
fibres, but it is included here in Clathria (Wilsonella) having detritus in fibres and spined terminations on auxiliary spicules similar to C. (W.)
australiensis. This further illustrates the unclear
generic boundary between many nominal genera
of Microcionidae, and supports the recognition of
a wide definition for Clathria.
Clathria (Wilsonella) australiensis (Carter,
1885) (Figs 83-84, Plate 2B, Table 15)
Wilsonella australiensis Carter, 1885f: 366; Hallmann,
1912: 239; Hallmann, 1920: 768; Burton, 1934a:
599.
Clathria australiensis; Dendy, 1896: 33; Whitelegge,
1901: 66, 84, 85, p1.11, fig.12; Hooper & Levi,
1993a: 1242, table 4; Hooper & Wiedenmayer,
1994: 275.
Clathria australiensis var. spinulata Hentschel, 1911:
374-375, text-fig. 47; Dendy, 1922: 71.
Clathriopsamma lobosa Lendenfeld, 1888: 149;
Whitelegge, 1901: 85; Hallmann, 1912: 239.
Thorecta ramsayii Lendenfeld, 1888: 149.
Sigmatella corticata var. elegans Lendenfeld, 1888:
199-201; Lendenfeld, 1889b: p1.40, fig.7.
Not Clathria australiensis; Levi, 1967b: 22, p1.2, fig.D,
text-fig.6.
Not Ophlitaspongia australiensis Ridley, 1884a: 442.
Not Echinochalina australiensis; Thiele, 1903a: 961962.
MATERIAL. LECTOTYPE: BMNH1886. 12.15.43
(fragment AMG2805): Port Phillip, Vic, 38 ° 09'S,
144 ° 52'E, 12m depth, coll. J.B. Wilson (dredge).
PARALECTOTYPE: BMNH1886.12.15.284: same
locality. HOLOTYPE of C. lobosa: AMG9053: Port
Jackson, NSW, 33 °51'S, 151 ° 16'E. HOLOTYPE of T.
ramsayi: AMG8820: same locality. SYNTYPES of
var. spinulata: ZMB4446, HM numbers unknown:
125-180 x4
51(60.4)-81
x 2.5-(3.7)5.5
65-80 x 4
14-(15.2)-17
12-(14.6)-19
14-15
28-(39.4)-59
x 0.5-(0.6)-
43-(55.6)-88
x 0.5-(0.8)1.5
40-75 x 1-2
Bunbury and Middleton Beach areas, WA, 33 ° 20'S,
115 °36'E, coll. W. Michaelsen & R. Hartmeyer
(dredge). OTHER MATERIAL: NSW - QMG301447,
QMG301458, AMG975, BMNH1886.12.15.288,
AMZI412, fragment NTMZ1526, AMZ3176,
AMZ3199, AMZ3140, AMZ4283. VICTORIA NMVRN748. S. AUST. - SAMTS4098 (fragment
NTMZ1646), SAMTS4116 (fragment NTMZ1627).
WESTERN AUSTRALIA - WAM623-81(1) (fragment NTMZ1710), QMG300622 (NCIQ66C-4266-C).
HABITAT DISTRIBUTION. Subtidal to 160m depth;
on rock, sand and algal bed substrates; known only
from Australia: throughout temperate Australian
waters - from Houtman Abrolhos, Perth, Bunbury
(WA) (Hentschel, 1911; present study), Nuyts Archipelago, St Vincent Gulf (SA) (present study), Port
Phillip (Vic) (Carter, 1885f; present study), Bega, Jervis Bay, Port Hacking, Port Jackson, Botany Bay, N.
Sydney, Tweed River region (NSW) (Lendenfeld,
1888 Whitelegge, 1901; present study), and extending
into the tropics as far as Low Isles, GBR, Qld. (Burton,
1934a) (Fig. 83G).
DESCRIPTION. Shape. Lobate, lobate-digitate,
club-shaped, thickly lamellate, or thickly encrusting-bulbous growth forms, up to 140mm high,
110mm wide, with subspherical, tubular, bulbous
or flabellate digits, up to 75n un high, 45mm wide,
25mm thick.
Colour. Pale orange alive (Munsell 7.5 YR 8/6),
pale pink, brown or yellow preserved.
Oscules. Large oscules, up to 4mm diameter, on
apical or lateral margins of surface digits/lobes.
Texture and surface characteristics. Surface optically even, microscopically rugose, with a
whitish arenaceous, slightly hispid dermal
membrane.
-
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184zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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FIG. 83. Clathria(Wilsonella)australiensis (Carter) (NTMZ1627). A, Subectosomal auxiliary subtylostyles. B,
Choanosomal auxiliary subtylostyles. C, Echinating acanthostyle. D, Oxhorn toxa. E, Palmate isochela. F,
Section through peripheral skeleton. G, Australian distribution. H, Holotype BMNH1886.12.15.43. I,
NMVRN748.
Ectosome and subectosome. Ectosome
membraneous, without specialised dermal
megasclere, varying from densely arenaceous,
with most or all dermal megascleres obscured by
large sand grains and spicule fragments, or with
light palisade of erect brushes of subectosomal
auxiliary subtylostyles, arising from ascending
primary choanosomal fibres; sand grains at surface slightly larger than those in choanosome.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
185
FIG. 84. Clathria (Wilsonella) australiensis (Carter) (A-B, AMG9053, QMG301447). A, Choanosomal
skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Echinating acanthostyle. E, Acanthostyle spines.
F, Palmate and modified isochelae. G, Oxhorn toxas. H-I, Continuum in basal and apical spination of auxiliary
subtylostyles.
MEMOIRS OF THE QUEENSLAND MUSEUM
186^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
differences in interpretation of character importance between the present study and that of
Hallmann (1920) (see also Hooper, 1990a).
Unlike Clathria (Dendrocia), in which there is
only a single category of coring and extra-fibre
megasclere, most Clathria (Wilsonella) have
more than one form of auxiliary style, one coring
the fibres (choanosomal megascleres) and one
outside of fibres (subectosomal megascleres). In
some cases (e.g., C. (W) australiensis, C. (W)
ensiae sp. nov.), these spicules are only slightly
different in geometry, although showing clear
differences in patterns of spination; in others
(e.g., C. (W) reticulata, C. (W) mixta) these
spicules have different geometry; whereas in one
(C. (W) abrolhosensis) there are no apparent
differences which is interpreted as a convergence
or subsequent loss of a spicule category. Clathria
(Dendrocia) and Clathria (Wilsonella) can also
be distinguished by their skeletal architecture being predominantly plumose in the former and
reticulate in the latter.
In material listed above most of the larger
auxiliary styles without spines on points appeared
to be located within spongin fibres (i.e.,
choanosomal spicules), whereas most of the
auxiliary styles with spines on both bases and
points were found predominantly outside fibres,
strewn within the mesohyl, and in the dermal
skeleton (i.e., subectosomal spicules). However,
this observation is difficult to verify in all cases
because of the dense core of sand particles in
fibres. In this species both sorts of spicules are
classed as auxiliary styles due to their very similar
geometry: true principal styles are absent (i.e.,
Wilsonella s.s.).
The principal diagnostic characteristics of C.
(W) australiensis and its affinities with other
species are discussed elsewhere (see Table 19 and
remarks for C. (W) tube rosa). This species differs
from most Australasian Clathria (Wilsonella) in
having spinose extremities on both the bases and
points of quasidiactinal auxiliary subtylostyles.
In this regard it is similar to its sibling species, C.
(W) abrolhosensis from the Houtman Abrolhos,
REMARKS. Clathria (W) australiensis is the WA (which is sympatric with C. australiensis)
only species retained by Hallmann (1920) in Wil- and the allopatric species C. (W) rugosa, from
sonella; other species were transferred to either New Caledonia (Hooper & Levi, 1993a; Table
Clathria or Paradoryx, depending on whether 19). Spicule geometry (megascleres and toxas)
they had palmate or arcuate-like isochelae are useful in distinguishing these species of
microscleres. Some of these, however, are further Clathria (Wilsonella) (Table 19).
Levi's (1967b) material from New Caledonia,
allocated here to Clathria (Dendrocia) or
Clathria (Thalysias), depending on their ec- described as C. australiensis, was referred to C.
tosomal skeletons, choanosomal skeletal struc- (W) rugosa (Hooper & Levi, 1993a), based on
ture and spicule diversity (i.e., there are differences in spicule geometry (especially shape
Choanosome. Choanosomal skeleton irregularly
reticulate, with clearly differentiated primary and
secondary spongin fibres, forming a vaguely longitudinal reticulation with cavernous meshes;
spongin fibres well developed; primary ascending fibres marginally thicker, producing ascending lines abundantly cored by detritus, lightly
cored by auxiliary styles, heavily echinated by
acanthostyles, particularly at fibre nodes; smaller
secondary spongin fibres mainly transverse, connecting with primary elements, with no or little
detritus, paucispicular tracts of auxiliary styles
and lightly echinated by acanthostyles; auxiliary
megascleres coring fibres occupy only a small
proportion of fibre diameter; detritus in fibres
consists of scattered sand grains and spicule fragments; mesohyl matrix heavy; choanocyte chambers oval to eliptical, some appear to be paired,
and chambers lined by microscleres.
Megascleres (Table 15). Choanosomal auxiliary
styles coring fibres differ from subectosomal
auxiliary styles only in being slightly thicker and
lacking characteristic apical microspines of the
latter. Coring spicules relatively thin, straight,
smooth, mostly hastate, with slightly subtylote
and occasionally microspined bases.
Subectosomal auxiliary styles, dispersed between fibres and in dermal skeleton, straight,
usually hastate, subtylote bases with microspines
on both points and bases.
Echinating acanthostyles small, evenly spinose
or with granular, vestigial spines, slightly subtylote bases, fusiform points.
Microscleres (Table 15). Palmate isochelae relatively large, unmodified.
Toxas oxhorn, uncommon, rare in some
specimens, often forming trichodragmata, usually with wide, angular, central curves and slightly
reflexed points.
Larvae. Ovoid, incubated parenchymella larvae,
up to 350p.m diameter seen in some material.
Associations. Commensal polychaetes common
in many specimens; Abrolhos specimen with
epizootic zoanthids on surface.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPO
^
187
and spination of auxiliary spicules), and the latter
having flabellate-digitate growth forms. The
quasidiactinal (strongylotes) modifications of
some of the subectosomal auxiliary spicules in
both species, with weakly spined points and
bases, is a unique trait within the Wilsonella
group, but is also known in a few other species of
Microcionidae (C. (Dendrocia) pyramida, C.
(Thalysias) major, C. (Clathria) chelifera). These
quasidiactinal spicules are convergent in
geometry with true tornotes and strongyles found
in other groups such as Iophonidae (see remarks
for Megaciella and Acarnus in the introductory
section above).
Hentschel (1911) created a subspecies
spinulata for material from WA, indicating that it
differed substantially from typical forms of the
species in spicule dimensions (particularly
isochelae), and supposedly lacked toxas. However, Hentschel's type has toxas and isochelae
dimensions are within the size range of other S
Australian populations, and all other aspects
(growth form, fibre characteristics, amount and
form of detritus in fibres) are identical between
populations. Hentschel (1911) suggested that
spinulata differed from other populations in
having spines on both the points and bases of
auxiliary spicules, but this feature occurs in all
other populations, and the WA population is conspecific with C. (W) australiensis.
Clathria (W.) australiensis is widely distributed
throughout temperate Australia, from the Houtman Abrolhos on the W coast (30 °S), around S
Australia to the Tweed River (28 °S). Burton
(1934a) recorded this species from Low Isles,
(16°S), but his voucher material has not been
examined, and his description is not detailed
enough to differentiate between C. (W.)
australiensis and C. (W) rugosa from New
Caledonia. Recent collections from this region
do not include the species. Examination of many
other specimens (cited above), from many locations throughout Australia, showed that C. (W)
australiensis is a heterogeneous species. Most
regional populations differ from each other
slightly in spicule geometry or spicule size, but
there is not enough available material of any of
these populations to determine whether these differences are consistent within populations, nor
are there any features in any of these regional
populations worthy of distinguishing separate
taxa (cf. Hentschel, 1911). Widespread sampling
of regional populations, collection of samples for
biochemical and genetic studies, and determination whether or not observed morhological dif-
ferences correlate with any fixed genetic differences is a worthwhile study for the future.
Clathria (Wilsonella) claviformis Hentschel,
1912
(Figs 85-86, Table 16, Plate 2D)
Clathria claveformis Hentschel, 1912: 366-367, p1.19,
fig.29.
MATERIAL. HOLOTYPE: SMF 1504 (fragment
MNHNDCL2238): Sungi Manumbai, near Kapala
Sungi, E. side of Aru I., Arafura Sea, Indonesia, 6°S,
134 ° 50'E, 28.iii.1908, 23m depth, coll. H. Merton
(dredge). OTHER MATERIAL: NT - NCIQ66C0528-0, NTMZ3082.
HABITAT DISTRIBUTION. 18-23m depth; sand,
rock and dead coral substrate; Timor Seas (Fig. 85G).
Also Arafura, SE. Indonesia.
DESCRIPTION. Shape. Claviform, 160-220mm
high, 60-150mm wide, with small cylindrical
base and stalk, and one or more lobate, clubshaped, or bulbous digits, up to 80mm long,
65mm diameter.
Colour. Pale 'dusty' beige and red-brown mottled
colour alive (Munsell 2.5Y 8/6 and 5R 8/4), yellow-grey preserved.
Oscules. Large oscules, up to 5min diameter, on
ends of bulbous digits, occasionally in crevices
between digits.
Texture and surface characteristics. Surface
even, with distinct, partially arenaceous, skinlike covering, and with several longitudinal
ridges on sides of digits and between bulbous
digits; texture rubbery, compressible, sandy.
Ectosome and subectosome. Ectosome lightly
and evenly arenaceous, hispid, with plumose tufts
of subectosomal auxiliary subtylostyles protruding through surface, sometimes in dense brushes;
subectosome cavernous, with sparsely cored,
arenaceous, ascending primary spongin fibres
forming large meshes in periphery.
Choanosome. Choanosomal skeleton irregularly
reticulate, cavernous, with light spongin fibres
and spicule tracts; spongin fibres divisible into
primary and secondary components; primary
fibres relatively large, with very little fibre spongin content, cored by both large and small sand
grains, Foraminifera and paucispicular tracts of
choanosomal principal styles, becoming sinuous
during towards periphery; secondary fibres consist of uni- or paucispicular spicule tracts bound
together by collagenous spongin and debris;
primary fibres moderately echinated, slightly
heavier towards periphery; secondary fibres with
188zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 85. Clathria (Wilsonella) claviformis Hentschel (holotype SMF1504). A, Choanosomal principal style. B,
Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Accolada toxas. E, Palmate isochelae. F,
Section through peripheral skeleton. G, Australian distribution. H, Holotype.
^
189
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
FIG. 86. Clathria (Wilsonella) claviformis Hentschel (holotype SMF1504). A, Choanosomal skeleton. B,
Peripheral skeleton. C, Fibre characteristics. D, Palmate isochelae. E, Echinating acanthostyle. F, Acanthostyle
spines. G, Base of principal subtylostyle. H, Bases of auxiliary subtylostyles. I, Accolada toxas.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
190zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
no or few echinating acanthostyles; mesohyl
matrix heavy, granular, with abundant
microscleres and detritus scattered throughout,
usually lining small oval choanocyte chambers.
Megascleres (Table 16). Choanosomal principal
styles thin, slightly curved, fusiform, with
rounded or slightly subtylote, lightly microspined
bases.
Subectosomal auxiliary subtylostyles hastate,
thin, mostly straight, with slightly subtylote
bases, usually microspined.
Echinating acanthostyles slightly subtylote,
with few, dispersed, large spines on base and
apex, usually with aspinose neck; spines located
on point usually more recurved than those on
base.
Microscleres (Table 16). Palmate isochelae large,
abundant, with some twisted examples.
Toxas thin, with slight angular central curvature and straight, unreflexed points.
REMARKS. C. (W) claviformis is relatively
easily differentiated from other arenaceous
species by its skeletal architecture, fibre characteristics and spiculation (see remarks for C. (W)
tube rosa below, and compare spicule dimensions
between species in Table 19). Hentschel's (1912)
mention of similar isochelae geometry in C. (W)
clavifonnis and other microcionid species such as
C. (Dendrocia) pyramida is misleading, since the
latter species has arcuate-like chelae whereas
those of C. (W.) claviformis are simply palmate.
The nearest relative of C. (W) claviformis is
probably C. (W) tube rosa, especially in specific
features of its arenaceous ectosome.
Clathria (VVilsonella) ensiae sp. nov.
(Figs 87-88, Table 17, Plate 2E)
MATERIAL. HOLOTYPE: NTMZ3561 (NCIQ66C2384-0 (fragment QMG05004): Marion Reef, off
Edithburg, S. Yorke Peninsula, SA, 35°09.5'S,
137°48.0'E, 10.ii.1989, 6m depth, coll. NCI (SCUBA).
PARATYPE: NTMZ3821 (NCIQ66C-3744-L) (fragment QMG300270): Trap Reef, Bicheno, Tas.,
41°51.7'S, 148°18.6'E, 30m depth, 26.ii.1990, coll.
NCI (SCUBA).
HABITAT DISTRIBUTION. 6-30m depth; on sand
covered rock substrate, with algae and seagrasses on
patch reef; Yorke Peninsula (SA), E coast (Tas) (Fig.
87F).
DESCRIPTION. Shape. Erect, digitate, flabellate
growth form, 205-350mm high, 70-150mm wide,
with multiple branches usually aligned in one
plane, composed of long, slender, flattened or
cylindrical, bifurcate digits, 70-190mm long, up
TABLE 16. Comparison between present and published records of Clathria (Wilsonella) claviformis
Hentschel. All measurements are in p,m, denoted as
range (and mean) of spicule length x spicule width
(N=25).
SPICULE
Choanosomal
principal styles
Subectosomal
auxiliary styles
Echinating
acanthostyles
Holotype
Specimens (N=2)
(SMF1504)
2584308.6)-364 x 6- 235-(287.2)-338 x 7(8.4)-10
(9.6)-13
164-(292.0)-371 x 3- 169-(289.2)-375 x 3(5.1)-8
(5.2)-7
64-(74.0)-82 x 479-(88.8)-103 x 4(6.4)-8
(6.4)-8
Chelae
16-(18.4)-22
14-(16.8)-20
Toxas
28-(140.8)-266 x
0.8-(1.1)-1.5
44-(113.6)-2I8 x
0.8-(0.9)-1.2
to 18nun diameter, frequently fused near their
basal ends, attached to a common base or on a
short cylindrical stalk.
Colour. Red-brown alive (Munsell 2.5R 8/8),
darkening in air (2.5R 6/10), brown preserved.
Oscules. Large oscules, up to 4nun diameter,
scattered evenly over all surfaces of digits, in life
slightly raised above the surface with a
membraneous lip.
Texture and surface characteristics. Surface
even, optically smooth, broken only by raised
oscules; texture firm, compressible, rubbery.
Ectosome and subectosome. Ectosome heavily
arenaceous, with peripheral spongin fibres fully
packed with mostly sand particles and some
foreign spicule fragments (holotype; vice versa in
paratype), and with sparse plumose tracts of subectosomal auxiliary styles, confined completely
below surface; spongin fibres in subectosomal
region ascend to surface, plumose, fully
arenaceous, with plumose brushes of subectosomal auxiliary spicules.
Choanosome. Choanosomal skeleton irregularly
reticulate in axis, plumo-reticulate near
periphery, clearly divided into primary, ascending fibres, 60-130Rm diameter, and secondary,
connecting, transverse spongin fibres, 30-70p..m
diameter; primary fibres fully arenaceous, incorporating both sand and foreign spicule fragments,
and a sparse core of choanosomal auxiliary styles
amongst the debris; secondary fibres without
sand particles, with some foreign spicules, and
also with a light core of choanosomal auxiliary
spicules; echinating acanthostyles not abundant
on fibres, usually echinating fibres at acute
angles, directed towards surface; mesohyl matrix
moderately heavy, with few foreign spicules and
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
191
TABLE 17. Comparison between present and published records of Clathria (Wilsonella) ensiae sp.nov. All
measurements are given in 1.1m, denoted as range (and
mean) of spicule length x spicule width (N=25).
Subectosomal
auxiliary styles
Holotype
(NCIQ66C2384I)
69-(111.4)-132 x
3.5-(4.2)-6
106-(130.7)-147 x
1.5-(2.6)-3.5
97-(132.0)-154 x 2(2.4)-3.5
1888)
(Figs 89-90, Table 18)
F,chinating
acanthostyles
28-(40.7)-54 x 2.5(3.1)-4
48-(61.1)-70 x 3.5(4.3)-5.5
Clathriopsamma reticulata Lendenfeld, 1888: 227;
Chelae
11-(14.2)-16
13-(14.4)-16
Toxas
absent
SPICULE
Choanosomal
principal styles
Paratype
(NCIQ66C3744L)
87-(108.6)-120 x 4(4.4)-5.5
could also be included in C. (Dendrocia) due to
close resemblance between choanosomal and
subectosomal styles, both classed here as
auxiliary spicules (i.e., Wilsonella s.s.). However,
like C. (W) australiensis, those styles coring
fibres differ subtly in their terminations from
those styles outside of fibres.
Clathria (Wilsonella) reticulata (Lendenfeld,
Hallmann, 1920: 771.
Clathria reticulata; Hooper & Wiedenmayer, 1994:
absentzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCB
-
very little sand; choanocyte chambers large, 180480p.m diameter, oval or eliptical.
Megascleres (Table 17). Choanosomal auxiliary
megascleres straight or slightly curved towards
base, slightly subtylote bases, hastate, abruptly
pointed; very similar geometry to subectosomal
auxiliary styles but shorter, slightly thicker.
Subectosomal auxiliary styles short, slender,
straight or rarely slightly curved at centre, slightly
subtylote bases, hastate, stepped points.
Echinating acanthostyles short, slender,
straight, slightly subtylote bases, fusiform points,
spined all over but spines slightly heavier on base
and point.
Microscleres (Table 17). Palmate isochelae relatively long, slender, unmodified.
Toxas absent.
ETYMOLOGY. Phonetic acronym in the National
Cancer Institute (NCI), in appreciation of the AIMS
group who provided the author with unrestricted access
to all their sponge collections.
REMARKS. There are some differences between
the two specimens of C. (W) ensiae in the size of
acanthostyles (Table 17). Similarly, primary
spongin fibres of the holotype are predominantly
cored with sand particles, whereas in the paratype
foreign spicules are more abundant than sand, but
in all other respects both these specimens are
identical, and these observed differences are considered to be relatively minor.
C. (W) ensiae differs from other species of the
Wilsonella group primarily in its flattened-flabelliform, erect, bifurcate, branching growth form
and in having auxiliary styles with peculiar hastate, telescoped points. Other features such as
skeletal structure and spicule dimensions can also
be used to distinguish allied species (Table 19).
Like C. (W) australiensis, the present species
275.
Not Echinochalina reticulata; Whitelegge, 1907: 506,
p1.45, fig.25; Hallmann, 1912: 287, p1.30, fig.2,
text-fig.66.
Not Dictyocylindrus reticulatus Carter, 1881a: 377.
Not Rhaphidophlus reticulatus; Hallmann, 1912: 177.
MATERIAL. LECTOTYPE: AMG9135 (dry): E.
coast of Australia, no other details known. PARALECTOTYPES: AMZ457: E coast of Australia, no other
details known. BMNH1925.11.1.576 (dry): Manly
Beach, NSW, 33°49'S, 151°18'E, no other details
known.
HABITAT DISTRIBUTION. Ecology unknown;
central E coast (NSW) (Fig. 890).
DESCRIPTION. Shape. Subspherical, reticulatebranching growth form, 80-150mm high, 5595nun wide, composed of lobate, bifurcating,
sometimes anastomosing tubular digits with
rounded margins, 30-50nun long, up to 18mm
diameter.
Colour. Dark brown in ethanol.
Oscules. Not seen (available material dry and
surface contracted).
Texture and surface characteristics. Surface
shaggy, reticulate; texture brittle in dry state.
Ectosome and subectosome. Ectosomal skeleton
lightly arenaceous, with plumose brushes or individual choanosomal principal styles protruding, together with a paratangential layer of
subectosomal auxiliary subtylostyles, lying near
bases of principals, and echinating acanthostyles
projecting into these.
Choanosome. Choanosomal skeleton irregularly
reticulate, with moderately heavy, large spongin
fibres forming relatively wide ovoid meshes,
lined by very large, typically curved oxeote toxas;
spongin fibres not easily divisible into primary or
secondary components based on fibre diameter,
although primary, ascending fibres contain
plumose, paucispicular tracts of both principal
192zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 87. Clathria (Wilsonella) ensiae sp.nov. (holotype NTMZ3561). A, Choanosomal auxiliary subtylostyle.
B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Palmate isochelae. E, Section through
peripheral skeleton. F, Australian distribution. G, Paratype QMG300270.
and auxiliary megascleres; secondary, transverse,
connecting fibres without coring spicules; both
sorts of spongin fibres contain a light core of
detritus, especially small sand grains; echinating
acanthostyles very abundant, including
peripheral fibres; mesohyl matrix heavy, darkly
pigmented, with abundant microscleries, especially bundles of whispy, sinuous toxas
(toxodragmata); extra-fibre auxiliary
megascleres organised into ascending subdermal
tracts, with few loose spicules scattered between
fibres.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
193
FIG. 88. Clathria (Wilsonella) ensiae sp.nov. (holotype NTMZ3561). A, Choanosomal skeleton. B, Peripheral
skeleton. C, Fibre characteristics. D, Palmate isochela. E, Echinating acanthostyles. F, Acanthostyle spines. G,
Base of principal subtylostyle. H, Bases of auxiliary subtylostyle.
MEMOIRS OF THE QUEENSLAND MUSEUM
194^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
BC
n
E
m..zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
LO
CV
23
FIG. 89. Clathria (W ilsonella) reticulata Lendenfeld (lectotype AMG9135). A, Choanosomal principal subtylostyle. B, Subectosomal auxiliary subtylostyle. C, Echinating acanthostyle. D, Oxeote toxa. E, Palmate isochelae.
F, Section through peripheral skeleton. G, Australian distribution. H, Lectotype. I, Paralectotype
BMNH1925.11.1.576.
^
195
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
FIG. 90. Clathria (Wilsonella) reticulata Lendenfeld (paralectotype AMZ457). A, Choanosomal skeleton. B,
Peripheral skeleton. C-D, Fibre characteristics. E, Echinating acanthostyle. F, Acanthostyle spines. G, Base of
principal subtylostyle. H, Bases of auxiliary subtylostyles. I. Oxeote toxa. J, Palmate isochelae.
196zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Megascleres (Table 18). Choanosomal principal
subtylostyles well differentiated from auxiliary
spicules, mostly straight, fusiform, with profusely microspined, slightly subtylote bases.
Subectosomal auxiliary subtylostyles thin,
fusiform, straight, slightly curved, sometimes
sinuous, with minutely rnicrospined, subtylote
bases.
Echinating acanthostyles with rounded bases,
relatively even spination, although basal and distal portions slightly more heavily spined than
points.
Microscleres (Table 18). Palmate isochelae differentiated into 2 size classes, smaller with approximately 75% of contort forms.
Toxas extremely abundant, long, thin, sinuous,
characteristically oxeote, with only very slight or
no central curvature and straight tapering points.
REMARKS. This species is poorly known from
3 specimens but differentiated from other species
in the Wilsonella group having only a very light
core of detritus in fibres, usually composed of
small sand and spicule particles, differentiated
primary and secondary fibres, and prominent
bundles of sinuous toxas lining aquiferous chambers (Fig. 90). Affinities with other species (Table
19) are discussed elsewhere (remarks under C.
(W) tuberosa and C. (W) australiensis). In some
respects (growth form, geometry of some
spicules, presence of two sizes of isochelae with
contort forms) the species is closest to C. (W)
tuberosa, but toxa geometry is quite different
between these two species.
Clathria (Wilsonella) tuberosa (Bowerbanlc, 1875)
(Figs 91-93, Table 19, Plate 2F)
Microciona tube rosa Bowerbank, 1875: 281, 282, 286;
Vosmaer, 1935a: 607.
Clathria tuberosa; Ridley, 1881: 121; Ridley, 1884a:
444-445, p1.42, fig.d; Hentschel, 1912: 365-366;
Hooper & Wiedenmayer, 1994: 275.
MATERIAL. HOLOTYPE: BMNH1877. 5.21.1312:
Straits of Malacca, Malaysia, vicinity of 2 ° N, 102 ° E,
coll. Capt. Parish (dredge). OTHER MATERIAL: INDONESIA - SMF978 (fragment MNHNDCL2346).
QLD - BMNH1881.10.21.325, BMNH1882.2.23.198,
253, 283, 334. NT - AMZ4559 (RRIMP FN989),
NTMZ777, NTMZ809, NTMZ920, NTMZ933,
NTMZ946, NTMZ1091, NTMZ2087, NTMZ2400,
NTMZ2708, QMG303336, NTMZ2157, NTMZ2189,
NTMZ1959, NTMZI980, NTMZ1987, NTMZ2020,
NTMZ2098, QMG303428, NTMZ2107, NTMZ234,
NTMZ235, NTMZ236 - NTMZ2496, NTMZ540,
NTMZ107, NTMZ112, NTMZ128, NTMZ554. WESSEL ISLANDS; NT - NTMZ3955.
TABLE 18. Comparison between present and published records of Clathria (Wilsonella) reticulata (Lendenfeld). All measurements are given in p.m, denoted
as range (and mean) of spicule length x spicule width
(N=25).
Lectotype
(AMG9135)
Paralectotype
(AMZ457)
Paralectotype
(BMNH
1925.11.1.576)
Choanosomal 173-(209.4)258 x 7-(9.1)principal
styles
11
182(240.2)289 x 8-(9.3)11
227-(251.5)296 x 8-(9.4)ii
SPICULE
Subectosomal I63-(236.8)1984243.4)187-(259.0)auxiliary
324 x 3-(4.0)-6 322 x 3-(4.2)-5 309 x 3-(4.6)-7
styles
52-(61.7)-71 x 54-(63.6)-75 x 65-(67.8)-72 x
Echinating
acanthostyles
4-(7.2)-9
6-(7.3)-9
3-(6.5)-9
Chelae I
5-(6.6)-9
4-(5.8)-7
6-(6.1)-9
Chelae II
12-(14.4)-17
13-(14.8)-17
14-(15.2)-17
Toxas
238-(402.7)684 x 0.8(2.4)-4
45-(231.3)433 x 0.5(1.9)-3.5
302-(427.7)593 x I -(2.2)-4
HABITAT DISTRIBUTION. Semi-encrusting on
rock, dead or live coral heads, epizootic on other sponges and gorgonians; usually associated with shallow
coral reef habitats; 2-19m depth range; prevalent in the
tropical, Australian and Indo-Malay shallow water
macrobenthic community, extending as far south as
13°S latitude: Torres Strait (FNQ) (Ridley, 1884a),
Bynoe Harbour, Darwin Harbour, Coral Bay, Port
Bremer, Wessel Is (NT). Also Straits of Malacca
(Bowerbank, 1875; Ridley, 1881) and Aru Is, Indonesia (Hentschel, 1912).
DESCRIPTION. Shape. Subspherical,
predominantly bulbous growth form, 60-135mm
diameter, less often club-shaped with apical
lobate digits, or pseudo-vasiform on low stalk
with convoluted, apical, lobate digits; surface
projections (or branches) rounded lobate, relatively close-set, attached to common centre,
which in turn is usually attached to substrate by a
small peduncle; lobate digits usually bifurcate
with rounded margins. In life lobes prominently
bulbous, evenly rounded; after preservation lobes
become slightly flattened and angular.
Colour. Live pigmentation dusty pale pink-red
(Munsell 2.5R 6/10) to pink (5RP 8/6), with a
darker choanosome (5RP 7/8); lighter ectosomal
colouration due to arenaceous nature of ectosome; dessicated colouration darkens to brown
(2.5Y 8/2), red-brown (5Y 8/4), or red-purple
(5RP 3/6), as paler ectosome collapses. In situ,
subdermal ridges and canals red-pink in life,
showing darker choanosomal pigmentation.
Oscules. Exhalant pores variable in diameter,
ranging from 1-3mm, each with prominent,
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
197
Clat hria ( Wilsonella) t uberosa (Bowerbank) (NTMZ2157). A, Choanosomal principal subtylostyle. B,
FIG. 91. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Echinating acanthostyle. C, Subectosomal auxiliary subtylostyles. D, Longer accolada and smaller wing-shaped
toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Known Australian distribution. H,
Specimen of Hentschel (1912) SMF978. I, Specimen of Ridley (1884) BMNH1882.2.23.198. J, NTMZ107 in
sit u.
198zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 92. Clathria (Wilsonella) tuberosa (Bowerbank) (NTMZ2708). A, Choanosomal skeleton. B, Fibre
characteristics (x164). C, Echinating acanthostyles. D, Acanthostyle spines. E, Bases of principal subtylostyles.
F, Palmate isochelae. G, Base of auxiliary subtylostyle. H, Longer accolada toxas. I. Smaller wing-shaped toxas.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
199
SEASON
TOTAL SAMPLES
SAMPLES WITH
LARVAE
WET
3
2
PREDRY
5
5
DRY
10
0
PREWET
10
5
FIG. 93. Clathria (Wilsonella) tube rosa (Bowerbank).
Seasonal production of incubated larvae in populations from NW Australia.
membraneous, raised lip (collapses upon preservation); oscules confined to apex of lobate digits,
with subdermal canals and grooves radiating
away from pores in cruciform pattern.
Texture and surface characteristics. Surface optically smooth, even in situ, distinctly
membranous; ectosomal membrane transparent
or slightly opaque when intact, stretched across
adjoining lobes, with darker subdermal pigmentation and fibre reticulation clearly visible below;
subdermal grooves and minute subdermal canals
produce a more-or-less microscopically reticulate surface; upon dessication ectosome collapses
to become optically reticulate, distinctly
arenaceous, with convoluted ridges and conules,
and large amounts of clear mucous usually
produced; texture stiffly compressible,
arenaceous, harsh to touch, minutely hispid.
Ectosome and subectosome. Ectosomal skeleton
heavily arenaceous, with delicate traces of sand
coring peripheral fibres, through which protrude
sparse tufts of subectosomal auxiliary styles,
usually raised on low surface conules; special
ectosomal megascleres absent; subectosomal
auxiliary subtylostyles also form tangential or
paratangential tracts perpendicular to dermal
crust, intermingled with foreign particles
auxiliary spicules variable in size, but no distinct
localisation of smaller or larger forms; subectosomal region obscured by abundant sand grains
coring peripheral subdermal fibres; individual
extra-fibre auxiliary styles are intermingled
amongst sand and fibres in subdermal region,
sometimes forming dense paratangential
plumose brushes, ascending to ectosome, but
usually producing sparse tangential subdermal
tracts; subdermal tracts clustered tightly around
fibres and sand matrix, bound together with abundant collagen; on peripheral fibres, choanosomal
principal spicules produce plumose brushes,
sometimes protruding through surface, but usually only obvious in places where ectosome has
collapsed and peripheral fibres are closest to surface.
Choanosome. Choanosomal skeleton irregularly
reticulate, with light spongin fibres fully cored by
sand grains and fewer choanosomal principal
megascleres, the latter in rows of 6-10 abreast in
larger fibres; spicule fragments also common
amongst detritus, particularly haplosclerid oxeas;
spongin fibres heavily echinated by acanthostyles; fibre branching produces irregular oval
meshes, 50-(334)-600p.m diameter, with irregular eliptical choanocyte chambers (38121p,m diameter), with light mesohyl matrix and
abundant microscleres, without sand or any
megascleres; spongin fibres not clearly divisible
into primary or secondary elements, but thinner
fibres (30-551J,m diameter) have coring
megascleres more visible (fewer detrital particles); larger spongin fibres 70-(104)-230p„m
diameter; megascleres core fibres in
paucispicular tracts, slightly more heavily aggregated in thicker fibres but partially obscured
by sand particles; mesohyl matrix in axis light,
with little foreign debris or auxiliary megascleres.
Megascleres (Table 19). Choanosomal principal
subtylostyles slightly curved towards basal end,
occasionally straight, with heavily microspined
bases, tapering to sharp fusiform points.
Subectosomal auxiliary subtylostyles straight,
variable in size, usually with microspined,
prominently subtylote bases, sharply tapering,
fusiform points.
Acanthostyles very variable in length and
width, straight, subtylote, fusiform, evenly
spi nose with granular spines (thinner spicules) or
heavy thorn-like spines (thicker spicules).
Microscleres (Table 19). Palmate isochelae typically very abundant, incompletely divided into
two size classes, with some twisted smaller examples.
Toxas abundant, thin, usually long, without
reflexed points, only slightly curved at centre,
although smaller examples may have more angular central curvature; occurring individually or
more often in toxodragmata within mesohyl
matrix.
Larvae. Incubated parenchymella larvae were
recorded in only 17% of specimens, collected
from Darwin and Cobourg Peninsula regions, NT,
during May, September, October and December,
suggesting a possible breeding period during the
wetter months (Fig. 93). Larvae orange-brown
pigmented, oval - elongate, ranging from 165280 x 110-160iLm. All larvae contained juvenile
megascleres scattered throughout central portion
of mesohyl, usually with heavy collagen. In the
few adult sponges seen incubating larvae, the
^ MEMOIRS OF THE QUEENSLAND MUSEUM
200zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
mesohyl was usually lightly orange in colour,
whereas in non-fecund specimens the mesohyl
matrix was invariably pigmented light brown.
Associations. Scyllid poly chaete worms
(Typosyllis spongicola) relatively common (31%
of specimens examined).
Morphological variation. Gross morphology:
characteristic, relatively consistent, varying only
in elongation of basal stem and clump of digits on
apex, ranging from low spherical bulbs (69%),
club-shaped (16%), or convoluted, semivasiform
growth forms (15%). Live colouration: consistent, only slight variation in pinkish hue. Extosomal skeletal structure: extends from densely
arenaceous with few dermal spicule brushes
(39%), arenaceous with abundant protruding
plumose subectosomal brushes (35%), to
arenaceous with plumose brushes of subectosomal spicules and single choanosomal styles
erect on surface (26%). Subectosomal skeletal
structure: varies from well-developed, plumose
ascending spicule tracts composed of subectosomal auxiliary styles (27%), few plumose
extra-fibre brushes with most extra-fibre spicules
closely bound by collagen (31%), or with all
extra-fibre spicules simply bound closely but external to spongin fibres (42%). Choanosomal
skeletal structure: relatively consistent, ranging
from simply irregularly reticulate (54%) to
regularly reticulate (46%), with oval meshes
(88%) or less commonly longitudinally elongated
meshes (12%). Mesohyl matrix: lightly pigmented (58%), moderately heavily pigmented
(23%), or heavily pigmented collagen (19%).
Echinating acanthostyles: forming dense (42%),
moderately dense (50%), or only lightly echinating fibres (8%). Megasclere geometry: consistent; majority of choanosomal subtylostyles with
microspined bases, 8% of specimens with smooth
subtylote bases. Acanthostyles vary only in
dimensions. Microsclere geometry: consistent although proportion of contort versus unmodified
TABLE 19. Comparison between morphological characteristics of some Australasian arenaceous Clathria
(Wilsonella) species, based on present and published records. All measurements are given in p.m.
MATERIAL
CHARACTER
1
2
3
4
5
6
Shape
massive
subspherical
tubular
elongate
subspherical
tubular
clay iform
digitate flabellate
digitate
fan
Digits
lobate
lobate
clubs/bulbs
cylindrical or
flattened
cylindrical
lobate
Live colour
pale orange
pale orange
red-brown
red-brown
unknown
unknown
Skeletal
architecture
irreg. relic
irreg. relic
regular relic
irreg. retic.
irreg. relic.
Differentiated
primary/seconda
ry fibre system
irreg. relic, axis,
plumo-,regular
relic. subect.
yes
yes
yes
yes
yes
no
Choanosomal
styles
89-175 x2.5-8
smooth or spined
base
96-214x4-6.5
smooth or spined
base
235-364 x6-13
spined base
69-132 x3.5-6
smooth base
172-388 x6-16
smooth base
192-298 x6-11
spined base
Subectosomal
styles
92-172 x1.5-5
spined apex and
base
99-148 x2.5-3.5
spined apex and
base
164-375 x3-8
spined base
97-154 x1.5-3.5
smooth base
162-541 x3-7
smooth base
160-325 x2.5-5.5
spined base
49-85x2-6
52-63x3-5.5
64-103x4-8
28-70x2.5-5.5
63-94x4-8
78-94x3-9
11-19
12-15
14-22
11-16
14-18
12-18
absent
absent
absent
absent
absent
3-8 twisted
28-89 x 0.5-1.5
angular centre
43-59 x0.8-1.5
angular centre
28-266 x 0.8-1.5
slightly angular
centre
absent
57-74 x1-2
rounded centre
91-435 x0.8-2
straight at centre
Acanthostyles
Large chelae
Small chelae
Toxas
Material:
1. W. australiensis (Carter) - present study.
2. C. lobosa Lendenfeld (1888:149) (=W. australiensis) zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFED
holotype AMU G9053; Port Jackson,NSW.
3. W. clavifonnis (Hentschel) - present study.
4. W. ensiae, sp.nov. - present study.
5. W. ramosa (Lindgren, 1897:482- schizotype BMNH 1929.11.26.48.
6. W. mixta (Hentschel, 1912:298) - holotype SMF 974.
-
201
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
fibres. Nevertheless, the two species are closely
related.
Clathria (W) australiensis has choanosomal
fibres divided into ascending and radial primary
elements, fully cored by detritus, with less heavily cored secondary transverse elements. Subectosomal auxiliary subtylostyles have spines on
both bases and points, typically with a prominent
terminal spine/point and also smaller spines occuring at least part of the way along the shaft.
Choanosomal subtylostyles are also auxiliary
spicules, with smooth or spined bases only. Acanthostyles are pointed and tapering; there is only
one size of isochelae; and toxas have strongly
curved (rounded or sharply angular) central curves.
Clathria (W) rugosa is a sister species of C.
(W) australiensis, differing in having prominent
subectosomal drainage canals ('astrorhizae').
Subectosomal auxiliary styles are also spined on
both ends, but spines are perched only on the very
extremity of the spicule point (not on the shaft);
acanthostyles have bulbous points; there is onlyzyxwvutsrqp
small isochelae varies from 0-20% of contort
spicules (12%), 20-40% (19%), 40-60% (24%),
60-80 (35%), to 80-10% of spicules (12%).
Spicule dimensions: Few specimens atypical but
variation apparently random with no statistical
significance between specimens irrespective of
seasonal or geographical distribution of samples.
REMARKS. C. (W) tuberosa is distinctivein the
field: pink colour, bulbous growth form, soft texture. However, it is more difficult to differentiate
descriptively. Pertinent differences are:
Choanosomal architecture and fibre characteristics of C. (W) reticulata are identical to those
of C. (W) tuberosa; acanthostyles are as equally
abundant in both species, but many acanthostyles
have heavier spines on the distal part than on
points; and there are two sizes of isochelae, 75%
of the smaller being contort. Thus, the major
features distinguishing the two taxa are the
straight or sinuous oxeote toxas in C. (W)
reticulata, which are never present in C. (W)
tuberosa, and the light deposits of debris into
MATERI AL
CHARACTER
Shape
7
8
9
fan palmate-
massive spherical
massive spherical
digitate
tubular
globular
cylindrical
Digits
10
11
elongate
massive spherical
subspherical
globular
tubular
cupriform
lobate
lobate
lobate
lobate
pale pink or pink-
pale pink or pink-
pale pink or pink-
pale pink or pink-
red
red
red
red
Live colour
bt.red-orange
Skeletal
architecture
irreg.retic.
irreg.retic.
irreg.retic.
irreg.retic.
yes
none
none
none
134-159x4.5-8
167 - 278 x4 - 11
144 - 286x4 - 8
l72-302x6-8.5
smooth base
most spined bases
most spined bases
most smooth bases
Differentiated
primary/ seconda
ry fibre system
Choanosomal
styles
162-206 x2.5-4
Subectosomal
styles
spined apex and
Acanthostyles
58-91 x4-7
none
1334226.3)-343
x4-(8.2)-14 most
spined bases
Small chelae
lobate tubular
unknown
partial
173-296 x7-11 all
spined bases
163-324 x3-7
154-324 x2-5
148-223 x2-5
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
x1.5-(4.1)-8
159 - 333 x2-6
spined bases
spined bases
spined bases
44-97 x4-9
43-75 x3-8
69-88 x3.5-5
base
spined bases
spined bases
(6.7)-11
52-75 x3-9
12 - 17
11 - 19
11 - 16
13 - 19
10 - (13.3) - 18
14 - 17
absent
4-8 twisted
4-7 twisted
6-8 rare
4-(6.1)-9 twisted
4-9 twisted
24-122 x0.8-3
Toxas
reticulate branches
129-(228.9)-375
60-(79.9)-112 x4-
Large chelae
12
slight curve at
centre
55-255 x0.5-1.5
rounded centre
95-215 x0.5-1.5
angular& rounded
centre
30-(140.3)-388
81-94 x1-2
rounded centre
x0.4-(1.I )-2
45-684 x0.5-4
sinuous oxeote
rounded centre
Material:
7. W. rugosa (Hooper & Levi, 1993) - New Caledonian population (included for comparative purposes).
8. W. t uberosa (Bowerbank, 1875:281); holotype BMNH 1877.5.21.1312; Straits of Malacca (19=25).
9. W. t uberosa; Ridley, 1884a:444; BMNH 1881.10, 21.325, 1882.2.23.198, 253, 283, 334; Tones Strait (N=125).
10. W. t uberosa; Hentschel, 1912: 365; SMF 978; Arafura Sea (N= 25).
11. W. t uberosa; present material - NW Australia (N= 650).
12. W. r et iculat a (Lendenfeld) - .present st udy.
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDC
202zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
a single small category of isochelae; and toxas are
only slightly curved at their centre.
Clathria (W) clavtformis, C. (W) ensiae, and
C. (W) ramosa are different from these other
species in spicule geometry, spicule size, skeletal
architecture and fibre characteristics. These IndoAustralasian species are relatively easy to distinguish from their gross morphology and field
characteristics although their skeletal characters
are usually at least partially obscured by the incorporation of sand into fibres.
With the exception of C. (W.) australiensis
species of Clathria (Wilsonella) have relatively
restricted, mostly allopatric distibutions: C. (W)
tube rosa is from N Australia and SE Asia; C. (W)
claviforrnis from the Arafura and Timor Seas; C.
(W) mixta from 2 disjunct populations in the
Arafura Sea and the S. Arabian coast; C. (W)
ramosa from the Java Sea; C. (W) ensiae in S
Australia and Tasmania; C. (W) reticulata SE
Australia. C. (W.) australiensis ranges from SW
Australia to S Queensland, whereas its cryptic
sibling C. (W) rugosa is restricted to New
Caledonia.
OTHER SPECIES OF CLATHRIA
(WILSONELLA)
Clathria (Wilsonella)cercidochela (Vacelet &
Vasseur, 1971)
Clathriopsammacercidochela Vacelet& Vasseur, 1971: 104105, text-fig.62, p1.3, fig.! [Tulear, Madagascar].
MATERIAL. HOLOTYPE: MNHNDJV24. W Indian Ocean.
Clathria (Wilsonella) ferrea (de Laubenfels,
1936)
Fisherispongiaferrea de Laubenfels, 1936b: 460, fig.44. [Atlantic coast of Panama].
Clathria (Microciona) ferrea; Van Soest, 1984b: 101-103,
text-fig.40, table 4 [Curacao].
Clathria ferrea; Zea, 1987: 172, text-fig.60 [Colombian
Caribbean].
Microciona ferrea; Pulitzer-Finali, 1986: 149 [West Indies].
MATERIAL. HOLOTYPE: USNM22239. Caribbean.
Clathria (VVilsonella) foraminifera (Burton &
Rao, 1932)
Aulenella foraminifera Burton & Rao, 1932: 345-346, p1.18,
fig.11, text-fig.11 [Gaspar Straits, Java Sea].
MATERIAL. HOLOTYPE: IMP790/1. E Indonesia.
Clathria (VVilsonella) lindgreni sp. nov.
Clathria ramosa Lindgren, 1897: 482-483; Lindgren, 1898:
308-309, p1.17, fig.9, p1.18, fig.15, p1.19, fig.16 [Belitung
I., Java Sea]; Hentschel, 1912: 367.
Thalysias ramosa; de Laubenfels, 1936a: 105.
Not Rhaphidophlus ramosus Kieschnick, 1896: 533;
Kieschnick, 1900: 53-54, p1.45, figs 47-50.
Not Echinoclathria ramosa; Hallmann, 1912: 277, p130,
fig.3.
Not Wilsonella ramosa; Hallmann, 1912: 243, 298.
Not Colloclathria ramosa Dendy, 1922: 74-76.
cf. Microciona prolifera tropus spinosa; Vosmaer, 1935a:
642.
MATERIAL. HOLOTYPE: NHRM (fragment
BMNH1929.11.26.48). Indonesia. Clathria (Thalysias)
ramosa (Kieschnick, 1896) has priority.
Clathria (Wilsonella) litos Hooper & Levi, 1993
Clathria (Clathriopsamma) litos Hooper & Levi, 1993a:
1243-1246, figs 9-10 [New Caledonia].
MATERIAL. HOLOTYPE: QMG301269. SW Pacific.
Clathria (Wilsonella) mixta Hentschel, 1912
Clathria mixta Hentschel, 1912: 298, 367, 368, p1.13, fig.8,
p1.19, fig.30 [Aru I., Arafura Sea]; Burton, 1959a: 244 [S.
Arabian coast].
Thalysias mixta; de Laubenfels, 1936a: 105.
cf. Clathria lobata or Clathria ulmus; Vosmaer, 1935a: 649.
MATERIAL. HOLOTYPE: SMF 974 (fragment
MNHNDCL2280). Indonesia, Arabian Gulf.
Clathria (Wilsonella) pseudonapya (de
Laubenfels, 1930)
Clathriopsamma pseudonapya de Laubenfels, 1930: 28; de
Laubenfels, 1932: 96-97, text-fig.57 [Pacific Grove,
California]; Sim & Bakus, 1986: 10 [California].
MATERIAL. HOLOTYPE: USNM21436. PARATYPE
BMNH1929.8.22.19. NE Pacific rim.
Clathria (Wilsonella) rugosa Hooper & Levi,
1993 (Table 19)
Clathria (Clathriopsamma) rugosa Hooper & Levi, 1993a:
1237-1243, figs 7-8, tables 4-5 [New Caledonia].
MATERIAL. HOLOTYPE: QMG300278 (fragment
NTMZ3880). PARATYPE QMG300696 (fragment
NTMZ3889). SW Pacific.
Clathria (Microciona) Bowerbank, 1862
Microciona Bowerbank, 1862b: 1109.
[Abila] Gray, 1867: 539 [preocci.
[Aaata] de Laubenfels, 1930: 271preocci.
Anaata de Laubenfels, 1932: 89.
Axocielita de Laubenfels, 1936a: 118.
Cionanchora de Laubenfels, 1936a: 108.
Fisherispongia de Laubenfels, 1936b: 460.
Folitispa de Laubenfels, 1936a: 119.
Hymantho Burton, 1930a: 503.
Hymeraphia, in part, Hentschel, 1912: 377; not
Hymeraphia Bowerbank, 1864: 189.
Leptoclathria Topsent, 1928a: 298.
Ophlitaspongia Bowerbank, 1866: 14; not Ophlitaspongia of authors.
Paratenaciella Vacelet & Vasseur, 1971: 103.
Pseudanchinoe Burton, 1929a: 433.
Seriatula Gray, 1867: 515
Sophax Gray, 1867: 521.
Wetmoreus de Laubenfels, 1936a: 112.
DEFINITION. Persistently encrusting growth
form, with hymedesmoid skeletal architecture
consisting of a basal layer of spongin, typically
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
203
with ascending, plumose, non-anastomosing,
spongin fibre nodes, and megascleres embedded
and erect on basal layer; ectosomal skeleton with
only a single undifferentiated category of
auxiliary megasclere.
TYPE SPECIES. Microciona atrasanguinea Bowerbank, 1862b: 1109 (by subsequent designation of
Bowerbank, 1864: 188).
REMARKS. Of 118 named species described in,
or subsequently referred to Microciona or one of
its synonyms, 103 appear to be valid, and 7 are
recorded from Australasia, including 2 new
species.
Clathria (Microciona) aceratoobtusa
(Carter, 1887)
(Figs 94-95, Table 20, Plate 3C)
Microciona acerato-obtusa Carter, 1887: 62, 67, 83,
p1.5, figs 7-10; Dendy, 1896: 18; Hentschel, 1911:
348, text-fig. 32a-f.
Axocielita aceratoobtusa; de Laubenfels, 1936a: 118.
Clathria aceratoobtusa; Rudman & Avern, 1989: 335;
Hooper & Wiedenmayer, 1994: 266.
cf. Microciona prolifera; Vosmaer, 1935a: 608, 637.
MATERIAL. HOLOTYPE: LFM (confirmed
destroyed during WWII): Kadan Kyun (King I.), Mergui Archipelago, Andaman Sea, Burma. NEOTYPE:
NTMZ3676: NW. side of N. I., Ko Wao Yai Group,
vicinity of Ko Samui, Gulf of Thailand, 9°46.7'N,
99°40.3'E, 12m depth, 6.vi.1990, coll. J.N.A. Hooper
(SCUBA). OTHER MATERIAL: NSW - NTMZ2835
(fragment QMG300543), NTMZ3125. QLD QMG303089, QMGL713 (fragment NTMZ1536).
SAHUL SHELF, WA - QMG301083, QMG301188.
INDONESIA - BMNH1946.11.25.244.
HABITAT DISTRIBUTION. Coral rubble, rock and
bivalve substrata; intertidal-14m depth; Shark Bay,
Cartier I., Hibernia Reef, Sahul Shelf (WA) (Hentschel,
1911, present study); Cairns and Shelburne Bay (FNQ)
(present study); Sydney and Iluka (NSW) (Rudman &
Avern, 1989, present study) (Fig. 940). Also IndoMalay Archipelago — Andaman Sea (Burma) (Carter,
1887), Gulf of Thailand (present study).
DESCRIPTION OF NEOTYPE. Thinly encrusting on bivalves, up to lnun thick; colour orangered alive (Munsell 1 OR 6/12); firm texture;
oscules not seen; surface microscopically hispid,
with choanosomal principal styles protruding up
to 100p,m from ectosome; subectosomal
auxiliary styles lie paratangential to surface, in
bundles or individually; choanosomal skeleton
leptoclathriid, with principal styles and echinating acanthostyles embedded in and perpendicular
to basal spongin fibres; principal styles form
plumose brushes, and both sorts of spicules also
scattered individually in skeleton; mesohyl
matrix heavy, dark brown, granular, with incorporated detritus, numerous toxas and auxiliary
styles dispersed; principal choanosomal styles
long, fusiform, rounded or very slightly subtylote, with smooth or minutely spined bases
(length 175-548p.m, width 11 22 p,m); subectosomal auxiliary subtylostyles polytylote, with
microspined swollen bases (length 264-387p.m,
width 1.5-4.5p,m); echinating styles short, slightly curved, robust, with prominently swollen,
usually rnicrospined bases and smooth shafts
(length 128-183p,m, width 5-12p,m); palmate
isochelae small, relatively homogeneous in size,
with many twisted forms (9-14p.m long); toxas
short, thickest at centre, tapering to sharp, slightly
reflexed points (length 58-92p,m, width 25.5p,m).
-
DESCRIPTION. Shape. Thinly encrusting, contiguous or discrete mats on rock or coral substrata, covering up to 120mm2, 0.4-2mm thick.
Colour. Bright orange-red alive (Munsell lOR
6/12-14), grey-brown in ethanol.
Oscules. Small exhalant apertures unevenly distributed over surface, up to 1.5mm diameter,
slightly raised or flush with surface; small
membraneous lip surrounding oscules when
alive, collapsing in air. Minute inhalant pores
irregularly dispersed, producing slightly reticulate appearance.
Texture and surface characteristics. Firm,
mucusy alive, minutely hispid; surface with irregularly dispersed, bifurcate subdermal
drainage canals meandering from oscules.
Ectosome and subectosome. Ectosomal skeleton
hispid, with points of large choanosomal principal styles protruding up to 200p.m from surface,
occurring individually or in paucispicular
plumose brushes of about 5 spicules; subectosomal auxiliary styles usually lie paratangential
to surface, sometimes forming tangential tracts
lying immediately subdermal; auxiliary styles
arise from skeleton at oblique angles, rarely
protruding through ectosome; tracts of auxiliary
spicules originate in basal half of skeleton, with
8-12 spicules per tract.
Choanosome. Choanosomal skeletal architecture
hymedesmoid in thin sections, microcionid in
thicker regions, with a relatively thick layer of
heavy spongin fibre lying on basal substrate, 2243 um diameter; bases of principal and echinating
styles embedded in basal spongin, perpendicular
to substrate, individually or in plumose bundles;
^ MEMOIRS OF THE QUEENSLAND MUSEUM
204zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
D1
FIG. 94. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hria ( Microciona) acerat oobt usa (Carter) (NTMZ2835). A, Choanosomal principal subtylostyles.
B, Polytylote subectosomal auxiliary subtylostyles. C, Echinating subtylostyles. D, Oxhorn toxas. E, Palmate
isochelae. F, Section through peripheral skeleton. G, Australian distribution.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
205
FIG. 95. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hria ( Microciona) acerat oobt usa (Carter) (QMG303089). A, Choanosomal skeleton. B, Close view
of hymedesmoid skeleton. C, Bases of choanosomal principal subtylostyles. D, Polytylote base of auxiliary
subtylostyle. E, Palmate isochela. F, Oxhorn toxa.
MEMOIRS OF THE QUEENSLAND MUSEUM
206^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
TABLE 20. Comparison between present and published records of Clathria (Microciona) aceratoobtusa
(Carter). All measurements are given in p,m, denoted
as range (and mean) of spicule length x spicule width
(N=25).
SPICULE
Choanosomal
principal
styles
Subectosomal
auxiliary
styles
Neotype
(NTMZ3676)
175(386.0)548 x11(15.5)-22
264-(324.3)387x 1.5(3.8)4.5
Hentschel
(1911)
up to 408 x 8
168 x 4
Specimens
(N=6)
235-(352.8)492 x12.5(17.2)-22
217(379.8)443 x 3.5(4.3)-6
97-(
194 x 5.2(6.8)-9
128-(149.9)183 x 5-(7.2)12
from 64 x 5
Chelae
9-(12.2)-14
9-12
10.5-(13.5)-16
Toxas
58-(71.5)-92 x
2-(3.1)-5.5
35-92 x 3-4
18-(57.3)-84 x
0.8-(2.2)-3.5
styles
in thick sections basal spongin fibres form small
erect nodes, 22-35 p.m thick, up to 48p,m high,
enveloping bases and parts of spicule shafts;
mesohyl matrix heavy, granular, darkly pigmented, incorporating irregularly dispersed sand
grains and other foreign debris, numerous toxas
occurring singly or in dragmata, more-or-less ascending tracts of subectosomal auxiliary
megascleres, and fewer isochelae; choanocyte
chambers minute, ovoid but rarely seen, 1218p,m diameter, mostly obscured by heavy collagen; large subectosomal cavities, 110-145p.m
diameter, visible where inorganic substrate is
fragmented and discontiguous.
Megascleres (Table 20). Choanosomal principal
subtylostyles long, thick, fusiform, typically
curved in basal third, with slightly subtylote
mostly smooth, less often microspined bases.
Subectosomal auxiliary subtylostyles usually
long, straight, thin, fusiform, with prominently
swollen, smooth or microspined bases.
Echinating subtylostyles entirely smooth or
occasionally with lightly microspined bases,
small, thick, fusiform, slightly curved or straight.
Intermediates between echinating and principal
styles also occur.
Microscleres (Table 20). Palmate isochelae small,
with long lateral alae fused to shaft for most of its
length; chelae relatively common, of a single size
class, homogeneous in size and geometry, approximately 70% with contort shafts.
Toxas very abundant, oxhorn, with slightly
rounded central curves, straight or slightly
reflexed points; central part thickest whereas tips
taper to fine points.
Associations. On the NSW coast this species has
been found in association with pairs of
nudibranchs grazing on the sponge, Rostanga
arbutus (AMC151078, 154589) (W.B. Rudman,
pers.comm.). These predators are identical in
their live colouration to the sponge, presumably
utilising the sponge's carotenoid pigments.
REMARKS. This species was originally
recorded from Mergui Archipelago and by
Hentschel (1911) from Shark Bay, WA. The
holotype was destroyed during WWII (its absence from the LFM collections has been checked
by Shirley Stone, BMNH, pers.comm.); the
neotype comes from an area in Thailand relatively close to the type locality. The first record of the
species in the Pacific Ocean is also made here.
Previous published descriptions of this species
are relatively poor and non-discriminatory; some
attributes of the type material are still uncertain.
Carter (1887) did not give any spicule dimensions, but his figures indicate that Mergui
specimens are very similar to present material.
There are some minor differences between my
material and descriptions by Carter (1887) and
Hentschel (1911). Hentschel's specimens from
Shark Bay were thickly encrusting with
stoloniferous, mammiform surface processes.
Choanosomal architecture varied from leptoclathriid, with a thin layer of spongin lying on
the substrate, to microcionid in thicker regions,
with fibre nodes and single, non-anastomosing
columns of spongin arising from the substrate.
Principal styles were fusiform, prominently subtylote, often with microspined bases. Palmate
isochelae were frequently contort. By comparison, Carter (1887) reported the holotype had
principal styles with hastate or styloid points, and
their bases were completely smooth and only
slightly subtylote. Similarly, there was no mention in Carter's description whether isochelae
were modified (contort). Vosmaer (1935a) expressed doubts about the conspecificity between
Carter's and Hentschel's material based on alleged differences between them in megasclere
and microsclere geometries, but this criticism is
unfounded. Both Carter and Hentschel reported
that their specimens were thinly encrusting on
living and dead serpulid worm tubes, bivalves
and gastropods; colour was brown to beige
preserved; toxa geometry was distinctive and
identical; and echinating megascleres were entirely smooth.
De Laubenfels (1936a) erected Axocielita for
this species, having smooth echinating
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP
^
207
TABLE 21. Spicule dimensions of of Clathria
(Microciona) antarctica (Topsent), giving comparisons between nominotypical material and other
type material. All measurements are given in p.m,
denoted as range (and mean) of spicule length x
spicule width (N=25).
Holotype of S.
SPICULE
Part of type of M.
toxtfera
basispinosa
(MNHNDT1612)
(BNMN1933.3.1739)
Choanosomal
principal styles
409-(519.9)-676 x 9- 293-(498.3)-618 x 9(13.8)-22
(12.6)-18
Subectosomal
auxiliary styles
2I3-(424.9)-899 x 4- 252-(360.9)-503 x 4(10.5)-16
(7.2)-10
52-(110.8)-2I4 x
2.5-(6.8)-10
78-(130.1)-265 x 3(8.7)-11
Chelae
absent
absent
Toxas
31-(46.1)-84 x 0.8(1.8)-3.0
18-(27.8)-35 x 1.5(2.3)4.0
Echinating
acanthostyles
megascleres (i.e., like Axociella) but also with a
Microciona-like encrusting growth form. This is
surprising given that Ophlitaspongia seriata, a
north Atlantic species, also has an encrusting
growth form, microcionid architecture and
smooth echinating spicules (which Simpson
(1968a) subsequently showed was a synonym of
Microciona based on cytology and other characters). These arguments demonstrate that the apparent generic boundaries between microcionids
based on growth form and spicule spination are
tenuous at best.
QMGL713 from the Cairns region, encrusting
on an ascidian, has skeletal architecture, fibre
characteristics and spiculation closely comparable to other material of C. (M.)aceratoobtusa
but lacks toxas completely. It is difficult to confirm the identity of this specimen (i.e., because
the species is largely characterised by the
geometry of its toxas), but given that all other
characters are the same it is included here.
Although only known from few specimens it is
likely that this species is widespread in the Indowest Pacific shallow-water fauna.
Clathria (Microciona) antarctica (Topsent, 1917)
(Figs 96-97, Table 21)
Stylostichon toxiferum Topsent, 1913a: 621-622, p1.4,
fig.7, p1.6, fig.14.
Not Hymeraphia toxifera Hentschel, 1912: 382.
Anchinoe toxifera var. antarctica; Topsent, 1917: 43,
p1.4, fig.5, p1.6, fig.5.
Pseudanchinoe toxifera; Burton, 1929a: 433-434; Burton, 1932a: 325; Burton, 1934b: 39; Burton, 1940:
115; Koltun, 1964a: 72; Koltun, 1976: 155, 188, figs
11-12.
Clathria toxifera; Van Soest, 1984b: 129.
Pseudanchinoe toxiferum; Koltun, 1976: 155, 188.
Stylostichon tuberculata Burton, 1934b: 35, p1.3, fig.2,
text-figs 6-9; Koltun, 1976: 188.
Microciona basispinosa Burton, 1934b: 38-39, p1.5,
fig.2, text-figs 11-12; Burton, 1938b: 17; Koltun,
1964a: 76; Desqueyroux, 1972: 31, figs 103-107;
Desqueyroux & Moyana, 1987: 49; Dawson, 1993:
36.
Clathria antarctica; Hooper & Wiedenmayer, 1994:
266.
MATERIAL. HOLOTYPE: MNHNDT1612: Gough
I., S. Atlantic, 40°20'S, 95°6.3'W, 200m depth,
22.iv.1904, coll. R.R.V. 'Scotia' (dredge).
HOLOTYPE of M. basispitzosa: NHRM997 III (fragment BMNH 1933.3.17.39): Port Albemarle, Falkland
Is, 18-30m depth, 11.ix.1902, Swedish Antarctic Expedition (dredge).
HABITAT DISTRIBUTION. Deeper water rock reefs;
16-610m depth (Koltun, 1976); Antarctica - Discovery
Inlet, Ross Sea, McMurdo Sound, Graham Land, Victoria Land, Enderby Land; Subantarctic — Macquarie I.
(Fig. 96F). Also SW Atlantic; SW and SE Pacific —
Tierra del Fuego, Falkland Is, Shag Rocks, South Georgia, Gough I., Argentina, Chile, Kerguelen, New
Zealand.
DESCRIPTION. Shape. Growth forms range
from thickly encrusting to massive, subspherical.
Colour. Brown in ethanol.
Oscules. Small, 2mm diameter, on apex of surface conules.
Texture and surface characteristics. Compressible; smooth surface with scattered prominent
conules, translucent surface.
Ectosome and subectosome. Moderately dense
plumose brushes of auxiliary subtylostyles, of a
single size category, arising from ends of erect
fibre nodes, protruding through surface and interdispersed with long principal spicules.
Choanosome. Skeletal architecture microcionid,
with hymedesmoid basal layer of spongin lying
on substrate echinated by erect acanthostyles of
various sizes, and erect fibre nodes at 200-40011An
intervals; fibre nodes non-anastomosing, forming
discrete skeletal columns in choanosome; erect
fibre columns, 40-100p,m diameter, cored by
long choanosomal subtylostyles, usually protruding through fibres in plumose bundles or individually, and also heavily echinated by smaller
acanthostyles in their basal portion only; fibres
form single, discrete columns of spongin and
spicules for most of their length but diverge into
2 or more branches in subectosomal region, ultimately producing ectosomal spicule brushes at
their ends; mesohyl matrix light, choanocyte
chambers 30-50p.m diameter, numerous spherical cells, and dispersed auxiliary spicules outside
MEMOIRS OF THE QUEENSLAND MUSEUM
208zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
E
D.
0
C\J
Clat hria ( Microciona) ant arct ica (Topsent) (holotype MNHNDT1612). A, Choanosomal principal
FIG. 96. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
subtylostyle. B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Wing-shaped toxas. E,
Section through skeleton. F, Antarctic distribution.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
209
fibres also form plumose columns;
mesohyl collagenous.
Megascleres (Table 21). Choanosomal principal subtylostyles
very long, slender, with fusifonn
points, slightly curved towards
apical end, subtylote or slightly
subtylote bases, smooth or
microspined bases.
Subectosomal auxiliary subtylostyles short, robust, straight or
slightly curved near basal end,
hastate points or at least less
fusiform than principal spicules,
with subtylote microspined bases.
Echinating acanthostyles variable size range, the larger ones
clearly intermediate between principal spicules and smaller spined
spicules; spicules straight or
slightly curved, moderately heavily spined, evenly spined, spination
becoming vestigial on larger
spicules, fusiform points, subtylote bases.
Microscleres (Table 21). Chelae
absent.
Toxas short, thick, wingshaped, with wide central curvature, curved at slight angle,
slightly reflexed arms.
REMARKS. Hentschel's (1912)
toxifera has seniority over
Topsent's (1913a) name, and
hence the next available name antarctica (Topsent, 1917) is used for
this species (Hooper & Wiedenmayer, 1994).
Koltun (1976) proposed that M.
basispinosa Burton was conspecific with S. toxiferum Topsent,
and this is now confirmed.
Koltun's (1976) proposed
synonymy of this species and S.
tuberculata Burton has not yet
been corroborated (types not yet
found). His proposal to include C.
(Clathria) pauper Brondsted,
1927, in this taxon is rejected, the
latter species having a plumoreticulate skeleton (as opposed to
exclusively plumose skeleton),
FIG. 97. Clathria (Microciona) antarctica (Topsent) (fragment of different pattern of spination on
holotype BMNH1933.3.17.39a). A, Echinating acanthostyles and toxa. acanthostyles, two toxa morB, Plumose skeletal structure.
phologies (C. (M.) antarctica
MEMOIRS OF THE QUEENSLAND MUSEUM
210^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
having only one), and spicule sizes differing substantially between the two species (compare
Table 21 and description of C. (C)pauper above).
Clathria (Microciona) grisea (Hentschel,
TABLE 22. Comparison between present and published records of Clathria (Microciona) grisea
(Hentschel). All measurements are given in gm,
denoted as range (and mean) of spicule length x
spicule width (N=25).
1911) (Figs 98-99, Table 22)
Holotype
(ZMB4435)
Specimen (N=1)
Choanosomal
principal styles
135-(198.2)-212 x 6(10.4)-12
241-(268.5)-298 x
11-(13.2)-16
Subectosomal
auxiliary styles
195-(218.9)-242 x 4- 246-(264.4)-283 x 4(5.8)-8
(6.3)-8
954103.4)-116 x 6- 109-(124.6)-158 x 8(9.4)-11
(7.8)-10
SPICULE
Leptosia grisea Hentschel, 1911: 353, text-fig.35.
Microcionagrissa [lapsus]; de Laubenfels, 1936a: 111.
Clathria grisea; Hooper & Wiedenmayer, 1994: 266
MATERIAL. HOLOTYPE: ZMB4435: NW. of Middle Bluff, Shark Bay, WA, 2548'S, 11326'E, 7-8m
depth, 21.ix.1905, coll. W. Michaelsen & R. Hartmeyer
(dredge). OTHER MATERIAL: WA- NTMZ2863
(fragments QMG300054, PIBOC 04-295).
HABITAT DISTRIBUTION. 7-25m depth; growing
on bivalves and Acropora cf. robusta; Shark Bay and
Pelsart Is, Houtman Abrolhos (WA) (Fig. 98F).
DESCRIPTION. Shape. Thinly encrusting, up to
3mm thick (holotype) or long cylindrical digitate
sponge, 480mm long, 70mm maximum width,
with few, slightly flattened, bifurcate, cylindrical
branches, up to 40mm diameter, and short basal,
holdfast attachment.
Colour. Red-brown alive (Munsell lOR 4/10),
brownish-grey preserved.
Oscules. Large oscules, up to 4mm diameter,
irregularly distributed on lateral sides of branches
in ramose material.
Texture and sutface characteristics. Surface
smooth, unornamented, with distinct skin-like
detachable covering; texture compressible, rubbery.
Ectosome and subectosome. Ectosome heavily
collagenous, up to 1801.1,m thick, including a light
crust of arcuate isochelae, and with tangential
fibres running longitudinally along surface; subectosomal skeleton consists of plumose brushes
of auxiliary subtylostyles, in bundles, standing
erect or semi-erect but not protruding beyond
surface; subectosomal spicule bundles arise from
ends of principal spicules erect on the substrate
(in holotype) or from peripheral fibres (in
specimen).
Choanosome. Holotype — choanosomal skeleton
hymedesmoid, with a basal layer of spongin fibre,
acanthose bases of principal styles and smaller
acanthostyles embedded in basal spongin, standing erect upon substrate. Specimen —
choanosomal skeleton irregularly plumose,
slightly reticulate, with sinuous, heavy spongin
fibres, up to 220..m diameter, cored by both
subectosomal auxiliary subtylostyles and principal styles, and echinated by plumose brushes of
both choanosomal principal styles and echinating
Echinating
acanthostyles
Chelae I
15-(16.8)-19
14-(15.2)-17
Chelae II
20-(22.9)-28
23-(25.5)-28
absent
absent
Toxas
acanthostyles poking through fibres into
mesohyl; mesohyl matrix light, choanocyte
chambers elongate-oval, up to 150p,m diameter,
with abundant arcuate isochelae dispersed
throughout.
Megascleres (Table 22). Choanosomal principal
styles thick, long, slightly curved at centre, subtylote, with heavily spined bases and sparsely
microspined shafts.
Subectosomal auxiliary subtylostyles, in dermal skeleton within choanosomal fibres, long,
thick, straight, with fusiform points and very
slightly subtylote, smooth bases.
Echinating acanthostyles long or short, relatively slender, straight, slightly subtylote, heavily
spined all over spicule except for aspinose point.
Microscleres (Table 22). Arcuate-like isochelae
divided into two size classes, without intermediates; larger chelae with very thick, strongly
curved shaft, small rounded lateral alae attached
to shaft for most of its length, front ala completely
free; smaller chelae with slightly curved shaft,
long lateral alae only partially attached to shaft.
Toxas absent.
REMARKS. There are some notable differences
in skeletal structure and spicule sizes between the
encrusting holotype and the branching specimen
described above (Table 22), but the two
specimens agree so closely in spicule diversity
and geometry that they are obviously conspecific. These differences may be due to the
holotype being immature, having smaller spicule
dimensions and a hymedesmoid skeleton,
whereas the larger branching specimen still
retains the ascending plumose (non-anastomosing) fibre nodes, typical of the Microciona condition.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
211
The major distinguishing features in C. (M.)
grisea are the arcuate isochelae and spined principal spicules. On this basis it is surprising that de
Laubenfels (1936a) did not refer it to Anaata
which he created specifically for this purpose
(i.e., to include species with acanthose principal
spicules, echinating acanthostyles, smooth subectosomal styles and arcuate isochelae). The
holotype of Leptosia grisea has a hymedesmoid
choanosome, without plumose fibre nodes (and
therefore strictly a member of Leptoclathria), and
only a single category of auxiliary spicule.
Anaata, Leptoclathria and Microciona are considered synonyms of Clathria.
Clathria (M.) grisea belongs to Hallmann's
(1912) spicata group of species (see comments
for C. (Thalysias) lendenfeldi), having a spicate
arrangement of principal and echinating spicules,
which protrude through fibres in a plumose manner (Hooper et al., 1990).
Clathria (Microciona) illawarrae sp. nov.
(Figs 100-101, Plate 3D)
MATERIAL. HOLOTYPE: QMG304572: Shellharbour, Illawarra, NSW, 34 ° 35'S, 150°52'E, 10.vi.1993,
coll. L. Miller (SCUBA).
HABITAT DISTRIBUTION. Shallow subtidal; on
rock reef, growing over bivalves and coralline algae;
central E coast (NSW) (Fig. 100G).
DESCRIPTION. Shape. Thinly encrusting, 0.31.5mm thick, following contours of substrate.
Colour. Pale yellowish-orange alive (Munsell
2.5Y 8/10), beige in ethanol.
Oscules. Minute, less than 2mm diameter, scattered over surface, with slightly raised surrounding membraneous lip; pores very small covering
entire surface.
Texture and surface characteristics. Soft, compressible, easily torn; porous, opaque, even,
fleshy surface, without any sculpturing or other
ornamentation.
Ectosome and subectosome. Single category or
large subectosomal auxiliary subtylostyles form
paratangential plumose brushes, protruding only
slightly through surface but extending well into
mesohyl.
Choanosome. Skeleton microcionid, with thin
basal layer of spongin lying on substrate, 3040p..m thick, containing incorporated sand grains;
erect spongin fibre nodes arise at approximately
200Rm intervals along basal spongin, 20-4011m
thick, 100-150p,m long, cored by erect
choanosomal principal styles in uni- or multispicular tracts, up to 5 spicules per bundle, form-
ing perfectly erect or slightly plumose brushes
ascending to but not protruding through surface;
fibre nodes discrete, not anastomosing with adjacent nodes, but some principal spicules from
adjacent nodes cross within mesohyl; paratangential plumose brushes of auxiliary spicules located in several places within mesohyl, forming
a tangential tract near basal spongin layer, forming stellate brushes midway along erect fibre
nodes, and forming plumose paratangential
brushes near surface; echinating acanthostyles
relatively sparse on both basal spongin and erect
fibre nodes; mesohyl matrix moderately heavy
with microscleres dispersed throughout;
choanocyte chambers not seen.
Megascleres. Choanosomal principal styles long,
thin, slightly curved or whispy near point, often
bent in distal third of spicule, with smooth tapering hastate bases and fiisiform points, occasionally slightly telescoped. Length 62-(129.4)-165Rm,
width 3-(3.6)-4.5p,m.
Subectosomal auxiliary subtylostyles long,
thin, straight, with elongated smooth subtylote
bases and hastate points. Length 1764206.5)228p,m, width 1.0-(2.3)-3.0Rm.
Echinating acanthostyles short, relatively
thick, cylindrical, usually thickest above basal
constriction, slightly spined, aspinose slightly
constricted neck, slightly swollen base, rounded
or fusiform point. Length 36-(53.2)-68p,m, width
2-(3.8)-6Rm.
Microscleres. Palmate isochelae very small, with
greatly reduced lateral alae, often no more than
ridge on shaft, and small front ala complete;
sometimes asymmetrical ends. Length 445.9)Toxas small, thick, u-shaped or forceps shaped,
with angular central curve and non-reflexed
arms. Length 6-(8.1)-11m, width 0.540.8)1.01/m.
ETYMOLOGY. For the type locality.
REMARKS. This species is one of the most thinly
encrusting microcionids, with most sections no
more than 300jLm thick. Its choanosomal
skeleton is typical of Microciona (erect fibre
nodes arising from a hymedesmoid basal
skeleton), but subectosomal auxiliary spicules
have an unusual distribution within the skeleton
forming both stellate brushes around the fibre
nodes and basal tangential tracts near the substrate. The species is also unusual amongst thinly
encrusting microcionids in that there are no subsurface drainage canals associated with the
aquiferous system, whereasthe surface is smooth,
porous and fleshy. These live surface features, the
MEMOIRS OF THE QUEENSLAND MUSEUM
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^
FIG. 98. Clathria (Microciona) grisea (Hentschel) (NTMZ2863). A, Choanosomal principal acanthostyles. B,
Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, Arcuate isochelae. E, Section through
peripheral skeleton. F, Australian distribution. G, NTMZ2863. H, Section through fragment of holotype
ZMB4435.
skeletal structure, spicule geometry, and spicule
sizes differentiate this species from other
Microciona in the Indo-west Pacific.
Clathria (Mierociona) lizardensis sp. nov.
(Figs 102-103, Plate 3E)
MATERIAL. HOLOTYPE - QMG304121: Blue
Lagoon, Lizard I., Cairns Section, Great Barrier Reef,
14°41.0'S, 145 °27.5'E, 9m depth, 03.iv.1994, coll.
J.N.A. Hooper, L.J. Hobbs, J.A. Kennedy & S.D. Cook
(SCUBA).
HABITAT DISTRIBUTION. Coral reef, patch reef in
lagoon, under coral overhangs, on ledges or exposed
coral heads, growing on live coral, coral rubble at base
of reef or on dead bivalves; 9-12m depth; Lizard I.
(FNQ) (Fig. 102G).
DESCRIPTION. Shape. Thinly or thickly
encrusting, bulbous in life, usually following
contours of substrate, up to about 101nm thick,
collapsing and less than 4mm thick when
preserved.
Colour. Pale red alive (Munsell 2.5R 6-5/10),
light brown in ethanol.
Oscules. Large, up to 4mm diameter alive, surrounded by raised membraneous lip, usually
situated on apex of bulbous (flaccid) surface, with
drainage canals radiating towards pores; oscules
and drainage canals not visible in preserved
material.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
213
FIG. 99. Clathria (Microciona) grisea (Hentschel) (NTMZ2863). A, Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal principal acanthostyle. D, Spination on principal acanthostyle. E, Echinating
acanthostyle. F, F,chinating acanthostyle spines. G-H, Smaller and larger arcuate isochelae.
Texture and surface characteristics. Soft, slimy,
easily peeled from substrate; smooth, fleshy flaccid surface in life, with slightly sculptured subectosomal drainage canals visible in live sponge; in
preserved material surface uneven, regularly
papillose.
Ectosome and subectosome. Membraneous, collagenous, rarely intact in histological sections,
with some detritus; tips of choanosomal principal
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
A
FIG. 100. Clathria (Microciona) illawarrae sp.nov. (holotype QMG304572). A, Choanosomal principal styles.
B, Subectosomal auxiliary subtylostyles. C, Echinating acanthostyles. D, U-shaped toxas. E, Palmate isochelae.
F, Section through peripheral skeleton. G, Australian distribution.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
215
FIG. 101. Clathria (Microciona) illawarrae sp.nov. (holotype QMG304572). A, Hymedesmoid basal skeleton.
B, Erect spongin fibre. C, Echinating acanthostyles. D, Acanthostyle spines. E, Reduced palmate isochelae. F,
U-shaped toxa.
216zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
CD
Clat hr ia ( Micr ociona) lizar densis sp.nov. (holotype QMG304121). A, Choanosomal principal style/
FIG. 102. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
subtylostyles. B, Subectosomal auxiliary subtylostyles. C, Accolada-U-shaped toxas. D, Echinating acanthostyles. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian distribution.
styles arising from fibre endings, and subectosomal auxiliary subtylostyles protrude through
surface in preserved material, but probably do not
when alive; no special ectosomal skeleton but
plumose bundles of subectosomal subtylostyles
clustered on intact parts of surface skeleton,
usually lying just below the ectosome.
Choanosome. Microcionid skeletal structure,
with very thick, relatively long spongin fibre
nodes, 450-1900p,m long, up to 420p.m diameter,
arising from hymedesmoid basal spongin fibre,
70-23011in diameter, lying directly on substrate;
fibre nodes discrete, erect, without any anastomoses between adjacent nodes, 300-770p.m
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
217
FIG. 103. Clathria (Microciona) lizardensis sp.nov. (holotype QMG304121). A, Choanosomal skeleton. B,
Ascending fibre node. C, Echinating acanthostyles. D, Acanthostyle spines. E, Palmate isochela. F, Accolada
and u-shaped toxas.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
218zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
apart, unbranched except at apex of each node
which bifurcates 1 or more times; fibres dark
brown, with heavy spongin, cored by multi- or
paucispicular plumose tracts of choanosomal
principal styles, with 1 or more principal styles
protruding from apex of each fibre node; fibres
moderately heavily echinated by acanthostyles
dispersed evenly over each ascending fibre node
and all basal fibres; exterior surface of most fibres
often with small amount of collagen and plumose
bundles of auxiliary spicules lying paratangential
to surface (but this is probably an artifact of
preservation, the mesohyl region collapsing
around the fibres, whereas in life the areas between fibres is likely to contain a more structured
aquiferous system); mesohyl matrix granular,
containing some detritus, abundant auxiliary
spicules, and rare microscleres; choanocyte
chambers difficult to see in preserved material,
small, oval, up to 40v.m diameter.
Megascleres. Choanosomal principal styles long
or short, thick, straight, cylindrical or clubshaped, variable basal terminations from tapering
hastate, evenly rounded or faintly subtylote,
fusiform points. Length 183-(272.3)-345p,m,
width 8-(l2.3)-1611m.
Subectosomal auxiliary subtylostyles long,
very slender, usually straight, rarely curved,
sometimes sinuous, with well developed smooth
subtylote bases, fusiform points. Length 211(306.2)-428p.m, width 2-(3.8)-6p.m.
Echinating acanthostyles relatively long, thick,
prominently subtylote, fusiform pointed, with
more-or-less evenly dispersed very small spines,
but abundant larger spines concentrated only on
base and point, giving appearance of aspinose
shaft. Length 81-(94.3)-112Rm, width 446.6)111.1.m.
Microscleres. Isochelae palmate, unmodified, uncommon, moderately large, with long broad front
ala, reduced lateral alae completely fused to shaft,
front and lateral alae approximately the same
size. Length 16-(22.4)-28p,m.
Toxas uncommon, accolada to u-shaped, long
or short, very thin, with slight central curvature,
straight arms or faintly reflexed arms. Length
22-(85.5)-112Rm, width 1.0-(1.2)-1.511m.
ETYMOLOGY. For the type locality.
REMARKS. This species has typical
`microcionid' skeletal structure, with long, discrete, virtually unbranched spongin fibre nodes
arising from a hymedesmoid basal fibre skeleton.
The species is also remarkable for the thickness
and density of its spongin fibres, which are even
heavier than those found in C. (T) corneolia from
New Caledonia (which was named for this character). The external colouration and bulbous surface processes seen in C. (M.) lizardensis are also
reminiscent of C. (T) corneolia, although spicule
geometry, spicule size and skeletal structure differ substantially between the two species (Hooper
& Levi, 1993a), and they do not appear to be
otherwise closely related. In its live external appearance this species could also be mistaken for
C. (M.) aceratoobtusa, but that species has entirely smooth echinating styles, curved principal
styles, oxhorn toxas and abundant isochelae.
Comparisons with other Indo-west microcionids
are discussed in the remarks for C. (M.)
aceratoobtusa.
OTHER SPECIES OF CLATHR1A
(MICROCIONA)
Clathria (Microciona) adioristica (de Laubenfels, 1953)
Dictyociona adioristica zyxwvutsrqponmlkjihgfedcbaZYXWVU
de Laubenfels, 1953a: 526-528, textfig.5 [ Gulf of Mexico] ; Wells et al.,1960: 217-218, textfigs 21,24 [ North Carolina] .
Clathria (Microciona) adioristica; Van Soest, 1984b: 104,
108-109, table 4 [ affinity with Clathria obliqua, possible
synonymy] .
MATERI AL. HOLOTYPE: USNM23403, paratype MLUMML4-214. NW. Atlantic, Caribbean.
Clathria (Microciona) affinis (Carter, 1880)
Microciona affinis Carter, 1880a: 41, 151, 153, p1.4,fig.15
[ Gulf of Manaar, Ceylon] ; Carter, 1881a: 368,384; Carter,
1882b: 111; Ridley & Dendy, 1887: 110; Dendy, 1889a:
38; Vosmaer, 1935a: 608; Burton, 1959a: 247 [ S. Arabian
coast, Zanzibar] .
Not Microciona affinis; de Laubenfels, 1936a: 111.
Not Hymeraphia affinis Topsent, 1889: 43, fig.8.
MATERI AL. HOLOTYPE: LFM dest royed, fragment
BMNH1936.3.4.597. Gulf of Manaar, Arabian Gulf.
Clathria (Microciona) africana (Levi, 1956)
Microciona africana Levi, 1956b: 402-403, text-fig.8 [ Dakar,
Sengal] .
MATERI AL. HOLOTYPE: MNHNDCL1276. NW. Africa.
Clathria (Microciona) angularis (Sara &
Siribelli, 1960)
Microciona angularis Sara & Siribelli, 1960: 69-71, fig.18
[ Bay of Naples] ; Siribelli, 1960: 12, fig.5C [ Naples] ; Sara
& Siribelli, 1962: 47 [ Gulf of Naples] ; Pulitzer-Finali,
1983: 610.
MATERI AL. HOLOTYPE: I MZUN100.4. Mediterranean.
Clathria (Microciona) anonyma (Burton,
1959)
Microciona anonyma Burton, 1959a: 250-251, fig.30 [ Zanzibar area, I ndian Ocean]
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
219
MATERIAL. HOLOTYPE: BMNHI936. 3.4.575. Central E
Africa.
MATERIAL. HOLOTYPE: MOM, fragment MNHNDT125.
Mediterranean. Possible synonym of Clat hria spinarcus
(Carter & Hope) (Maldonado, 1992: 1152).
Clathria (Microciona) armata (Bowerbank,
1862)zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathria (Microciona) atrasanguinea (Bower-
Microcionaarm at a Bowerbank, 1862a: 779, 1858, p1.24, figs
26-28; Bowerbank, 1864:41, p1.4, figs 96-98; Bowerbank,
1866: 124, 129-131, 141 [Britain]; Gray, 1867: 535; Norman, 1869:330; Schmidt, 1870:76; Bowerbank, 1874: 60,
p1.23, figs 17-24; Carter, 1874b: 405, p1.21, fig.27; Carter,
1874c: 456-457; Carter, 1876: 310; Carter, 1880a: 40-41,
151 [Gulf of Manaar]; Koehler, 1886a: 62 [English Channel]; Topsent, 1888: 117, 124, 125, 141, 156, 158, p1.6,
fig.8; Topsent, 1891a: 528 [Roscoff]; Stephens, 1916:234
[W coast, Ireland]; Stephens, 1921; Vosmaer, 1935a: 6-7,
666; Alander, 1942: 62 [Sweden]; Levi, 1956b: 399-400,
text-fig.6 [Dakar, Senegal]; Koltun, 1959: 181-182, textfig. 141 [White Sea]; Levi, 1960a: 73, text-figs 16, 17
[Roscoff,Sidmouth]; Poggiano, 1965: 3, 7, 11-14, textfig.5, tables 1, 2 [Italy]; Pulitzer-Finali, 1983: 573-574,
610, text-fig. 69 [Mediterranean]; Wintermann-Kilian &
Kilian, 1984: 134 [Colombia]; Ackers, Moss & Picton,
1992: 143 [Ireland].
Microciona arm at us; Bowerbank, 1882: 7, 18, 53; Sollas,
1884: 614; Vosmaer, 1884a: 121; Vosmaer, I885b: 353;
Koehler, 1886a: 11, 55; Carter, 1889a: 287; Carter & Hope,
1889: 99, 101-106; Dendy, 1889c: 17; Hope, 1889: 333,
336, 337; Chatin, 1890: 889; Topsent, 1890c: 202, 204;
Topsent, 1892a: 17; Topsent, 1893d: 445; Norman, 1892:
6, 11; Hanitsch, 1894: 176; Topsent, 1894a: 8,23; Topsent
1899: 105; Topsent, 1900: 255; Topsent, 1904a: 189;
Loisel, 1898:38; Minchin, 1898:529; Minchin, 1909:215;
Stephens, 1912: 27; Ferrer Hernandez, 1914: 41.
Clat hria arm at a; Topsent, 1925:649 [discussion]; Van Soest
& Stone, 1986: 45 [Norway].
Scopalina arm at a; Wright, 1868: 224.
Am philect us arm at us; Vosmaer, 1880: 118-119; Vosmaer,
1889: 353; Svarcevskij, 1906: 342; Babic, 1921: 87-88
[Adriatic]; Babic, 1922: 261-262, text-fig.6.
Esperia arm at a Fristedt, 1885: 36-38.
Not Microciona arm at us; de Laubenfels, 1936a: Ill.
Microciona svarchevskyi de Laubenfels, 1936a: 111; Levi,
1960a: 73.
cf. Microciona prolifera; Vosmaer, 1935a: 607.
MATERIAL. HOLOTYPE: BMNH1910.1.1.66
(1930.7.3.209). Caribbean, NE Atlantic, Gulf of Manaar, NW
Africa, Mediterranean. Many of these records are suspect
given their disjunct distribution and this taxon is likely to
consist of a species complex.
Clathria (Microciona) ascendens
(Cabioch, 1968)
Microcionaascendens Cabioch, 1968a: 239, text-fig.11 [Ros-
bank, 1862)
Microciona at rasanguinea Bowerbank, 1862c: 824, 1109,
1110, 1135, p1.30, fig. 1, p1.74, fig.2 [British Seas]; Bowerbank, 1864: 188, 286, p1.33, fig.368, p1.34, fig.369; Bowerbank, 1866: 7, 124, 138-141 [Britain]; Bowerbank, 1874:
63, p1.24, figs 14-19; Topsent, 1888: 141, 157; Topsent,
1890c: 202,204; Topsent, 1891a: 528 [Roscoff]; Topsent,
1892c: 17 [Banyuls]; Dendy, 1922: 60, p1.13, fig. la-e
[Egmont Reef]; Burton, 1934b: 37, text-fig.10; Burton,
1938a: 30, p1.4, fig.24 [Madras]; Lilly et al., 1953 [recordLough Inc, Ireland]; Levi, 1960a: 72-73, text-fig.I5
[English Channel, Atlantic]; Sara & Siribelli, 1962: 47
[Gulf of Naples]; Levi, 1965: 18-19, text-fig.21 [Red Sea];
Simpson, I968a: 33, text-fig.! [Plymouth,England];
Juniper & Steele, 1969: 161 [Portsmouth, England]; Van
Soest & Weinberg, 1980: 10 [Lough Inc. Ireland]; BouryEsnault, 1971: 326 [Banyuls]; Ackers, Moss & Picton,
1992: 142 [Ireland].
Microciona at rosanguinea; Gray, 1867: 535; Norman, 1869:
330; Schmidt, 1870: 76; Carter, I870a: 332, 339, 340;
Carter, 1871a: 272, 274; Carter, 1871b: 8; Carter, 1872a:
106, 111, p1.10, figs 17-20; Carter, 1874c: 457; Carter,
1875: 195; Carter, 1876: 308; Carter, 1880a: 38-41, 151
[Gulf of Manaar]; Carter, 1880b: 59 [Indian Ocean];
Carter, 1881a: 384 [record]; Bowerbank, 1882: 7, 18, 54;
Koehler, 1885: 53, 55; Vosmaer, 1885b: 209; Koehler,
1886a: 61, 62 [English Channel]; Carter, 1887b: 355;
Carter & Hope, 1889: 102, 104-106; Dendy, 1889c: 18;
Hope, 1889: 334; Topsent, 1889: 39; Topsent, 1890c: 202;
Topsent, 1891d: 232; Topsent, 1893d: 445; Topsent,
1894a: 8, 10, 23; Norman, 1892: 6; Hanitsch, 1894: 176;
Heider, 1895: 280; Loisel, 1898: 38; Topsent, 1900: 255
[note]; Woodland, 1908: 140, 145; Minchin, 1909: 217;
Burton & Rao, 1932: 344-345; [coasts of Bengal, Burma,
India and Arabian Sea]; de Laubenfels, 1936b: 448-449
[Panama]; Pulitzer-Finali, 1983: 610; Rodriguez
SolOrzano et al., 1991: 177 [Galicia, Spain].
Microciona at rasanguineum ; Cuenot, 1903: 4 [Arcachon].
Clat hria ( Microciona) at rasanguinea; Van Soest, 1993: 103
[Mauritius].
Am philect us at rasanguineus; Vosmaer, 1880: 115.
Plum ohalichondriaat rasanguinea; Hanitsch, 1890: 207-208,
210 [England].
Scopalina lophyropoda; Schmidt, 1868: 26, 40.
Scopalina at rosangut hea; Schmidt, 1866a: 149; Schmidt,
1866b: 15.
cf. Microciona prohfera; Vosmaer, 1935a: 604, 607.
coff, France]; Rodriguez SolOrzano et al., 1979: 44, 59-60,
text-fig.15 [Galicia, Spain].
MATERIAL. HOLOTYPE: RMBS. NE Atlantic.
MATERIAL. HOLOTYPE: BMNH1930.7. 3.225, paratypes
BMNH1930.7.3.226, 1910.1.1.68. Caribbean, NE Atlantic,
Mediterranean, Red Sea, Arabian Gulf, W Indian Ocean, W
India, Gulf of Manaar, Bay of Bengal, Andaman Sea.
Clathria (Microciona) assimilis Topsent, 1925
Clathria (1Vlicrociona) basifixa
Clat hria assim ilis Topsent, 1925: 649; Topsent & Olivier,
(Topsent, 1913)
1943: 1 [no diagnosis; Adriatic].
Pseudanchinoe assim ilis; de Laubenfels, 1936a: 109 [note].
Microcionaassim ilis; Levi, 1960a: 76 [Adriatic, Naples, Marseilles]; Siribelli, 1960: 18, text-fig.7C [Naples]; Poggiano, 1965: 3, table 1; Vacelet, 1969: 207, text-fig.46
[Mediterranean]; Pulitzer-Finali, 1977: 61 [Bay of
Naples]; Pulitzer-Finali, 1983: 610; Pansini, 1987: 170
[Alboran Sea].
cf. Clat hria com pressa; Topsent, 1925: 649.
Ophlit aspongia basifixa Topsent, 1913b: 39 [Norway]; Bur-
ton, 1935c: 74 [Japan; probable misidentification]; de
Laubenfels, 1954: 162 [note].
Clat hria ( Microciona) basifixa; Van Soest & Stone, 1986:45
[Norway].
MATERIAL. HOLOTYPE: MOM, fragment
MNHNDT1957. NE Atlantic.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
220zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Hymantho bitoxa Burton, 1930a: 503, text-fig.2 [Norway];
Alander, 1942: 63 [Sweden].
Microciona levis; Fristedt, 1887: 416.
Clathria bitoxa; Van Soest, 1984b: 90 [generic synonymy];
Van Soest & Stone, 1986: 47 [note].
Microciona bitoxa Rodriguez SolOrzano & Rodriguez Babio,
1993: 62 [Iberian Peninsula].
MATERIAL. HOLOTYPE: BMNH1910.1. 1.787.v. NE Atlantic.
Microciona affinis; de Laubenfels, 1936a: 111.
Clathria (Microciona)affinis; Van Soest, 1984b: 93-95, 108,
text-fig.36, table 4 [Curacao, West Indies].
Clathria cf. affinis; Kobluk & Van Soest, 1989: 1216
[Bonaire].
Not Microciona affinis Carter, 1880a: 41, p1.14, fig. 15; Vosmaer, 1933: 608.
MATERIAL. HOLOTYPE: MNHNDT1841, paratype
MNHNDT3584. Caribbean, NE Atlantic. Clathria
(Microciona) affinis (Carter, I880a) has priority.
Clathria (Microciona) brepha
(de Laubenfels, 1930)
Clathria (Microciona) carnosa (Bowerbank,
1862)
Clathria (Microciona) bitoxa (Burton, 1930)
Aaata brepha de Laubenfels, 1930: 27 [California]
Anaata brepha; de Laubenfels, 1936a: 91, text-fig.53.
Clathria brepha; Van Soest, 1984b: 7 [generic synonymy].
MATERIAL. HOLOTYPE: USNM2I427. PARATYPES
BMNH1929.8.22.36, 57. NE Pacific.
Clathria (Microciona) brondstedi sp. nov.
Hymedesmia pennata Brondsted, 1932: 12 [Faeroe Is].
Anaata pennata; de Laubenfels, 1936a: 109.
Clathria pennata; Van Soest,1984b: 7 [generic synonymy for
Anaata].
Not Desmacella pennata Lambe, 1895: 129.
MATERIAL. HOLOTYPE: UZM (not located). NE Atlantic.
Clathria ( Microciona) pennata (Lambe, 1895) has seniority.
Clathria (Microciona) bulboretorta (Carter,
1880)
Microcionabulboretorta Carter, I880a: 41, 42, 151, 153, p1.4,
fig.3a-e [Gulf of Manaar, Ceylon]; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: LFM destroyed. Gulf of Manaar.
Clathria (Microciona) bulbotoxa Van Soest,
1984
Clathria (Microciona)bulbotoxa Van Soest, 1984b: 103-104,
p1.7, figs 5-8, text-fig.41, table 4 [Curacao, West Indies].
Microciona bulbotoxa; Pulitzer-Finali, 1986: 149-150 [West
Indies].
MATERIAL. HOLOTYPE: ZMAPOR4789. Caribbean.
Clathria (Microciona) calla (de Laubenfels,
1934)
Axociella calla de Laubenfels, 1934: 16 [Puerto Rico].
Avocielita calla; de Laubenfels, 1954: 149 [note]; Sim &
Byeon, 1989: 40, p1.5, figs 3-5 [Korea; probable misidentification].
Clathria calla; Boury-Esnault, 1973: 286, text-fig.46
[Brazilian Basin]; Zea, 1987: 170, text-fig.59, p1.2, fig. 3
[Colombian Caribbean].
Clathria(Microciona)calla; Van Soest,1984b: 100-101, p1.7,
fig.1, text-fig.39, table 4 [Curacao, Florida; affinity with
Clathria coralloides from Mediterranean].
Microciona calla; Pulitzer-Finali, 1986: 150 [West Indies].
Microciona rarispinosa Hechtel, 1965: 42-44, text-fig.8 [Port
Royal, Jamaica]; Wintermann-Kilian & Kilian, 1984: 135
[Colombia].
Tenaciella obliqua; Alcolado, 1976:5; Alcolado, 1980: 10.
MATERIAL. HOLOTYPE: USNM. Caribbean, tropical SW
Atlantic.
Clathria (Microciona) campecheae nom. nov.
II ymeraphia affinis Topsent, 1889: 43, fig.8A [Banc de
Campeche]; Topsent, 1904a: 162-3 [Azores].
Microciona camosa Bowerbank, I 862a: 804, 1110 [Britain];
Bowerbank, 1866: 133; Vosmaer, 1935a: 607
? Halichondria incrustans; Schmidt, 1866a: 150
MATERIAL. HOLOTYPE: BMNH1930.7. 3.203, fragment
BMNH1910.1.1.666. NE Atlantic, tropical SW. Atlantic.
Clathria (Microciona) claudei sp. nov.
Microciona acanthotoxa Levi & Levi, 1989: 81, fig.49
[Philippines].
MATERIAL. HOLOTYPE: MNHNDCL341 I. Philippines.
Clathria acanthotoxa (Stephens) has seniority.
Clathria (Microciona) cleistochela Topsent,
1925
Clathria cleistochela Topsent, 1925: 650-651, fig.9 [Gulf of
Naples].
Microciona cleistochela; de Laubenfels, 1936a: 1 1 1 [note];
Levi, 1960a: 72, fig. I 4 [ Naples,Banyuls]; Siribelli, 1960:
12-14, fig.5B [Naples]; de Laubenfels, 1951b: 214 [Black
Sea]; Pulitzer-Finali, 1983: 610; Boury-Esnault & Lopes,
1985: 193-194, fig.42 [Azores].
MATERIAL. HOLOTYPE: MOM, fragment MNHNDT329.
Mediterranean, NE Atlantic.
Clathria (Microciona) coccinea (Bergquist,
1961)
Microciona coccinea Bergquist, 1961a: 38, fig.8a,b [N. New
Zealand]; Bergquist & Sinclair, 1968: 427, 428, fig. 1 a
[morphology and larvae]; Bergquist & Sinclair, 1973: 43;
Bergquist et al., 1970: 248, 254; Evans & Bergquist, 1977:
195-196; Bergquist & Fromont, 1988: 102-103, p1.47,
fig.f, p1.48, fig.a; Rudman & Avern, 1989: 335; Dawson,
1993: 36 [note].
Not Thalisias coccinea Duchassaing & Michelotti, 1864: 84,
p1.18, fig.5 [St.Thomas]; Wiedenmayer, 1977a: 253, table
49.
MATERIAL. HOLOTYPE: NMNZ unregistered. New
Zealand.
Clathria (Microciona) ctenichela (Alander,
1942)
M wrociona ctenichela Alander, 1942: 61-62, p1.15, fig.20
[Sweden].
Clathria (Microciotia) ctenichela; Van Soest & Stone, 1986:
44-45 [Norway].
MATERIAL. HOLOTYPE: ZMA. NE Atlantic.
Clathria (Microciona) dendyi (Bergquist &
Fromont, 1988)
Microciona dendyi Bergquist & Fromont, 1988: 100-102,
p1.47, figs d,e [Slipper 1.]; Dawson, 1993: 37 [note].
MATERIAL. HOLOTYPE: NMNZPOR114. New Zealand.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
221
Clathria (Microciona) densa (Burton, 1959)
Microciona densa Burton, 1959a: 248, text-fig.28 [S Arabian
coast]
MATERIAL. HOLOTYPE: BMNH1936.3. 4.456. Arabian
Gulf.
Clathria (Microciona) dianae (Schmidt, 1875)
Suberites dianae Schmidt, 1875: 116, p1.1,fig.1 [Norway];
Czerniawsky, 1880: 70.
Microciona dianae; Thiele, 1903b: 394, 395, 398, p1.21,
fig.28a-e; Vosmaer, 1935a: 608, 630; de Laubenfels,
1936a: 1 1 1 [note].
Not A rtemisina dianae Topsent, 1907: 69; Vosmaer, 1935a:
630.
Clathria dianae; Van Soest & Stone,1986: 47 [note].
MATERIAL. HOLOTYPE: unknown. NE Atlantic.
Clathria (Microciona) ditoxa (Stephens, 1916)
Eurypon ditoxa Stephens, 1916:239-240 [W coast of Ireland];
Stephens, 1921: 51, p1.5; Burton, 1959b: 44-45 [Iceland].
Microciona ditoxa; Levi, 1960a: 66 [W coast of Ireland,
Atlantic].
Dictyociona ditoxa; de Laubenfels, 1936a: 110 [note].
MATERIAL. HOLOTYPE: INMSR151.9.1916. NE Atlantic.
Clathria (Microciona) duplex Sara, 1958
Clathria duplex Sara, 1958: 262-264, fig.24 [Gulf of Naples].
Microciona duplex; Levi, 1960a: 69 [Naples]; Siribelli, 1960:
14-15, fig.6A [Naples]; Sara, 1963: 210 [Gulf of
Policastro]; Pulitzer-Finali, 1983: 610; Rodriguez
SolOrzano et al., 1991: 179, fig.3 [Galicia, Spain].
MATERIAL. • HOLOTYPE: IMZUN93.26.x.58. Mediterranean.
Clathria (Microciona) echinata (Alcolado,
1984)
Axociella echinata Alcolado, 1984: 7 [Cuba].
Clathria echinata; Kobluk & Van Soest, 1989: 1216;
Meesters et al., 1991: 194 [Curacao, Bonaire].
Clathria (Microciona) simpsoni Van Soest, 1984b: 97-99,
p1.7, figs 2-4, text-fig.38, table 4 [Puerto Rico, Curacao];
Pulitzer-Finali, 1986: 150 [West Indies].
Clathria simpsoni; Zea, 1987: 168, text-fig.58, p1.3, fig.1
[Colombian Caribbean].
MATERIAL. HOLOTYPE: Cuba. Holotype of simpsoni:
ZMAPOR3332. Caribbean.
Clathria (Microciona) elliptichela (Alander,
1942)
Microciona elliptichela Alander, 1942: 58-61 [Sweden].
Clathria elliptichela; Van Soest & Stone, 1986:45 [note].
MATERIAL. HOLOTYPE: ZMA. NE Atlantic.
Clathria (Microciona) fallax (Bowerbank,
1866)
Microciona falla.v Bowerbank, 1866: 124, 128, 129, 135
[Hastings, Britain]; Vosmaer, 1935a: 607; Curtis, 1970:
260-261 [cytology]; Ackers, Moss & Picton, 1992: 147
[Ireland].
Sophax fallav; Gray, 1867: 521.
MATERIAL. LECTOTYPE-BMNH 1910.1. 1.71.
PARALECTOTYPE BMNH1930.7.3.198, fragment
USNM5047. NE Atlantic.
Clathria (Microciona) fascispiculifera (Carter,
1880)
Microciona fascispiculifera Carter, 1880a: 44, 45, 151, 153,
p1.4, fig.7a-g [Gulf of Manaar]; Hallmann, 1916c: 637
[note]; [?] Vosmaer, 1935a: 608.
Damoseni fascispiculifera; de Laubenfels, 1936a: 110.
MATERIAL. HOLOTYPE: LFM destroyed, no extant fragment in BMNH. Gulf of Manaar.
Clathria (Nlicrociona) fraudata (Bowerbank,
1874)
Microciona fraudata Bowerbank, 1874: 273, 275, 277, p1.83,
figs 7-11 [Polperro, Fowey Hbr.]; Vosmaer, 1935a: 607.
MATERIAL. HOLOTYPE: BMNH1930.7.3. 205. NE Atlantic.
Clathria (Microciona) frogeti (Vacelet, 1969)
Microciona frogeti Vacelet, 1969: 208, text-fig.47 [Mediterranean].
Microciona fregeti [sic.]; Pulitzer-Finali, 1983: 610 [list].
MATERIAL. HOLOTYPE: MNHN missing. W Mediterranean.
Clathria (Microciona) gradalis Topsent, 1925
Clathria gradalis Topsent, 1925: 651-653 [Gulf of Naples];
Topsent & Olivier, 1943: 1 [Monaco]; Sara, 1958: 258260, text-fig.22 [Gulf of Naples]; Sara, 1960a: 461 [Ischia].
Clathria gradalis var. atoxa; Topsent, 1928a: 299, p1.10,
fig.14 [Boavista I., Senegal].
Microciona gradalis; de Laubenfels, 1936a: 111; Levi, 1960a:
75 [W Mediterranean]; Sara & Siribelli, 1960: 67 [Bay of
Naples]; Siribelli, 1960: 16, text-fig.6B [Naples]; Sara &
Siribelli, 1962:47-48 [Gulf of Naples]; Poggiano, 1965:3,
table 1; Cabioch, 1968a: 244 [Roscoff, N. France];
Vacelet, 1969: 207 [W Mediterranean]; Pulitzer-Finali,
1977: 63 [Bay of Naples]; Pulitzer-Finali, 1983: 610.
MATERIAL. HOLOTYPE: MOM, fragment MNHNDT328.
Mediterranean, NE Atlantic, NW Africa.
Clathria (Microciona) haematodes (de Laubenfels,
1957)
Microciona haematodes de Laubenfels, 1957: 240, text-fig.6
[Oahu, Hawaii]; Bergquist, 1977: 67 [Hawaii].
MATERIAL. HOLOTYPE: USNM23533. Hawaii.
Clathria (Microciona) haplotoxa (Topsent,
1928)
Leptoclathria haplotoxa Topsent, 1928a: 298, p1.10, fig.16
[Madeira I.]; [cf.] Topsent, 1934b: 24.
Microciona haplotoxa;Topsent, 1934a: 92-93 [Gulf of Gabes,
Tunisia]; Levi, 19566: 400-402, text-fig.7 [Dakar,
Senegal]; Levi, 1960a: 70 [Madeira, Tunisia]; PulitzerFinali, 1983: 610.
MATERIAL. HOLOTYPE: MNHNDT1101. Mediterranean,
NE Atlantic, NW Africa.
Clathria (Microciona) hentscheli sp. nov.
Ilymeraphia lendenfeldi Hentschel, 1912: 378-379, p1.20, fig.
35 [Mimien Bay, Am I., Arafura Sea].
Eurypon lendenfeldi; de Laubenfels, 1936a: 110.
Not Clathria lendenfeldi Ridley & Dendy, 1886: 474.
MATERIAL. LECTOTYPE: SMF 1705. Indonesia. Clathria
(Thalysias) lendenfeldi Ridley & Dendy has priority.
222zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Clathria (Microciona) heterotoxa
(Hentschel, 1929)
Microciona heterotoxa Hentschel, 1929: 891-892, 970, p1.14,
fig.5 [White Sea]; Koltun, 1959: 182-183, text-fig.142
[Arctic, USSR].
Dictyociona heterotoxa; de Laubenfels, 1936a: 110 [note]; de
Laubenfels, 1953a: 527.
MATERIAL. HOLOTYPE: HM or ZMB (not located).
Arctic.
Clathria (Microciona) hymedesmioides Van
Soest, 1984
Clathria (Microciona) hymedesmioides Van Soest, 1984b:
104-105, p1.7, figs 9-10, text-fig.42, table 4 [Curacao].
MATERIAL. HOLOTYPE: ZMAPOR4790. Caribbean.
Clathria (Microciona) ixauda (Levi, 1969)
Microciona ixauda Levi, 1969: 965, text-fig.7a [Vema
Seamount].
MATERIAL. HOLOTYPE: MNHNDCL1415. S Atlantic.
Clathria (Microciona) jecusculum (Bowerbank, 1866)
Hymeniacidon jecusculum Bowerbank, 1866: 198 [Harris I.,
Hebrides].
Microciona jecusculum; Bowerbank, 1874: 273-275, p1.83,
figs 1-6; Carter, 1876: 237 [Cape St. Vincent; Faroe Is];
Vosmaer, 1933: 607 [imperfectly known].
MATERIAL. HOLOTYPE: unknown; fragments
BMNH1954.3.9.176, 177. NE Atlantic.
Clathria (Microciona) kentii (Bowerbank,
1874)
Microciona kentii Bowerbank, 1874: 311, 312, 317-319,
p1.89, figs 9-13 [Jersey, Strangford Lough]; Vosmaer,
1935a: 607.
MATERIAL. HOLOTYPE: BMNH1910.1.1.77, fragment
USNM5044. NE Atlantic.
Clathria (Microciona) laevis (Bowerbank,
1866)
Microciona laevis Bowerbank, 1866: 124, 127-128 [Britain];
Stephens, 1917: 12, p1.1, fig.3 [N. of Bolus Head, Ireland];
Vosmaer, 1935a: 607; Burton, 1959b: 43 [Iceland].
Not Microciona laevis; Fristedt, 1887: 415.
Abila laevis; Gray, 1867: 539.
Hymantho laevis; de Laubenfels, 1936a: 111; Alander, 1942:
63 [Sweden].
MATERIAL. HOLOTYPE: BMNH1930.7.3. 215. NE Atlantic.
Clathria (Microciona) laevissima
(Dendy, 1922)
MATERIAL. HOLOTYPE: USNM22827. W central Pacific.
Clathria (Microciona) leighensis sp. nov.
Microciona rubens Bergquist, 1961a: 38, text-fig.9 [NNew
Zealand]; Bergquist & Green, 1977b: 289-302 [ontogeny];
Bergquist & Fromont, 1988: 103, p1.48, figs b-c [N New
Zealand]; Dawson, 1993: 37 [note].
Not Thalassodendron rubens Lendenfeld, 1888: 223.
MATERIAL. HOLOTYPE: NMNZ unregistered. New
Zealand. C. (Clathria) rubens (Lendenfeld, 1888) has
priority.
Clathria (Microciona) levii (Sara & Siribelli,
1960)
Microciona levii Sara & Siribelli, 1960: 71-73, text-fig.19
[Bay of Naples]; Siribelli, 1960: 6-8, text-fig.2 [Naples];
Poggiano, 1965:3, table 1; Pulitzer-Finali, 1983:610 [list].
MATERIAL. HOLOTYPE: IMZUN31. Mediterranean.
Clathria (Microciona) longispiculum
(Carter, 1876)
Microciona longispiculum Carter, 1876: 231, 237, 238, 470,
p1.12, fig. 1 h, p1.15, fig.31a-c [N. Scotland]; Vosmaer,
1935a: 608 [insufficiently known].
MATERIAL. HOLOTYPE: BMNH1887.10. 29.3. NE Atlantic.
Clathria (Microciona) longistyla
(Burton, 1959)
Microciona longistyla Burton, 1959a: 249-250, text-fig.29 [S.
Arabian coast]; Sim & Kim, 1988: 26 [Korea]; Sim &
Byeon, 1989: 40 [Korea; possible misidentifications].
MATERIAL. HOLOTYPE: BMNH1936.3.4. 583. Arabian
Gulf, ? S China Sea.
Clathria (Microciona) macrochela
(Levi, 1960)
Microciona macrochela Levi, 1960a: 70, text-fig.13 [Roscoff].
MATERIAL. HOLOTYPE: MNHNDCL940. NE Atlantic.
Clathria (Microciona) microjoanna
(de Laubenfels, 1930)
Microciona microjoanna de Laubenfels, 1930: 27 [Carmel,
California]; de Laubenfels, 1932: 93-95, text-fig.55
[California]; Bakus, 1966:433-435, text-fig.4, table 4 [San
Juan Arch., Washington State]; Lee & Gilchrist, 1985:
24-32 [biochemistry]; Bakus & Green, 1987: 72 [S.
California].
MATERIAL. HOLOTYPE: USNM21468. PARATYPE
USNM21469, fragment BMNH1929.8.22.28. NE Pacific.
Clathria (Microciona) micronesia
(de Laubenfels, 1954)
Hymedesmia laevissima Dendy, 1922: 81-82, p1.15, fig.1
[Mauritius].
Folitispa laevissima; de Laubenfels, 1936a: 119 [note].
MATERIAL. HOLOTYPE: BMNH1921.11. 7.69. W Indian
Ocean.
Microciona micronesia de Laubenfels, 1954: 145-146, textfig.93 [Majuro Atoll].
MATERIAL. HOLOTYPE: USNM22833. W central Pacific.
Clathria (Microciona) lajorei (de Laubenfels,
1954)
Clathria (Microciona) microxea
(Vacelet & Vasseur, 1971)
Anaata lajorei de Laubenfels, 1954: 147-148, text-fig. 95
[Ailing-lap-lap Atoll].
Clathria lajorei; Van Soest, 1984a: 129 [generic synonymy].
Paratenaciella microxea V acelet & Vasseur, 1971: 103, textfig.61 [Tulear, Madagascar].
MATERIAL. HOLOTYPE: MNHNDIV27. W Indian Ocean.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
223
Clathria (Microciona) mima
(de Laubenfels, 1954)
Ophlitaspongia mima de Laubenfels, 1954: 161-162, textfig.105 [Majuro Atoll, central W. Pacific]; WintermannKilian & Kilian, 1984: 135 [Colombia].
MATERIAL. HOLOTYPE: USNM22839. NW central
Pacific, Caribbean.
Clathria (Microciona) namibiensis (Uriz,
1984)
Microciona namibiensis Uriz, 1984b: 111-113, text-figs 3
A-B, 5D; Uriz, 1988a: 87 [Namibia].
MATERIAL. HOLOTYPE: ABIPB-12. SW Africa.
Clathria (Microciona) normani (Burton, 1930)
Hymantho norrnani Burton, 1930a: 503, text-fig.! [Norway].
Clathria normani; Van Soest, 1984b: 90 [generic synonymy];
Van Soest & Stone, 1986: 46-47 [note].
MATERIAL. HOLOTYPE: BMNH1910.1.1. 791.iii. NE Atlantic.
Clathria (Microciona) novaezealandiae
(Brondsted, 1924)
Microciona novaezealandiae Brondsted, 1924: 463-464, textfig.19 [Slipper Is]; Bergquist & Fromont, 1988: 103-104;
Dawson, 1993: 37.
Wetmoreus novaezealandica; de Laubenfels, 1936a: 112.
MATERIAL. HOLOTYPE: UZM not located, no fragment in
BMNH. New Zealand
Clathria (Microciona) osismica (Cabioch,
1968)
Microciona osismica Cabioch, 1968a: 240-244, text-fig. 12
[Roscoff, France].
MATERIAL. HOLOTYPE: RMBS. NE Atlantic.
Clathria (Microciona) parthena
(de Laubenfels, 1930)
Microciona parthena de Laubenfels, 1930:27 [California]; de
Laubenfels, 1932: 95-96, text-fig.56 [California]; Henkart
et al., 1973: 3045-3050, text-figs 1-5 [biochem.];
Cauldwell et al., 1973: 3051-3058, text-figs 1-5
[biochem.]; Lee & Gilchrist, 1985: 24-32 [biocheml; Sim
Bakus, 1986: 9 [California]; Bakus & Green, 1987:
72-73 [S. California].
MATERIAL. HOLOTYPE: USNM21383. PARATYPE
BMNH1929.9.30.6. NE Pacific.
Clathria (Microciona) pennata (Lambe, 1895)
Desmacella pennata Lambe, 1895: 129, p1.4, figs 6a-d
[Sooke, Vancouver I., Canada]; Ricketts & Calvin, 1948:
34 [Vancouver I., Canada].
Ophlitaspongia pennata; de Laubenfels, 1927: 265-266, textfigs 1-4 [Laguna Beach and Monterey Pen., California]; de
Laubenfels, 1930: 28; Burton, 1930a: 521; de Laubenfels,
1932: 103, text-fig.62 [var. californiana]; Hewatt, 1946:
193 [California]; Hartman (in Light, 1954): 20 [California]; de Laubenfels, 1954: 162 [note]; Burton, 1959a: 247
[key to species]; de Laubenfels, 1961: 198, fig.1 [California and Washington State]; Bakus, 1966: 435-440, textfig.5, tables 5-6 [et var., San Juan Arch., Washington];
Anderson, 1973:5668 [associates]; Lee & Gilchrist, 1985:
24-32 [biochem.]; Sim & Bakus, 1986: 10; Bakus & Green,
1987:73 [var. californiana]; Sim & Byeon, 1989: 37, p1.1,
figs 1-4 [var. californiana; Korea]; Lee & Klontz, 1991:
61 [chemistry].
Tylodesma pennata; Koltun, 1959: 96 text-fig.51, 1-3 [S.
Kuriles].
Biemna pennata; Koltun, 1958: 54.
Ophlaaspongia affinis basifixa; Burton, 1935c: 74 [Possiet
Bay, Sea of Japan].
Microciona pennata; Simpson, 1968a: 40, p1.15 [Washington
State].
Clathria (Ophlitaspongia)pennata; Rudman & Avem, 1989:
335 [associates; probable misidentification of sponge].
Not Pandaros pennata Duchassaing & Michelotti,1864: 88.
Not Hymedesmia pennata Brondsted, 1932: 12 [see C.
brondstedi sp. nov.].
MATERIAL. HOLOTYPE: USNM7488. PARATYPE
NMCIC1900-2826. NE Pacific, NW. Pacific, Japan.
Clathria (Microciona) plana (Carter, 1876)
Microciona planum Carter, 1876: 238, 472 [Cape St. Vincent,
Hebrides].
Microciona plana; Topsent, 1889: 41-42 [Banc de
Campêche]; Norman, 1892: 9.
Amphilectus planus; Vosmaer, 1880: 121; Dendy, 1889: 18.
Clathria planer; Topsent, 1894b: 30; Van Soest, 1984b: 108
[unrecognisable].
cf. Microciona prolifera; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: BMNH1890.4. 10.13. NE Atlantic.
Clathria (Microciona) plinthina (de Laubenfels,
1954)
Microciona plinthina de Laubenfels, 1954: 144-145, textfig.92 [Ailing-lap-lap Atoll].
MATERIAL. HOLOTYPE: USNM22949. W central Pacific.
Clathria (Microciona) poecilosclera (Sara &
Siribelli, 1960)
Microciona poecilosclera Sara & Siribelli, 1960: 73-75, textfig.20 [Bay of Naples]; Siribelli, 1960: 18, text-fig.7B
[Naples]; Poggiano, 1965:3, table 1; Pulitzer-Finali, 1983:
610.
MATERIAL. HOLOTYPE: IMZUN350. Mediterranean.
Clathria (Microciona) polita (Ridley, 1881)
Hymedesmia polita Ridley, 1881: 121-122, p1.10, fig.9
[Sandy Point, Magellan Straits; supposed affinities with
Clathria tuberosa (Bowerbank)].
Microciona polita; Topsent, 1900: 113 [English Channel;
identified with doubt].
MATERIAL. HOLOTYPE: BMNH1879.12.27. 22. SW Atlantic, NE. Atlantic.
Clathria (Microciona) primitiva (Koltun,
1955)
Microciona prirnitiva Koltun, 1955a: 16-17, text-fig.6 [Behring Sea]; Burton, 1959: 43 [Iceland]; Koltun, 1959: 184,
text-fig. 144 [USSR].
Not Clathriella primitiva Burton, 1935c: 73.
MATERIAL. HOLOTYPE: ZIL, fragments
BMNH1932.1.1.241, 572. Boreal region.
Clathria (Microciona) proxima (Lundbeck,
1910)
Hymedesmia proxima Lundbeck, 1910: 81 [Denmark Strait].
Anaata proxima; de Laubenfels, 1936a: 109 [imperfectly
known].
224zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Clathriaproxima; Van Soest, 1984b: 7 [generic synonymy for
Anaata].
MATERIAL. HOLOTYPE: ZRS. NE Atlantic.
Clathria (Microciona) pugio (Lundbeck, 1910)
Hymedesmia pugio Lundbeck, 1910: 94 [Denmark Strait].
Anaata pugio; de Laubenfels, 1936a: 109.
Clathria pugio; Van Soest, 1984b: 7 [generic synonymy for
Anaata].
MATERIAL. HOLOTYPE: ZRS. NE Atlantic.
Clathria (Microciona) pustulosa (Carter, 1882)
Halichondria pustulosa Carter, 1882a: 285, p1.11, fig.1
[vicinity of Pantagonia and Falkland Is].
Not Halichondria pustulosa; Carter, 1886g: 450 [Port Phillip
Bay, Victoria].
Anaata pustulosa; de Laubenfels, 1936a: 109.
MATERIAL. HOLOTYPE: BMNH not found, fragment
BMNH1954.3.9.233. SW Atlantic.
Clathria (Microciona) quadriradiata (Carter,
1880)
Microciona quadriradiata Carter, 1880a: 42, 43, 151, 153,
p1.4, figs 4a-d [Gulf of Manaar]; [?] Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: LFM destroyed. Gulf of Manaar.
Clathria (Microciona) rhopalophora
(Hentschel, 1912)
Hymeraphia rhopalophora Hentschel, 1912: 380, p1.20,
fig.37 [Aru I., Arafura Sea].
Microciona rhopalophora; Burton, 1959a: 248 [Maldives];
Thomas, 1970b: 206, text-fig.7 [Cocos-Keeling Basin,
Gulf of Manaar].
Eurypon rhopalophora; de Laubenfels, 1936a: 111.
MATERIAL. HOLOTYPE: HM (fragment 5MF992). Indonesia.
Clathria (Microciona) scotti Dendy, 1924
Clathria scotti Dendy, 1924a: 352, p1.10, fig.!, p1.14, figs 5-8
[E of North Cape, New Zealand].
Pseudanchinoe scold; de Laubenfels, 1936a: 109 [note];
Bergquist & Fromont, 1988: 110-111, p1.51, figs b-d [N.
New Zealand]; Dawson, 1993: 39 [note].
MATERIAL. HOLOTYPE: BMNH1923.10. 1.128,
paratypes BMNH1923.10.1.129-131, AMZ2568. New
Zealand.
Clathria (Microciona) seriata (Grant, 1826)
Spongia seriata Grant, 1826: 116.
Halichondria seriata; Johnston, 1842:74, 125, 197, 248, 258,
p1.14, fig.2; Gray, 1848: 12,16.
Halic lona seriata; Bowerbank, 1861: 235; Bowerbank,
1862a: 769, 824, p1.29, fig.10.
Chalitza seriata; Bowerbank, 1864: 24,2275, p1.17, fig.287;
Bowerbank, 1866: 139, 294, 361, 376-378; Wright, 1868:
228; Norman, 1869:298-299; Schmidt, 1870: 3, 77; Carter,
1871b: 196.
Clathriaseriata; Schmidt, 1866b: 10, 24, p1.1, fig.7; Vosmaer,
1935a: 619 [uncertain affinity]; Hanitsch, 1889: 158;
Hanitsch, 1890: 205-207 [England]; Babic, 1921: 84
[Adriatic]; Babic, 1922: 244-245, text-fig.T [Adriatic].
Seriatula seriata; Gray, 1867: 515.
Desmacidon seriata; Schmidt, 1868: 12.
? Desmacodes seriatus; Vosmaer, 1880: 107.
Ophlitaspongia seriata; Bowerbank, 1874:6, 167, p1.65, figs
1-4; Carter, 1875: 196; Bowerbank, 1882: 14, 24, 120,
186-188; Carter, 1883b: 314; Vosmaer, 1885: 357; MacMunn, 1888: 12, 14, 20; Dendy, 1889c: 14; Topsent,
1890c: 204; Topsent, 1891a: 529; Norman, 1892: 8;
Minchin, 1900: 20, fig.32; Thiele, 1905: 450-451;
Kirkpatrick, 1907: 274; Kirkpatrick, 1908a: 26; Weltner,
1910a: 23; Hallmann, 1912: 254; Stephens, 1912: 3, 28;
Stephens, 1916: 234 [Ireland]; Ferrer Hernandez, 1914:
43; Lilly et al., 1953 [Ireland]; de Laubenfels, 1954: 161162 [note]; Burton, 1959a: 247 [key to species of Ophlitaspongia]; Levi, 1960a: 64-65, text-fig.9 [Atlantic];
Levi, 1963: 59-60, text-fig. 69, p1.9B-C [Cape Town,
South Africa]; Bergquist & Sinclair, 1968: 427, 428, textfig.1B [larvae, New Zealand]; Bergquist & Hogg, 1969:
207, 210; Fry, 1970: 135-157 [ecology]; Fry, 1971: 155178 [larvae]; Bergquist &Sinclair, 1973:37-39; Fry, 1973:
159-170 [ecology]; Van Soest & Weinberg, 1980: 10
[Lough Ine, Ireland]; Lee & Gilchrist, 1985: 24-32
[biochemistry]; Rudman & Avern, 1989: 335 [associates];
Ackers, Moss & Picton, 1992: 147 [Ireland]; Dawson,
1993: 38 [note].
Echinoclathria seriata;Topsent, I893d: 445; Hanitsch, 1894:
179; Hanitsch, 1895: 212; Heider, 1895: 281; Topsent,
1896: 114; Loisel, 1898: 38; Minchin, 1900: 20;
Whitelegge, 1907: 503.
Microciona seriata; Simpson, 1968a: 37, pls 9-10, text-fig.2
[Plymouth, England].
Not Rhaphidophlus seriatus Thiele, 1899: 14, p1.1, fig.6, p1.5,
fig.7 [Celebes].
Ophlitaspongia papilla Bowerbank, 1866: 14, 378-380
[Vazon Bay, Guernsey]; Bowerbank, 1874, p1.70, figs 1-4;
Bowerbank, 1882: 187 [Westport Bay, Guernsey]; Koehler, 1886a: 62 [English Channel]; Hallmann, 1912: 254
[note].
? Clathria papilla; Schmidt, 1870: 77; Vosmaer, 1880: 155.
Echinoclathria papilla; Hanitsch, 1894: 8-10, 16, 25, 26.
Ophistospongia papilla; Gray, 1867: 515.
MATERIAL. HOLOTYPE: BMNH1847.9.7.14, fragments
BMNH1910.1.1.2368, 2369. NE Atlantic, S Africa, New
Zealand, Mediterranean. This species is undoubtedly composite, consisting of at least two sibling species (Atlantic and
Indo-Pacific populations).
Clathria (Microciona) sigmoidea (Cuartas,
1992)
Microciona sigmoidea Cuartas, 1992: 85-88, figs 53-57, 67
[Mar del Plata, Argentina].
MATERIAL. HOLOTYPE: MCNPC04-81-43. SW Atlantic.
C1athria (Microciona) simae sp. nov.
Axociella cylindrica; Sim & Byeon, 1989: 39-40, p1.5, figs
1-2 [S. Korea].
Not Esperiopsis cylindrica Ridley & Dendy, 1886: 340; Ridley & Dendy, 1887: 79-80, p1.19, figs 2a-b.
Not Axociella cylindrica; Hamann, 1920: 780-783, p1.37,
figs 2-4, text-fig.2.
Not Rhaphidophlus cylindricus Kieschnick, 1900: 53, p1.44,
fig. 10.
MATERIAL. HOLOTYPE: Department of Biology, Han
Nam University, Korea 18/vii/1987. S China Sea. C. (Axociella) cylindrica (Ridley & Dendy, 1886) has priority.
ETYMOLOGY. Named for Dr C.J. Sim.
Clathria (Microciona) similis (Thiele, 1903)
Hymeraphia finials Thiele, 1903a: 957, fig.22 [Ternate,
Moluccas]; Hentschel, 1912: 377 [Am I., Arafura Sea].
Eurypon sinidis; de Laubenfels, 1936a: 111.
Not Microciona similis Stephens, 1915:441.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
225
Not Eurypon similis; Uriz, 1988a: 53-54, text-fig.29.
MATERIAL. HOLOTYPE: ZMB7215. Indonesia.
Clathria (Microciona) spinarcus (Carter &
Hope, 1889)
Microcionaspinarcus Carter & Hope, 1889:99-106, p1.6, figs
1-6 [Hastings, England]; Carter, 18896: 250; Hope, 1889:
339; Topsent, 1890c: 199, 202, 205; Topsent, 1892a: 113
[Banyuls]; Topsent, 1892c: 17; Topsent, 1894a: 8, 11;
Topsent, 1896: 115; Ferrer Hernandez, 1914: 14; Topsent,
1928a: 62; Levi, 1960a: 74, 76, text-fig.18 [Atlantic];
Boury-Esnault, 1971: 324-325 [Banyuls]; Van Soest &
Weinberg, 1980: 6-8, 10, text-fig.8 [Lough Inc, Ireland];
Boury-Esnault & Lopes, 1985: 193, fig.41 [Azores]; Uriz,
1988a: 88-89 [Namibia]; Uriz, 19886: 68 [Namibia]; Maldonado, 1992: 1152 [Aldoboran Sea]; Ackers, Moss &
Picton, 1992: 146 [Ireland].
Microciona armata, in part; Carter, 1874c: 457; Carter &
Hope, 1889: 101-106.
Ligrota spinarcus; de Laubenfels, 1936a: 126 [transferred
with hesitation].
Microciona acanthotoxa; Lilly et al., 1953 (Van Soest &
Weinberg, 1980: 10).
cf. Microciona prolifera; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: BMNH1910.1.1. 501, fragment
BMNH1954.3.9.175. NE Atlantic, W Mediterranean, SW
Africa.
Not Euryponsimilis; Uriz, 1988:53 [see C. urizae, nom. nov.].
MATERIAL. HOLOTYPE: RSME1921.143. 1447, fragments INM.31.1914, BMNH1939. 3.20.11. South Africa.
Clathria (Microciona)similis (Thiele, 1903a) has seniority.
Clathria (Microciona) strepsitoxa (Hope,
1889)
Microciona strepsitoxa Hope, 1889: 334-338, 342, p1.16, figs
1-10; Topsent, 1890c: 199, 205; Topsent, 1891a: 529
[Roscoff]; Topsent, 1892c: 17 [Banyuls]; Topsent, 1894a:
8; Topsent, I934a: 90-92 [Gulf of Gabes]; Alander, 1942:
62 [Sweden]; Levi, 1960a: 67, text-fig.10 [English Channel, Atlantic, Mediterranean]; Siribelli, 1960: 4, textfig. IA [Naples]; Sara & Siribelli, 1960: 67-69, text-fig.17
[Bay of Naples]; Poggiano, 1965: 3, table 1; Boury-Esnault, 1971: 326 [Banyuls]; Pulitzer-Finali, 1977:62 [Bay
of Naples]; Rodriguez SolOrzano et al., 1979: 44, 58-59,
text-fig. 14 [Galacia, Spain]; Van Soest & Weinberg, 1980:
6, text-fig. 7 [Lough Ine, Ireland]; Pulitzer-Finali, 1983:
573, 610 [Mediterranean]; Wintermann-Kilian & Kilian,
1984: 134 [Colombia]; Boury-Esnault & Lopes, 1985:
192-193, fig.40 [Azores]; Ackers, Moss & Picton, 1992:
145 [Ireland].
cf. Microciona prolifera; Vosmaer, 1935a: 608, 640.
MATERIAL. HOLOTYPE: BMNH1910.1.1. 500. NE Atlantic, Mediterranean, Caribbean.
Clathria (Microciona) tenuis (Stephens, 1915)
Clathria (Microciona) spinatoxa (Hoshino,
1981)
Microciona spinatoxa Hoshino, 1981: 155, text-fig.67, p1.7,
fig.! [Sasajima].
MATERIAL. HOLOTYPE: MMBSSIS-033. Japan.
Clathria (Microciona) spinosa (Wilson, 1902)
Microciona spinosa Wilson, 1902: 396-397 [St.Thomas,
Puerto Rico]; Simpson, I968a: 37, pls 9-10, text-fig.2
[Bahamas]; Simpson, 19686: 26, pls 6-8, tables 5-7;
Wiedenmayer, 1977: 141-142, text-fig. 145 [Bimini].
Axociella spinosa; de Laubenfels, 1936a: 113 [Florida]; de
Laubenfels, 1949a: 16 [Bimini]; Storr, 1964: 42 [W coast
Florida]; Hechtel, 1965: 43.
Clathria (Microciona) spinosa; Van Soest, 19846: 95-96,
table 4, text-fig.37 [Curacao].
Clathria spinosa; Zea, 1987: 167, text-fig. 57 [Colombian
Caribbean].
cf. Microciona prolifera; Vosmaer, 1935a: 608, 645.
MATERIAL. HOLOTYPE: USNM7680. Caribbean.
Clathria (Microciona) spongigartina (de
Laubenfels, 1930)
Aaata spongigartina de Laubenfels, 1930: 27 [Carmel,
California].
Anaata spongigartina; de Laubenfels, 1932: 89-91, text-fig.
52; Sim & Bakus, 1986: 12 [California].
Clathria spongigartina; Van Soest, 19846: 7 [generic
synonymy].
MATERIAL. HOLOTYPE: USNM21428, fragment
BMNH1929.8.22.13. NE Pacific.
Microciona tenuis Stephens, 1915:443, p1.40, fig.5 [Saldanha
Bay]; Levi, 1963: 67.
Hymantho tenuis; de Laubenfels, 1936a: Ill.
MATERIAL. HOLOTYPE: RSME missing. South Africa.
Clathria (Microciona) tenuissima (Stephens,
1916)
Eurypon tenuissima Stephens, 1916: 240 [W coast Ireland;
780m depth]; Stephens, 1921.
Dictyociona tenuissima; de Laubenfels, 1936a: 110 [note]; de
Laubenfels, 1953a: 528.
Microciona tenuissima; Levi, 1960a: 69 [W coast Ireland];
Biblioni, 1993: 312-3, fig.8 [W. Mediterranean].
Not Leptolabis tenuissima Hentschel, 1911: 360-361, textfig.41 [Shark Bay, Western Australia].
MATERIAL. HOLOTYPE: INMSR151.11. 1916. NE Atlantic.
Clathria (Microciona) tetrastyla (Hentschel,
1912)
Hymeraphia tetrastyla Hentschel, 1912: 379-380, p1.20,
fig.36 [Am I., Arafura Sea].
Eurypon tetrastyla; de Laubenfels, 1936a: 111.
MATERIAL. HOLOTYPE: SMF 954. Indonesia.
Clathria (Microciona) thielei (Hentschel, 1912)
Hymeraphia thieleiHentschel, 1912:377-378 [Aru I., Arafura
Seat
Not Ophlitaspongia thielei Burton, 1932a: 322, p1.55, fig.8,
text-fig.32; Koltun, 1964a: 70.
MATERIAL. HOLOTYPE: SMF 1708. Indonesia.
Clathria (Microciona) stephensae sp. nov.
Clathria (Microciona) toximajor Topsent, 1925
Microciona similis Stephens, 1915: 441, p1.40, fig.6 [Saldanha Bay]; Levi, 1963: 58-59, text-fig.67, p1.9A [St. Helena
Bay]; Hechtel, 1965: 43-44 [note].
Axocielita sitnilis; de Laubenfels, 1936a: 118.
Not Hyrneraphia similis Thiele, 1903a: 957.
Clathria toximajorTopsent, 1925: 653-655, text-fig.11 [Gulf
of Naples]; Sara, 1960a: 461 [Ischia].
Microciona toximajor, Levi, 1960a: 67 [Naples]; Siribelli,
1960: 6, text-fig.1B [Naples]; Sara & Siribelli, 1962: 48
[Gulf of Naples]; Poggiano, 1965: 3, table 1; Riitzler, 1966
226zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
[Banyuls]; Boury-Esnault, 1971: 325 [Banyuls]; PulitzerFinali, 1983: 610 [list].
MATERIAL. HOLOTYPE: MOM, fragment MNHNDT326.
Mediterranean
Clathria (Microciona) toxirecta (Sara &
Siribelli, 1960)
Microciona toxirecta Sara & Siribelli, 1960: 75-77, textfig.21 [Bay of Naples]; Siribelli, 1960: 14, text-fig.5A
[Naples]; Sara & Siribelli, 1962: 48 [Gulf of Naples];
Pulitzer-Finali, 1983: 610 [list].
MATERIAL. HOLOTYPE: IMZUN382. Mediterranean.
Clathria (Microciona) toxitenuis Topsent,
1925
Clathria toxitenuis Topsent, 1925:655 [Gulf of Naples]; Sara,
1958: 261-262, text-fig.23 [Gulf of Naples]; Sara, 1960a:
462 [Ischia].
Pseudanchinoe toxitenuis; de Laubenfels, 1936a: 109 [note].
Microciona toxitenuis; Levi, 1960a: 69, text-fig.11 [Marseilles, Naples]; Sara & Siribelli, 1960:69-71 [Bay of Naples];
Siribelli, 1960: 10-12, text-fig.4 [Naples]; Sara & Siribelli,
1962: 48 [Gulf of Naples]; Labate, 1964: 334-335, p1.2,
fig.8 [Adriatic Sea]; Sara, 1964: 230 [Ligurian Sea]; Poggiano, 1965: 3, table 1; Cabioch, 1968a: 244 [Roscoff,
France]; Pulitzer-Finali, 1977: 62 [Bay of Naples]; BouryEsnault, 1971: 325 [Banyuls]; Pulitzer-Finali, 1983: 573,
610 [Mediterranean]; Wintermann-Kilian & Kilian, 1984:
134 [Colombia; possible misidentification].
MATERIAL. HOLOTYPE: MOM, fragment MNHNDT325.
Mediterranean, NE Atlantic.
Clathria (Microciona) tumulosa (Bowerbank,
1882)
Microciona tumulosa Bowerbank, 1882: 7, 18, 50-52, p1.11,
figs 1-4 [Westport Bay, Ireland]; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: BMNH1910.1.1.82. NE Atlantic.
Clathria (Microciona) tunisiae sp. nov.
Microciona chelifera Levi, 1960a: 70, fig. 12 [Sicily-Tunisian
Strait]; Pulitzer-Finali, 1983: 610.
MATERIAL. HOLOTYPE: MNHN missing. W Mediterranean. Spanioplon (= Clathria) cheliferum Hentschel, 1911
has seniority.
Clathria (Microciona) urizae sp. nov.
Eurypon similis; Uriz, 1988a: 53-55, text-fig. 29 [Namibia].
Not Hymeraphia similis Thiele, 1903a: 957.
Not Microciona sitnilis Stephens, 1915: 441 [see C.(M)
stephensae sp. nov.].
MATERIAL. SPECIMENS: ABIP7B-58, 7B-59, 7B-61. SW
Africa. Note: E. similis of Uriz (1988a) is conspecific with
neither Clathria (Microciona) similis (Thiele, 1903a), nor
Microciona similis Stephens, 1915 (= C. (M.) stephensae sp.
nov.), and requires a new name.
ETYMOLOGY. Named for Dr. M.J. Uriz.
Clathria (Microciona) vacelettia nom. nov.
Microciona curvichela Vacelet & Vasseur, 1965: 106-108,
p1.9, fig.31 [Madagascar].
Not Wilsonella curvichela Hallmann, 1912: 247.
MATERIAL. HOLOTYPE: MNHN missing. W Indian
Ocean. C. (D.) curvichela (Hallmann, 1912) has priority.
TRANSFERS
Other species described in Clathria
(Microciona), or a synonym, but now transferred
to other genera.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP
Microciona ambigua Bowerbank, 1862b: 1110; Bowerbank,
1864: 188; Bowerbank, 1866: 124, 136-138 [Britain];
Gray, 1867: 535; Norman, 1869: 330; Schmidt, 1870: 76;
Bowerbank, 1874: 65, p1.25, figs 1-9; Marenzeller, 1878:
2, 4, 5, 14-15, 370, p1.1, fig.3, p1.2, figs 3-3a; Urban, 1880:
257; Bowerbank, 1882:7,18,53; Topsent, 1891a: 528,543,
554 [Roscoff, France]; Norman, 1892:6,11; Vosmaer,
1935a: 607.
Hastatusambiguus; Fristedt, 1885: 31-32, p1.3, fig. la-h; Fristedt, 1887: 443, 465.
Plocamia ambigua; Topsent, 1894a: 8, 14, 21-22, 23, 26;
Topsent, 1895: 214, 216; Topsent, 1896: 115, 127; Topsent, 1898: 226; Topsent, 1900: 112-113; Topsent, 1904a:
10, 24, 26, 154-155, 201 [et var. elegans; Azores]; Arnesen, 1903: 22-23, p1.3, fig.5 [Norway]; Thiele, 1903b:
389, 395, 397, p1.21, fig.21; Arndt, 1913: 119; Topsent,
1913b: 6, 7,32, 63, p1.5, fig.15; Hentschel, 1914: 120;
Burton, 1930a: 494 [Norway]; Hentschel, 1929: 895, 973
[White Sea].
Plocamionida ambigua; Topsent, 1927: 1-19; Alander, 1942:
53 [Sweden]; Lilly et al., 1953 [Lough Ine, Ireland]; Burton, 1959b: 39 [Iceland]; Vacelet, 1969: 208 [Mediterranean]; Van Soest & Weinberg, 1980: 10 [Lough Inc.
Ireland]; Uriz & Rosell, 1990:387-388, figs 4g-k [Mediterranean]; Ackers, Moss & Picton, 1992: 137 [Ireland].
Scopalina ambigua; Schmidt, 1866a: 149; Schmidt, 1866b:
15; Schmidt, 1868: 26, 40.
Amphilectus ambiguus; Vosmaer, 1880: 116.
Stylostichon ambiguum; Hanitsch, 1894: 176, 195.
Hymedesmia indistincta Bowerbank, 1874: 303-306, p1.87,
figs 1-10.
Myxilla indistincta; Vosmaer, 1880: 129.
Hymeraphia indistincta; Hanitsch, 1894: 181, 196.
Plocamia microcionides; Carter, 1876: 390 [Cape St. Vincent]; Topsent, 1891a: 529, 544-545 [Roscoff]; Topsent,
1892a: 117.
MATERIAL. HOLOTYPE: BMNH1930.7.3.227, fragment
BMNH1910.1.1.65. Referred to Anchinoidae, Plocamionida.
Microciona bihamigera Waller, 1877: 261 [Torbay,Britain;
nomen nudum]; Waller, 1878: 1, pis 1-2 [new, cf. Zool.
Rec. 1877]; Ridley & Dendy, 1887:139; Vosmaer, I935a:
608.
Stylostichon bihamigera; Lilly et al., 1953 [Lough 1ne, Ire.].
Pronaxbihamigera; Van Soest & Weinberg, 1980: 10 [Lough
Ine,lreland].
MATERIAL. HOLOTYPE: unknown. Referred to
Anchinoidae, Pronax.
Microciona dives Topsent, 1891a: 529, 543-544, 554, p1.22,
figs 2-3 [Roscoff, France].
Stylostichon dives; Pulitzer-Finali, 1983: 567 [Mediterranean]; Ackers, Moss & Picton, 1992: 136 [Ireland].
cf. Hymedesmia zetlandica; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: MOM, fragments
BMNH1910.1.1.498-9. Referred to Anchinoidae, probably
Plumohalichondria.
Microciona fictitia Bowerbank, 1866: 124-126 [Guernsey,
Britain]; Vosmaer, 1935a: 607.
Hytnedestnia fictitia; Alander, 1942: 36 [ Sweden].
Anchinoe fictitius; Stephens, 1916: 242 [W coast of Ireland].
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
227
Phorbas fictitius; Lilly et al., 1953 [Lough the, Ireland]; Van
Soest & Weinberg, 1980: 10 [Lough Inc. Ireland].
MATERIAL. HOLOTYPE: BMNH1930.7.3. 199, fragment
USNM5043. Referred to Anchinoidae, Phorbas.
Microciona intexta Carter, 1876: 238-239, p1.15, fig.43a-c
[Cape St. Vincent, Hebrides]; Vosmaer, 1935a: 607.
Rhabderemia intexta; Topsent, 1892a: 116; Topsent, 1904a:
152 [transferred with hesitation]; Hooper, 1990: 72; Van
Soest & Hooper, 1993: 337.
Rhabdosigma intexta; Hallmann, 1916b: 520; Hallmann,
1917: 398-399.
MATERIAL. HOLOTYPE: BMNH1890.4. 10.12. Referred
to Rhabderemiidae, Rhabderemia.
Microciona minutula Carter, 1876: 479, p1.16, fig.5 I [justified emendation]; Carter, 1880a: 44; Vosmaer, 1935a:
608.
Rhabderemiaminutula; Topsent, 1904a: 152-153, p1.1, fig. 10,
p1.13, fig. 13 [Banyuls]; Levi, 1956b: 393, fig.2; Boury-Esnault, 1971: 306 [Banyuls]; Biblioni & Gili, 1982: 231;
Pulitzer-Finali, 1983: 533-534, text-fig.51 [Mediterranean].
Microciona pusilla Carter, 1876: 239, p1.16; Carter, 1880c:
437; Topsent, 1889: 41, text-fig. 7.
Rhabderemia pusilla; Topsent, 1892a: 116; Hallmann, 1917:
399; Dendy, 1922: 85; Van Soest, 1984b: 534; Hooper,
1990: 72 [note]; Van Soest & Hooper, 1993: 323.
Rhabderemia indica, in part; Sara, 1961: 44, text-fig. 8;
Pulitzer-Finali, 1983: 534.
MATERIAL. HOLOTYPE: BMNHI902.11. 16.32, fragment
BMNH1954.3.9.178. Referred to RhabderemiicLae, Rhabderemia.
Axinella monticularis Ridley & Dendy, 1886: 481; Ridley
& Dendy, 1887: 185, p1.38, fig.5 [Cape Verde Is].
Aulospongus monticularis; Hallmann, 1917: 373 [footnote];
Hooper, 1991: 1307 [note].
Microciona monticularis; Burton, 1956: 132 [Sao Vincente,
W. Africa].
Aulospongiella monticularis; Burton, 1956: 141.
MATERIAL. HOLOTYPE: BMNH1887.5.2.20, paratype
BMNH1887.5.2.273. Referred to Raspailiidae, A ulospongus.
Spongia plumosa Montagu, 1818: 116 [Devon, UK].
Hymeniacidon plumosa; Bowerbank, 1866:195, figs 141-143.
Pronax plutnosa; Gray, 1867: 536.
Microciona plumosa; Bowerbank, 1874: 61-63, p1.24, figs
7-13; Topsent, 1891b: 128 [France].
Myxilla (?) plumosa; Ridley & Dendy, 1887: 145-146 [Bahia,
Brazil].
Stylostichon plumosa; Topsent, 189Ia: 529; Lilly et al., 1953
[Lough Ine, Ireland].
Plumohalichondria plumosa; Kerville, 1901: 175 [Normandy].
Hymedesmia plumosa; Vosmaer, 1935a: 607.
Pronax plumosus; Van Soest & Weinberg, 1980: 10 [Lough
[tie, Ireland].
MATERIAL. HOLOTYPE: unknown, fragments
BMNH1930.7.3.216, 224. Referred to Anchinoidae, Pronax.
MATERIAL. HOLOTYPE: BMNH1902.11. 16.32. Referred
to Rhabderemiidae, synonym of Rhabderemia minutula
(Dendy, 1905: 180).
Microciona quinqueradiata Carter, 1880a: 43, 153, p1.4, figs
5a-e [Gulf of Manaar].
cf.Microciona prolifera; Vosmaer, 1935a: 608.
MATERIAL. HOLOTYPE: LFM destroyed, no fragment in
BMNH. Referred to Raspailiidae, Cyamon.
Microciona simplicissima Norman, 1869: 330; Bowerbank,
1874: 198, 204-205, p1.73, figs 16-19 [Shetland]; Vosmaer, 1935a: 607.
Tedania simplicissima; Hanitsch, 1890: 192.
Bubaris simplicissima; de Laubenfels, I936a: 131.
Not Leptosia simplicissima Hentschel, 1911: 359-360, textfig.40 [Shark Bay, Western Australia].
Not Protoclathria simplicissima Burton, 1932a: 321, p1.56,
fig.2, text-fig.31 [Falkland Is].
MATERIAL. HOLOTYPE: BMNH1930.7. 30.212. Referred
to Axinellidae, Bubaris.
Microciona spinulenta Bowerbank, 1866: 124, 132, 133
[Britain]; Gray, 1867: 534; Vosmaer, 1935a: 607.
Pocillon spinulenta; Topsent, 1893b: 34 [plus Isodictya implicita Bowerbank].
MATERIAL. HOLOTYPE: BMNH1930.7. 3.213. Referred
to Myxillidae, lophon.
Microciona virgula Sara & Siribelli, 1960: 77-79, text-fig.22
[Bay of Naples]; Siribelli, 1960: 16-17, text-fig.7A
[Naples].
MATERIAL. HOLOTYPE: IMZUN237: 0.5. Synonym of
Antho involvens (Sara, 1964: 228-229).
Clathria (Dendrocia) Hallmann, 1920
Dendrocia Hallmann, 1920: 767.
Paradoryx Hallmann, 1920: 767.
Wilsonella in part; sensu Hallmann, 1912: 242 (not
Carter, 1885f: 366).
DEFINITION. Single undifferentiated category
of smooth auxiliary spicule (style, subtylostyle or
modified style) forming plumose or plumoreticulate choanosomal tracts, ectosomal brushes
and dispersed between skeletal tracts; echinating
acanthostyles usually heavily spined and distributed evenly over skeletal tracts; microscleres
include isochelae ranging from typical palmate
form (straight shaft, lateral alae fused to shaft),
modified palmate forms (thickened, curved shaft,
partially detached lateral alae) to anchorate-like
forms (alae detached from shaft, shaft with lateral
ridge); toxas if present include oxhorns.
Microciona pusilla Carter, 1876: 239, p1.16, figs 51a-d [?
TYPE SPECIES. Claihria pyramida Lendenfeld,
tropical]; Carter, 1880c: 437 [name emended to M.
minutula]; Topsent, 1889: 41, text-fig. 7 [Banc de
Campeche].
Rhabderemia pusilla; Topsent, 1892a: 116; Hallmann, 1917a:
399; Dendy, 1922: 85; Pulitzer-Finali, 1983: 533-534,
fig.51 [Mediterranean]; Van Soest, 1984b: 108. Van Soest
& Hooper, 1993: 323.
REMARKS. Seven species are included in
Clathria (Dendrocia), all of which are endemic
to temperate Australian waters, with an
hypothesised Gondwanan origin.
1888: 222 (by original designation).
^ MEMOIRS OF THE QUEENSLAND MUSEUM
228zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathria (Dendrocia) curvichela
(Hallmann, 1912)
(Figs 104-105)
Wilsonella curvichela Hallmann, 1912: 247-249,
p1.34, fig.4, text-fig.51.
Paradoryx curvichela; Hallmann, 1920: 768.
Clathria curvichela; Hooper & Wiedenmayer, 1994:
263.
Not Microciona curvichela Vacelet & Vasseur, 1965:
106.
MATERIAL. HOLOTYPE: AMZ59 (part) + E926a
(part): 21km S. of St. Francis I., SA, 32°44'S,
133 ° 18'E, 60m depth, coll. FIV `Endeavour' (dredge).
HABITAT DISTRIBUTION. 60m depth; substrate unknown; S Aust (Fig. 104E).
DESCRIPTION. Shape. Branching digitate,
130mm high, 35mm maximum width, with short,
cylindrical, bifurcate, anastomosing, tapering
branches up to 14mm diameter, and short subcylindrical stalk, 15mm long, 8mm diameter.
Colour Live colouration unknown, yellowishbrown in ethanol.
Oscules. Minute oscules, up to 1 mm diameter,
dispersed on lateral margins of branches
Texture and surface characteristics. Smooth,
even, compressible.
Ectosome and subectosome. Ectosome not
hispid, although peripheral spongin fibres
produce small surface projections; subectosomal
auxiliary subtylostyles tangential to surface, or
protruding through ectosome only a short way.
Choanosome. Choanosomal skeletal architecture
almost regularly reticulate, with heavy, thick
spongin fibres incompletely divided into primary
and secondary components, differentiated only
by presence or absence of coring auxiliary
megascleres, respectively; spicule tracts in
primary fibres vaguely ascend to surface in multispicular bundles; secondary fibres pauci- or
aspicular, usually running parallel to surface; all
fibres very heavily echinated by acanthostyles,
dispersed evenly over fibres; mesohyl matrix
heavy but only lightly pigmented, with scattered
microscleres.
Megascleres. Choanosomal principal styles absent or undifferentiated from auxiliary spicules.
Subectosomal auxiliary subtylostyles (coring
fibres and scattered below membraneous ectosome) straight or slightly curved, hastate, with
subtylote, slightly pointed, smooth bases. Length
132-(149.5)-186p.,m, width 1.543.5)-411m.
Acanthostyles subtylote, with large and even
spination. Length 64474.1)-8911m, width 4(6.8)-811m.
Microscleres. Isochelae palmate, large, with
greatly curved shaft; lateral alae fused with shaft
about 3/4 way along ala; front ala ranges from
well developed to vestigial; poorly silicified
sigma-like forms also present. Length 22423.8)32p,m.
Toxas absent.
REMARKS. Hallmann (1912) originally assigned this species to Wilsonella because it had
only one type of auxiliary spicule coring the
fibres and scattered in the subectosomal skeleton.
However, it lacks detritus in fibres, and principal
spicules, and has modified palmate isochelae
(curved, thickened with partially detached lateral
alae) and heavily echinated fibres indicates that
it belongs with Derzdrocia. Hallmann (1920)
erected Paradoryx for this and several other
species (C. oxyphila, C. piniformis, C. dura and
C. elegantula), of which the present species has
the most heavily echinated fibres and the most
regular fibre reticulation, although slightly
plumose near the periphery. Other species of
Dendrocia have predominantly plumose (or
plumo-reticulate) choanosomal skeletons.
Clathria (Dendrocia) dura Whitelegge, 1901
(Figs 106-107, Table 23)
Clathria dura Whitelegge, 1901: 83, 84, 117, p1.11,
fig. 11; Hooper & Wiedenmayer, 1994: 263.
? Wilsonella dura; Hallmann, 1912: 242, 244, 245,
298; Shaw, 1927: 426; Guiler, 1950: 9.
Paradoryx dura; Hallmann, 1920: 768.
cf. Microciona prolifera; sensu Vosmaer, 1935a: 611,
644.
Not Antherochalina dura Lendenfeld, 1887b: 788.
Not Clathria dura var. mo//is Hentschel, 1911: 370.
MATERIAL. LECTOTYPE: AMG3046: (dry) Tuggerah Beach, NSW, 3318'S, 151 °30'E, coll. NSW
Fish Commission (trawl). PARALECTOTYPE:
AMG3046a: (dry) same locality.OTHER
MATERIAL: NSW- AMZ1052.
HABITAT DISTRIBUTION. Ecology unknown; Tuggerah Beach, Balmoral Beach and Port Jackson (NSW)
(Whitelegge, 1901), Maria I. (Tas.) (Shaw, 1927;
Guiler, 1950) (Fig. 106E).
DESCRIPTION. Shape. Flabellate or flabellate-
digitate, lobate, planar or multiplanar growth
form, 95-110mm high, 85-210mm wide, composed of fused anastomosing or free digits and
flattened lobes, on small compressed basal stalk,
8-15rrim diameter, and branches bifurcate and
taper towards apex.
Colour Sandy yellow preserved.
Oscules. Oscules mostly confined to lateral margins of branches.
^
229
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
FIG. 104. Claihria (Dendrocia) curvichela (Hallmann) (portion of holotype AMZ59). A, Subectosomal auxiliary
subtylostyle. B, Echinating acanthosty le. C, Modified palmate isochelae. D, Section through peripheral skeleton.
E, Australian distribution. F, Holotype.
MEMOIRS OF THE QUEENSLAND MUSEUM
230^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 105. Clathria (Dendrocia) curvichela (Hallmann) (portions of holotype A-B, AME926a, AMZ59). A,
Choanosomal skeleton. B, Fibre characteristics (x744). C, Subectosomal auxiliary subtylostyle. D, Echinating
acanthostyle. E, Acanthostyle spines. F, Base of auxiliary subtylostyle. G-I, Modified palmate isochelae with
full (G) or vestigial front alae (I).
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQP
231
FIG. 106. Clathria (Dendrocia) dura Whitelegge (holotype AMG3046). A, Subectosomal auxiliary style/
subtylostyles. B, Echinating acanthostyles. C, Modified palmate isochelae. D, Section through peripheral
skeleton. E, Australian distribution. F, Holotype.
Texture and surface characteristics. Firm,
flexible; even, microscopically reticulate and
porous.
Ectosome and subectosome. Ectosome with nearly continuous palisade of acanthostyles standing
erect on peripheral fibres, with plumose brushes
of subectosomal auxiliary styles projecting and
forming bundles; spicule bundles correspond in
position with ascending choanosomal primary
spongin fibres; subdermal region also with scattered subectosomal megascleres lying tangential
to surface, interdispersed with projecting acanthostyles.
^ MEMOIRS OF THE QUEENSLAND MUSEUM
232zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 107. Clathria (Dendrocia) dura Whitelegge (paratype AMG3046a). A, Choanosomal skeleton. B,
Peripheral skeleton. C, Fibre characteristics. D, Modified palmate isochelae. E, Echinating acanthostyle. F,
Acanthostyle spination. G, Subectosomal auxiliary style. H-I, Bases and apex of auxiliary styles.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
233
TABLE 23. Comparison between present and published records of Clathria (Dendrocia) dura
(Whitelegge). Measurements in pm, denoted as range
(and mean) of spicule length x spicule width (N=25).
SPICULE
Choanosomal
principal
styles
Lectotype
Paralectotype
Specimen
(AMG3046)
(AMG3046a)
(N=1)
absent
absent
absent
78-(88.2)-102
x 3 - (4.1) - 5.5
82496.5)-115
x 3 - (3.6) - 5
56-(64.6)-75 x
4-(5.6)-9
14-(17.4)-19
61-(66.3)-75 x
4 - (5.1) - 8.5
16-(18.8)-22
absent
absent
Subectosomal 58487.4)-105
auxiliary
x 2 - (3.9) - 6
styles
48-(57.0)-66 x
Echinating
acanthostyles
4-(5.5)-8
17-(18.6)-22
Chelae
Toxas
absent
Choanosome. Choanosomal skeletal architecture
irregularly reticulate, with sinuous, relatively
thick spongin fibres; fibres incompletely divided
into primary ascending, and secondary transverse
components; primary fibres with a multispicular
core of subectosomal auxiliary styles, identical to
those in ectosomal skeleton, together with heavy
tracts of acanthostyles secondarily incorporated
into fibres; secondary fibres with multi- or
paucispicular core of both subectosomal and
acanthostyle megascleres; acanthostyles echinate
fibres in moderate quantities, at right angles to
skeleton, also lying inside the spicule tracts;
mesohyl matrix heavy, darkly pigmented,
granular with scattered megascleres and
microscleres of all kinds.
Megascleres (Table 23). Choanosomal principal
megascleres absent or undifferentiated from
auxiliary spicules.
Subectosomal auxiliary styles quasidiactinal,
curved or straight, hastate, with tapering,
rounded, styloid, or pointed, smooth bases.
Acanthostyles rounded or slightly subtylote,
with large, evenly dispersed spines.
Microscleres (Table 23). Isochelae abundant, palmate, with lateral alae fused to shaft for only
about 1/2 length of ala; shaft greatly curved,
thickened ('hunchback'), front ala well
developed.
Toxas absent.
Larvae. Incubated parenchymella, up to 2701Lm
in diameter, with heavy mesohyl matrix.
REMARKS. In possessing a single category of
structural megasclere this species is assigned to
C. (Dendrocia). Its quasidiactinal megascleres
are reminiscent of those found in C. (D.)
pyramida; the modified palmate isochelae
(curved, thickened, partially detached lateral
alae) are similar to those found in other species
assigned to Paradoryx by Hallmann (1920) (e.g.,
C. (D.) curvichela); and skeletal architecture and
spongin fibre characteristics are reminiscent of
those in C. (C.) sartaginula. The presence of
acanthostyles incorporated into skeletal spicule
tracts is now known to occur in several species,
collectively termed here the `phorbasiformis'
group (including C. (Thalysias) phorbasiformis,
C. (T) orientalis, C. (Dendrocia) dura, C. (D.)
impelfecta, C. (D.) myxilloides and C. (Clathria)
squalorum), and this feature is analogous to (but
not homologous with) that seen in the Crellidae.
The presence of acanthostyles in the ectosomal
skeleton is also seen in the Crellidae. Whereas
species of Crella have a thick tangential ectosomal crust of acanthose megascleres, in C.
(D.) dura these acanthostyles stand erect on
peripheral skeletal fibres, pierced by plumose
tracts of smooth subdermal (auxiliary) spicules.
The synonymy cited above for C. (D.) dura
requires further comment. Lendenfeld (1887a)
described Antherochalina dura from Port Jackson, but the species is unrecognisable from his
brief description. Lendenfeld's type material was
not located in the AM collections, and only a slide
of the holotype was found in the BMNH collections (BMNH1886.8.27.684). Apparently a dry
specimen of the species was also deposited in the
BMNH collections (BMNH1886.8.27.608, possibly the holotype), but this has not yet been
rediscovered. Re-examination of the holotype
slide confirms that A. dura is a synonym of
Phakellia flabellata, as supposed by Burton
(1934a).
Clathria (Dendrocia) elegantula Ridley &
Dendy, 1886
(Figs 108-109)
Clathria elegantula Ridley & Dendy, 1886: 474; Rid-
ley & Dendy, 1887: 149, p1.28, figs 3,3a, p1.29, figs
14a-b; Guiler, 1950: 7; Carpay, 1986: 26; Hooper &
Wiedenmayer, 1994: 263.
Clathria elegantula var. occidentalis Hentschel, 1911:
372-374, text-fig.46.
Wilsonella elegantula; Hallmann, 1912: 241.
Paradoryx elegantula; Hallmann, 1920: 768.
MATERIAL. HOLOTYPE: BMNH1887.5.2.91:
Moncoeur I., Bass Strait, Tas, 39 ° 14'S, 146 ° 30'E, coll.
HMS 'Challenger' (dredge). HOLOTYPE of var. occidentalis: ZMB4445: Top of inner bar, Shark Bay,
WA, 25°30'S, 113 ° 03'E, 6-9m depth, coll. W.
Michael sen & R. Hartmeyer (dredge).
HABITAT DISTRIBUTION. 6-76m depth; on sand
and shell grit substrate; Bass Strait and E coast (Tas)
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^
Clat hria ( Dendrocia) elegant ula Ridley & Dendy (holotype BMNH1887.5.2.91). A, Subectosomal
FIG. 108. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
auxiliary style/ subtylostyle. B, Echinating acanthostyles. C, Modified sigmoid palmate isochelae. D, Section
through peripheral skeleton. E, Australian distribution. F, Holotype. G, Section of peripheral skeleton of variety
occident alis (holotype ZMB4445).
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
235
FIG. 109. Clathria (Dendrocia) elegantula Ridley & Dendy (holotype BMNH1887.5.2.91). A, Choanosomal
skeleton. B, Peripheral skeleton. C, Fibre characteristics. D, Modified sigmoid palmate isochelae. E, Echinating
acanthostyle and modified form. F, Acanthostyle spination. G-H, Bases and apex of subectosomal auxiliary
subtylostyles.
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^
(Ridley & Dendy, 1886; Carpay, 1986), central W
coast (WA) (Hentschel, 1911) (Fig. 108E).
DESCRIPTION. Shape. Massive, lobate fan,
73mm long, 75mm wide, with erect, flattened,
digitate lobes, 22-31mm long, 8-20mm wide,
lobes with uneven margins, arising from sprawling encrusting base.
Colour Live colouration unknown, pale beige
preserved.
Oscules. Differentiated inhalant and exhalant surfaces of lobes; ostia scattered, 1-2mm diameter,
oscules confined to sieve-plates.
Texture and surface characteristics. Harsh; uneven, prominently microconulose, translucent
dermal membrane, covering reticulate fibrous
surface, stretched between microconules.
Ectosome and subectosome. Fibrous, reticulate
ectosomal skeleton, with sparse tangential subectosomal auxiliary subtylostyles lying directly on
surface; sinuous plumose tracts of these spicules
below, ascending from choanosomal skeleton.
Choanosome. Choanosomal skeleton plumoreticulate, cavernous, with well developed
primary and secondary spongin fibres; primary
ascending fibres sinuous, almost dendritic, cored
by several discrete multi- or paucispicular tracts
of subectosomal auxiliary subtylostyles, occupying only small portion of fibre diameter; primary
fibres interconnected by reticulate secondary
fibres, and terminating in plumose spicule
bundles at periphery; secondary transverse fibres
pauci- or aspicular, thin; echinating acanthostyles
most abundant in axial region, sparse or absent in
peripheral skeleton; mesohyl matrix light, with
abundant scattered chelae.
Megascleres. Choanosomal principal
megascleres absent or undifferentiated from
auxiliary spicules.
Subectosomal auxiliary subtylostyles straight,
rarely slightly curved, quasidiactinal, hastate
points, with tapering, rounded or slightly pointed,
smooth bases. Holotype: Length 145-(160.6)168Rm, width 1.5 -(2.6)-4p,m. (Hentschel's
specimen: 133-(142.3)-152Rm, width 243.5)4p,m).
Acanthostyles slender, evenly spined, with
rounded or slightly subtylote bases, sharp points.
Length 53-(64.1)-74,m, width 2-(3.6)-5p,m.
(Length 52468.2)-84pm, width 3-(6.4)-81.1.,m).
Microscleres. Isochelae large, palmate, sigmoid
curved, with lateral alae nearly completely fused
to shaft, sometimes well developed, sometimes
vestigial and reduced to small ridge on shaft; front
ala usually reduced. Length 13415.4)-1711m.
(Length 13417.2)-2011m).
Toxas absent.
REMARKS. The nearly dendri tic, predominantly
plumose skeletal architecture and the concentration of acanthostyles in the axial region are quite
distinctive features for this species. These characters, together with the possession of a single
category of structural spicule in both fibres and
the ectosomal skeleton indicate that the species is
best placed in C. (Dendrocia). In having
quasidiactinal modified, vestigial structural
megascleres with principal and auxiliary spicules
having similar geometry, and vague similarities
in skeletal structure, this species is included in the
`oxyphila' species group (also containing C. (C.)
rap hana, C. (C.) oxyphila and C. (C.) piniformis).
Spongin fibres are also characteristic (whereby
each fibre may have several discrete spicule
tracts), but in most other respects this species is
very similar to C. (D.) dura Whitelegge (both
having sinuous spongin fibre systems and heavily
echinated fibres). These two species differ in the
size of their auxiliary styles (notably straight,
longer, more slender, with subtylote bases in C.
(D.) elegantula, whereas in C. (D.) dura these
spicules are curved, short, thick, with tapering
hastate (or sometimes pointed) bases)), and the
incorporation of acanthostyles into skeletal fibre
tracts in C. (D.) dura.
There is a remarkably close concordance in
these features between Bass Strait and Shark Bay
populations, despite their widely separated
geographical distribution. These samples differ
only slightly in the more robust acanthostyles
seen in Shark Bay material, but no other morphological differences were seen to justify subspecies separation (Hentschel, 1911).
Clathria (Dendrocia) imperfecta Dendy, 1896
(Figs 110-111)
Clathria imperfecta Dendy, 1896: 35; Ayling et al.,
1982: 103; Hooper & Wiedenmayer, 1994: 264.
Wilsonella imperfecta; Hallmann, 1912: 242.
MATERIAL. HOLOTYPE: NMVG2369 (RN376)
(fragment BMNH1902.10.18.335): Port Phillip Bay,
Vic, 38°09'S, 144°52'E, 36m depth, coll. J.B. Wilson
(dredge).
HABITAT DISTRIBUTION. 36m depth; substrate unknown; Port Phillip Bay (Vic) (Fig. 110E).
DESCRIPTION. Shape. Erect, bulbous-lobate,
with small cylindrical lobate digits.
Colour Colour in life recorded as dull brownorange exterior, yellow interior.
^
237
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
Oscules. Minute, less than 2mm diameter, mainly
on apex of lobes.
Texture and surface characteristics. Surface irregular, uneven, micropapillose, with subdermal
ridges and grooves. Texture is crumb-of-bread,
easily torn.
Ectosome and subectosome. Microscopically
hispid, with thick brushes of subectosomal
auxiliary styles from peripheral skeleton protruding through dermal crust, composed of same
megascleres, lying paratangential to or standing
erect on surface; subectosomal architecture thickly plumose, arising from ultimate ascending
choanosomal fibres.
Choanosome. Choanosomal skeleton irregularly
plumo-reticulate, with prominent primary ascending spicule tracts interconnected by irregularly dispersed secondary tracts; fibres
weakly developed in spongin but fully cored by
subectosomal auxiliary styles and also acanthostyles secondarily incorporated into tracts; fibres
moderately echinated; fibre anastomoses form
irregular oval-elongate meshes; mesohyl matrix
heavy but only lightly pigmented, with auxiliary
styles and sparse detritus scattered thrroughout.
Megascleres. Choanosomal principal styles absent or undifferentiated from auxiliary
megascleres.
Subectosomal auxiliary spicules fusiform,
mostly straight, occasionally slightly curved
towards bases, with tapering, or rounded, or
slightly subtylote smooth bases. Length 189(213.3)-234p.m, width 1.544.4)-7.5p.m.
Acanthostyles long, rounded or faintly subtylote, with large and evenly distributed spines.
Length 102-(114.8)-122p,m, width 5-(5.8)-7p.m.
Microscleres absent.
Larvae. Oval-elongate parenchymella, 345412x275-370p.m, without larval spicules but
with heavy mesohyl and differentiated cells
clearly visible.
REMARKS. This species was assigned to Wilsonella by Hallmann (1912) because it has only
a single category of smooth megasclere, but was
tacitly returned to Clathria by Hallmann (1920)
when he restricted Wilsonella to the type.
Clathria (Dendrocia) impeifecta is more appropriately placed in Dendrocia because it lacks
detritus incorporated into fibres and has only one
undifferentiated category of smooth auxiliary
style, whereas Wilsonella (sensu Hallmann,
1912) and Paradoryx (sensu Hallmann, 1920)
have two categories of auxiliary styles, one found
in fibres and the other on the ectosomal skeleton.
All three nominal genera lack true principal
megascleres. This species is similar to C.
(Thalysias) orientalis, C. (T) phorbasifonnis, C.
(D.) myxilloides and C. (D.) dura in having acanthostyles incorporated into fibres, intermingled
amongst the smooth coring megascleres (termed
the `phorbasifonnis' species complex).
Clathria (Dendrocia) myxilloides
Dendy, 1896
(Figs 112-113, Plate 3A)
Clathria myxilloides Dendy, 1896: 35; Hallmann,
1920: 768; Ayling et al., 1982: 104; Hooper &
Wiedenmayer, 1994: 264.
Wilsonella myxdloides; Hallmann, 1912: 242.
MATERIAL. HOLOTYPE: NMVG2376 (RN729)
(fragment BMNH1902.10.18.334): Port Phillip Bay,
Vic, 38 ° 09'S, 144 ° 52'E, coll. J.B. Wilson (dredge).
OTHER MATERIAL: S AUST- QM G300613
(NCIQ66C-2202-K, fragment NTMZ3535).
HABITAT DISTRIBUTION. 18m depth; on rock substrate; Port Phillip Bay (Vic), Kangaroo I. (SA) (Fig.
112E).
DESCRIPTION. Shape. Massive, subspherical,
lobate digitate growth form.
Colour. Orange-red alive (Munsell 5R 5/10),
grey-brown in ethanol.
Oscules. Small, less than 3mm diameter, scattered evenly over surface.
Texture and surface characteristics. Harsh, compressible; shaggy, uneven, microconulose, appearing pock-marked alive due to scattering of
small oscules.
Ectosome and subectosome. Relatively dense
discrete bundles of subectosomal auxiliary styles,
identical to those coring the fibres, forming erect
palisade on surface. Tracts of isochelae also
found in peripheral skeleton, scattered
throughout heavily pigmented peripheral spongin; subectosomal region relatively disorganised,
paratangential, merging into peripheral
choanosomal fibres lying almost immediately
subectosomal.
Choanosome. Choanosomal skeletal architecture
irregularly plumo-reticulate, without clearly differentiated primary or secondary components although fibre diameter varies considerably and
fibres become sinuous towards surface; fibres
composed of only light spongin, heavily cored by
multispicular tracts of subectosomal auxiliary
styles and heavily, irregularly echinated by acanthostyles lying parallel with, or at acute angles to
spicule tracts but also secondarily incorporated
into fibres; mesohyl matrix light, with smaller,
MEMOIRS OF THE QUEENSLAND MUSEUM
238zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
FIG. 110. Clathria (Dendrocia) impelfecta Dendy (holotype NMVG2369). A, Subectosomal auxiliary styles. B,
Echinating acanthostyle. C, Section through peripheral skeleton. D, Parenchymella larva in situ (diameter
400um). E, Australian distribution. F, Holotype.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
239
FIG. 111. Clathria (Dendrocia) impetfecta Dendy (holotype NMVG2369). A, Choanosomal skeleton. B, Fibre
characteristics (x419). C, Echinating acanthostyle. D, Acanthostyle spination. E, Bases of subectosomal
auxiliary styles.
MEMOIRS OF THE QUEENSLAND MUSEUM
240^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 112. Clathria (Dendrocia) myxilloides Dendy (holotype NMVG2376). A, Subectosomal auxiliary subtylostyle. B, Echinating acanthostyle. C, Anchorate isochelae. D, Section through peripheral skeleton. E, Australian
distribution. F, QMG300613 alive. G, Holotype.
thinner auxiliary spicules, and numerous
isochelae organised into tracts surrounding large
choanocyte chambers.
Megascleres. ^Principal^choanosomal
megascleres absent, or undifferentiated from
auxiliary spicules.
Subectosomal auxiliary styles differ slightly in
size according to location within skeleton: those
coring fibres and in ectosomal skeleton relatively
homogenous, straight or slightly curved, hastate,
with smooth, tapering or slightly subtylote bases,
some appearing quasi-diactinal. Length 287(311.4)-330p,m, width 3.5-(4.9)-6Rm; auxiliary
styles also scattered between fibres probably
younger forms of main structural megascleres,
being thin, whispy, hastate, usually curved or
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
241
FIG. 113. Clathria (Dendrocia) myxilloides Dendy (holotype NMVG2376). A, Choanosomal skeleton. B, Fibre
characteristics. C, Acanthostyle spination. D, Echinating acanthostyles. E, Anchorate, unguiferous and sigmoid
isochelae.
242zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
sinuous, with rounded or tapering bases. Length
212-(233.0)-284Rm, width 2-(2.2)-31.1.m.
Acanthostyles long, slender, straight or slightly
curved near basal end, evenly and heavily spined.
Length 84-(112.6)-133t.tm, width 6-(7.4)-10iLm.
Microscieres. Isochelae relatively large, abundant, anchorate, with well formed lateral alae
detached from shaft nearly completely, lateral
ridge on shaft (possibly vestigial point of attachment of lateral alae); some modified unguiferous
forms with pointed alae, or sigmoid forms with
greatly reduced alae also present. Length 21(24.4)-29Rm.
Toxas absent.
REMARKS. This species is similar to C. (D.)
impetfecta in growth form and incorporation of
some acanthostyles into fibres (see remarks for C.
(Thalysias) phorbastfonnis). It also shows
similarities to C. (D.) pyratnida in growth form
and megasclere morphology, although all three
species differ in isochelae geometry.
Clathria (Dendrocia) pyramida Lendenfeld,
1888
(Figs 114-115, Table 24, Plate 3B)
Clathria pyramida Lendenfeld, 1888: 222; Capon &
MacLeod, 1987: 1200; Chernoff, 1987: 160;
Hooper et al., 1992: 263; Hooper & Wiedenmayer,
1994: 264.
Wilsonella pyramida; Hallmann, 1912: 240, 244;
Shaw, 1927: 426; Guiler, 1950: 9.
Dendrocia pyramida; Hallmann, 1920: 767.
Clathriaalata Dendy, 1896: 34; Hentschel, 1911: 375377, text-fig.48; Hentschel, 1923: 387, fig.356;
Ayling et al., 1982: 100; Van Soest, 1984b: 120.
Wilsonella alata; Hallmann, 1912: 241.
Dendrocia alata; Hallmann, 1920: 767.
MATERIAL. LECTOTYPE: AMG9047: Port Jackson, NSW, 33°51'E, 151 ° 16'E, no other details known.
PARALECTOTYPE: BMNH1887.1. 24.62 (fragment
AMG3579): same locality. LECTOTYPE of C. alata:
NMVG2280 (fragment BMNH1902.10.18.330): Port
Phillip Bay, Vic, 38°09'S, 144°52'E, coll. J.B. Wilson
(dredge, RN 752). PARALECTOTYPES of C. alata:
NMVG2281 (fragment AMG2686): same locality (RN
763). NMVG2282 (fragment BMNH1902.10.18.31):
Sorrento Jetty, Port Phillip Bay, 38°33'S, 145 ° 21'E
(dredge, RN 792). NMVG2283 (fragment
BMNH1902.10.18.49): same locality (RN 801).
NMVG2284 (fragment BMNH1902.10.18.332): same
locality (RN 842). NMVG2285 (fragment
BMNH1902.10.18.333): same locality (RN 843).
OTHER MATERIAL: VIC- AME298, AMZ1145,
AM unreg. (small vial, label 'donated by A Dendy'),
AM unreg. (fragment BMNH1887.4.27.103) (small
vial, label 'Clathria tethyopsis, donated by A Dendy,
Lendenfeld or Burton MS name'). NSW- AMZ3216,
NTMZ2667, QMG304507, QMG304522,
QMG304574. S AUST- SAMTS4095 (fragment
NTMZ1629), SAMTS 6290, QMG3005 03
(NCIQ66C-2119-T) (fragment NTMZ3520),
QMG301354, QMG301358, QMG301374,
QMG301382, QMG304041, QMG304049,
QMG304051, QMG304053, QMG304054,
QMG304063, QMG304064, QMG304069,
QMG304070. WA- QM G300604 (NCIQ66C-4271J), QM G300191 (NCIQ66C-4651-V). (Not AM
unreg. (label 'Clathria alba. Port Jackson, NSW', =
Lendenfeld or Burton MS name; = Crella incrustans)).
HABITAT DISTRIBUTION. 0.8-25m depth; on sand,
rock reef or muddy bottoms; associated with oysters
and algal beds, Spondylus, Pinna and Chalamus asperrimus bivalves, rock or wood jetty substrates; Illawarra, Port Jackson, N. Sydney, Port Hacking and Durras
I. (NSW); Maria I. (Tas); Port Phillip Bay (Vic); St.
Vincent Gulf, Yorke Peninsula and Kangaroo I. (SA);
Albany and Pelsart Is, Houtman Abrolhos (WA) (Fig.
114E).
DESCRIPTION. Shape. Massive, lobate, lobo-
digitate, irregular growth form with small lobate
surface projections.
Colour. Dark brown or orange-brown live exterior (Munsell 5YR 4/6-5/10), paler
choanosome, brown in ethanol.
Oscules. Few, relatively large oscules (up to 8mm
diameter) on apex of lobes, each with slightly
raised membraneous lip, and with membraneous,
divided exhalant canals visible inside oscule; oscules collapse on preservation.
Texture and surface characteristics. Firm, compressible; smooth, relatively even, bulbous,
fleshy surface in live state; optically smooth,
membraneous surface in preserved state, with
irregularly rugose microscopic conulose and
lobate projections.
Ectosome and subectosome. Microscopically
hispid, with dense crust of subectosomal
auxiliary styles forming continuous, erect,
regular (straight) or irregular (stellate, paratangential), plumose palisade.
Choanosome. Skeletal architecture irregularly
plumo-reticulate, with sinuous spongin fibres ascending to surface in meandering tracts; fibres
anastomose more frequently at axis than in
peripheral skeleton; peripheral fibres and skeletal
tracts often diverge becoming plumose in subectosomal region, or forming a paratangential layer
immediately below erect ectosomal skeleton;
fibres contain moderately light to very heavy
spongin, cored by multispicular, sinuous tracts of
subectosomal auxiliary styles; fibres not obviously divided into primary or secondary components,
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
243
although fibre diameter varies within same sections of skeleton; acanthostyles heavily echinating most fibres, occasionally rare or absent from
some; mesohyl matrix very heavy but only lightly
pigmented, with abundant isochelae and
auxiliary megascleres; microalgae present in
mesohyl of some specimens.
Megascleres (Table 24). Choanosomal principal
megascleres absent, or at least undifferentiated
from auxiliary spicules.
Subectosomal auxiliary styles are thin, hastate,
straight or slightly curved towards basal end,
usually with smooth, slightly subtylote bases that
taper towards end into a small point (partially
mucronate), points hastate.
Acanthostyles vary considerably in length,
usually club-shaped, subtylote, with large spines
mostly confined on base and more-or-less
aspinose towards point; spines usually robust.
Microscleres (Table 24). Isochelae large, very
abundant, primarily palmate, heavily silicified,
usually with thickened and slightly curved shaft,
large lateral alae completely fused to shaft resembling swings', with front ala free, but sometimes
with reduced alae and sigmoid curvature.
Toxas absent.
Larvae. Larvae oval-elongate parenchymella, up
to 518x34011m, with larval raphidiform spicules
scattered throughout the axis and forming stellate
tufts at the posterior pole.
REMARKS. Type material of Clathria pyramida
and of C. alata are virtually identical in their
growth form, surface features, spicule geometry,
spicule distribution and fibre characteristics, differing only slightly in choanosomal skeletal construction (predominantly plumo-reticulate versus
predominantly plumose to slightly plumo-reticulate), and ectosomal structure (either producing a
continuous erect palisade (C. alata s.s.) or
punctuated palisade resulting in stellate plumose
brushes (C. pyramida s.s.), respectively). But
there are no consistent or significant differences
between these species in the spongin content of
fibres, density of spicule tracts, presence or absence of conecting secondary fibres, or spicule
dimensions (Table 24), as supposed by Hallmann
(1912: 241). On this basis the two species are
merged here. For all other specimens examined
the main variability involves the degree to which
fibres were compacted (mesh size), the spongin
content of skeletal tracts ('fibre' diameter), and
spicule dimensions between different geographic
populations (Table 24). In this latter regard
populations from NSW and WA had relatively
larger, more robust acanthostyles than other
populations (VIC, SA). In particular, spicules
from a small population in the Gulf of St Vincent,
SA, were noticeably more poorly silicified and
less robust than 'typical' specimens found in all
other localities.
In spiculation and skeletal architecture C. (D.)
pyramida is remarkably similar to Crella incrustans var. digitata ('cotype' AME503), although
upon careful examination of both species there
are obvious differences in skeletal structure
(Crella with a tangential ectosomal layer), composition of the ectosomal skeleton (styles versus
acanthostyles or acanthoxeas), and spicule
geometry (quasi-diactinal auxiliary styles versus
anisoxeas or quasi-monactinal megascleres,
respectively). Nevertheless, these species
demonstrate remarkably close convergences in
several prominent features making them easily
confused in the field and laboratory.
Specimens from the south coast of NSW have
peculiar biochemistry and significant biological
activity against both gram positive and gram
negative bacteria (Hooper et al., 1992). Unlike
many other bioactive sponges, however, the activity in C. (D.) pyramida was found to be related
to a unique, modified free sugar, 5-Thio-D-mannose (Capon & MacLeod, 1987). This discovery
represents the only known occurrence to date of
that class of thiosugar in nature, although the
compound has a glucose-based synthesised
analogue which has been known for several
years. Thio sugars have considerable pharmaceutical and medical potential: they inhibit the
release of insulin and transport of glucose, and are
able to cause reversible inhibition of sperm-cell
development without displaying acute toxicity
(R. Capon, pers. comm.). The taxonomic significance of those compounds is not clear, but it
is possible to speculate on the biological role of
those chemicals. Recent evidence (A. Butler,
pers.comm.) suggests that C. (D.) pyramida
together with a few other species (e.g., Crella
incrustans) occur in association with (are
epizootic on) scallops (Chlamys asperrima) from
South Australian waters. Chemoff (1987) found
that the presence of those epizootics increased the
survival of the scallop in both field and caging
experiments and it is possible that the sponges
provide some sort of chemical defence of the
host: thio-mannose sugar may play some sort of
role in that defence.
244zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 114. Clathria (Dendrocia) pyramida Lendenfeld (holotype AMG9047). A, Subectosomal auxiliary subtylostyle. B, Echinating acanthostyle. C, Modified palmate isochelae. D, Section through peripheral skeleton.
E, Australian distribution. F, Holotype. G, Paralectotype of C. alata NMVG2283. H, QMG300238 in situ.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
245
FIG. 115. Clathria (Dendrocia) pyramida Lendenfeld (QMG301358). A, Choanosomal skeleton. B, Peripheral
skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spination. F-G, Modified
thickened and sigmoid palmate isochelae.
MEMOIRS OF THE QUEENSLAND MUSEUM
246^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hria ( Dendrocia) pyram ida
TABLE 24. Comparison between published records and different populations of zyxwvutsrqponmlkjihgfedcbaZYXWVUTS
Lendenfeld. All measurements are given in pm, denoted as range (and mean) of spicule length x spicule width
(N=25) (bracket = number of specimens per locality examined).
-^ -
-
Lectotype
(AMG9047)
SPICULE
Lectotype of C.
Alata
(NMVG2280)
Specimens NSW
(N=7)
Specimens VIC
(N=10)
Specimens SA
(N=16)
Specimens WA
(N=2)
Choanosomal
principal styles
absent
absent
absent
absent
absent
absent
Subectosomal
auxiliary styles
208-(220.6)-231
x 4-(6.4)-8
211-(229.4)-243
x 3-(4.8)-7
190-(209.5)-225
x 4-(5.1)-7
2024222.6)-247
x 4-(5.9)-9
195-(213.4)-228
x 4-(4.9)-7
82-(103.9)-145 x
5-(8.4)-12
68-(89.6)-I25 x 4(7.2)-12
64-(78.6)-90x 4(7.4)-11
214(227.4)-242
x 4-(5.3)-7
82-(103.5)-155 x
6-(9.0)-11
20423.6)-27
20-(22.5)-25
17-(22.1)-30
22-(24.3)-27
Echinating
acanthostyles
Chelae
87-(99.0)-135 x 8- 75-(85.2)- 118 x 7(10.2)-12
(9.2)-11
21-(23.8)-27
23-(24.5)-27
Clathria (Dendrocia) scabida (Carter, 1885)
(Figs 116-117, Table 25)
Carter, 1885b: 112, p1.4, figs
4-5; Carter, 1886g: 449.
Micr ociona scabida; Dendy, 1896: 31; Hal'mann,
1912: 150; Vosmaer, 1935a: 608.
? St ylot ellopsis or Clat hrissa scabida; Hallmann, 1912:
151.
Anaat a scabit a Papsusl; de Laubenfels, 1936a: 109.
Clat hria scabida; Hooper & Wiedenmayer, 1994: 264
Halichondria pust ulosa, in part; Carter, 1886g: 450.
Not Halichondria pust ulosa Carter, 1882a: 285, p1.11,
fig. 1.
Halichondr ia scabida
MATERIAL. HOLOTYPE: BMNH1887.7.11.9 (fragment AMG2760): Port Phillip, Vic, 38°09'S,
144 ° 52'E, coll. J.B. Wilson (dredge). OTHER
MATERIAL: VIC - NMVRN413, NMVRN1025,
NMVRN1038, BMNH1887.7.11.26 (slide AMG2916).
HABITAT DISTRIBUTION. 38m depth; substrate unknown; Port Phillip and Westernport Bays (Vic) (Fig.
116F).
DESCRIPTION. Shape. Low, globular, erect, arborescent, digitate sponge, 80-95mm high, 4570mm wide, with subspherical, lobate branches,
28-42mm long, 12-28mm diameter, on a short
stalk, 22mm long, 12-15mm diameter.
Colour Live colouration orange-brown, brown
exterior and yellow-brown interior preserved.
Oscules. Large oscules, up to 5mm diameter,
situated in groups (seive-plates) on all sides of
branches; seive-plates sunken, resembling
porocalyses of the Tethyidae.
Texture and surface characteristics. Surface uneven, fibrous, with numerous large pores, seiveplates and subdermal canals covered with
prominent membraneous skin, raised into small
papillae and ridges.
Ectosome and subectosome. Ectosome hispid,
with subectosomal auxiliary subtylostyles form-
ing sparse but continuous dermal palisade of erect
brushes, sometimes tangential, protruding from
surface, piercing heavy crust of isochelae lying
on ectosomal membrane; subectosomal region
plumose, with sinuous tracts of subectosomal
subtylostyles diverging from ends of
choanosomal fibres, and with numerous, relatively large subdermal cavities lined by isochelae.
Choanosome. Choanosomal skeleton irregularly plumo-reticulate, with sinuous, heavy, spongin
fibres cored by subectosomal auxiliary styles and
fewer larger acanthostyles, and echinated by two
sizes of acanthostyles; smaller acanthostyles
echinate fibres in relatively heavy, even concentrations, whereas larger acanthostyles most
common at fibre nodes and echinate fibre terminations in radiating brushes; spongin fibres not
divided into primary or secondary components,
and fibre anastomoses occur irregularly
throughout skeleton; mesohyl matrix very heavy,
with abundant isochelae scattered and also congregated around fibres and choanocyte chambers.
Megascleres (Table 25). Choanosomal principal
megascleres absent or at least completely undifferentiated from auxiliary spicules.
Subectosomal auxiliary subtylostyles, coring
fibres and in dermal membrane, long, thin, mostly
straight, hastate, with elongated, subtylote,
smooth bases.
Larger acanthostyles subtylote, slightly
curved, with sparse but large spines along shaft
and base, and usually with an aspinose apex.
Smaller acanthostyles subtylote, evenly and
heavily spined.
Microscleres (Table 25). Isochelae extremely
abundant, anchorate-like, with curved shaft,
lateral alae nearly completely detached from
shaft, completely free front ala, and lateral ridge
on shaft (possibly vestigial point of alae attach-
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
247
TABLE 25. Comparison between present and published records of Clathria (Dendrocia) scabida (Carter).
Measurements in i.tm, denoted as range (and mean) of
spicule length x spicule width (N=25).
SPICULE
Holotype
(BMNH1887.7.11.9)
Specimens (N.3)
absent
absent
Choanosomal
principal styles
201-(237.2)-273 x 3- 1964226.5)-278 x 2(4.4)-5.5
(3.7)-5
153-(191.6)-219 x 193-(201.1)-221 x6Echinating
acanthostyles I
5.5-(7.1)-9
(7.3)-9
78490.4)-98 x 382-(95.3)-105 x 5Echinating
acanthostyles II
(6.8)-10
(6.4)-8
16-(18.2)-21
19-(20.9)-24
Chelae I (sigmoid)
Subectosomal
auxiliary styles
Chelae II (arcuatelike)
24 - (29.2) - 33
28-(31.4)-34
Chelae III
(bipocilla-like)
14416.2)-18
18-(19.3)-22
35-(72.4)-120x 1(1.8) 3 uncommon
70-(121.1)-156x 1-
Toxas
-
(2.2)-3 common
known species of Clathria (Dendrocia) with toxa
microscleres.
Clathria (Axociella) Hallmann, 1920
Axociella Hallmann, 1920: 779; Bergquist & Fromont,
1988: 116.
Axosuberites Topsent, 1893a: 179.
Tenaciella Hallmann, 1920: 772.
DEFINITION. Well differentiated axial and
extra-axial skeletal architecture (reminiscent of
Raspailiidae); axial skeleton markedly compressed, reticulate; extra-axial skeleton radial,
plumose or plumo-reticulate composed of large
subectosomal auxiliary styles-subtylostyles;
specialised ectosomal skeleton present composed
of smaller auxiliary spicules; echinating
megascleres absent although principal spicules
may protrude through fibres at acute angles.
TYPE SPECIES. Esperiopsis cylindrica Ridley &
Dendy, 1886: 340 (by original designation).
ment); reduced chelae also present, unguiferous,
with pointed alae.
Toxas oxhorn, with wide central curvature and
slightly reflexed arms.
REMARKS. Carter's (1885a) holotype
(BMNH1887.7.11.9), and his (1886g) specimen
of H. pustulosa, BMNH1887.7.11.26 are identical, as suspected from their respective descriptions, whereas H. pustulosa from the Falkland Is
(Carter, 1882a) is different.
Clathria scabida has skeletal architecture, fibre
structure, ectosomal characteristics and spicule
geometry characteristic of Dendrocia, whereas
the presence of a second, larger size class of
acanthostyle (which may be incorporated into
skeletal spicule tracts as well as echinating the
fibre endings and fibre nodes) is unusual to the
genus. Possession of hymedesmoid-plumose or
plumose skeletal architecture links several
microcionids termed the scabida' group (C. (D.)
scabida, C. (Microciona) si,nilis, C. (M.)
hentscheli, C. (M.) tetrastyla, C. (M.) thielei, C.
(Thalysias) tingens sp. nov. and C. (T) distincta).
Earlier records of this species (Carter, 1885a,
1886g; Hallmann, 1912) differentiate isochelae
microscleres into 3 or 4 morphs, but all isochelae
in both specimens are not as vastly different as
supposed by Carter (1885a). They are not arcuate,
as supposed by previous authors, but anchorate
with completely detached lateral alae and ridges
on the lateral sides of shaft. This is the only
REMARKS. Axociella is represented in
Australasian waters by six species, one new, all
of which have very well differentiated skeletal
structures closely resembling the compressed
skeletons common in Raspailidae. The existence
of microcionids with compressed axial skeletons
and differentiated axial and extra-axial skeletons
supports the proposal for a closer relationship
between Raspailiidae and Microcionidae, as
proposed by Hooper (1990a, 1991), and formalised further by Hajdu et al. (1994) in their
subordinal classification of Poecilosclerida. Crucial characters differentiating the Axociella group
from typical Raspailiidae are the possession of
chelae and absence any true echinating spicules
in the former, versus possession of a specialised
ectosomal skeleton (composed of small styles or
oxeas in brushes surrounding larger protruding
spicules) in the latter (see Hooper, 1991).
Clathria parva Levi (from S and SW Africa)
and Axosuberites fauroti Topsent (from the Gulf
of Aden), are also referred here to C. (Axociella)
because they have well differentiated axial and
extra-axial regions, whereas 3 NZ species
(Bergquist & Fromont, 1988) are retained in
this group only on a tentative basis since their
skeletal structures are not typical of Axociella.
Other species referred to Axociella e.g., de
Laubenfels, 1936a) are simply encrusting and/or
lack echinating acanthostyles (a secondary loss
common amongst Microcionidae and
Raspailiidae), and do not belong to Axociella as
defined here.
248zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 116. Clathria (Dendrocia) scabida (Carter) (NMVRN1038). A, Subectosomal auxiliary style/ subtylostyle.
B, 2 sizes echinating acanthostyle. C, Anchorate-like isochelae. D, Oxhom toxas. E, Section through peripheral
skeleton. F, Australian distribution. G, Holotype BMNH1887.7.11.9. H, NMVRN1025.
^
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249
'FIG. 117. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clat hr ia ( Dendr ocia) scabida (Carter) (NMVRN1025). A, Choanosomal skeleton. B, Peripheral
skeleton. C, Fibre characteristics. D, Echinating acanthostyles. E, Acanthostyle spination. F, Oxhorn toxas. G,
Anchorate isochelae. H, Uniguiferous anchorate isochelae.
MEMOIRS OF THE QUEENSLAND MUSEUM
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250zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathria (Axociella) canaliculata
(Whitelegge, 1906)
(Figs 118-119)
Esperiopsis canaliculata Whitelegge, 1906: 471,
p1.43, fig.7
Tenaciella canaliculata; Hallmann, 1920: 773, p1.36,
figs 1-2, p1.37, fig.1, text-fig.1; Ristau, 1978: 585586.
Artemisina canaliculata; Ristau, 1978: 585-586.
Rhaphidophlus canaliculata; Van Soest, 1984b: 115.
Clathriacanaliculata; Hooper & Wiedenmayer, 1994:
262.
MATERIAL. LECTOTYPE: AMG4325: Off Wata
Mooli, Bulgo, Woolongong, NSW, 34 °30'S,
151 ° 10'E, 104-126m depth, 01.iii.1900, coll. FIV
'Thetis' (dredge). PARALECTOTYPE: AMZ988: Off
Woolongong, NSW, 34°25'S, 151 ° 10'E, 220-224m
depth, 01.iii.1900., coll. FIV 'Thetis' (dredge).
OTHER MATERIAL: QLD- QMG300460 (fragment
NTMZ1562).
HABITAT DISTRIBUTION. Growing in soft sediments, associated with gorgonian beds in deeper waters
of the continental shelf; 94-126m depth; Wollongong
(NSW), S. Stradbroke I. (SEQ) (Fig. 118G).
DESCRIPTION. Shape. Erect, whip-like,
cylindrical branching digits, 150-190mm long,
with short cylindrical stalk 30-50mm long, 1015mm diameter, and few thick subcylindrical
branches, mostly in 1 plane, bifurcating and occasionally anastomosing.
Colour Drab grey in ethanol.
Oscules. Small, less than 2mm diameter, dispersed over lateral margins of branches.
Texture and surface characteristics. Firm,
flexible; surface smooth, fleshy, velvetty, feltlike, slightly porous, without obvious ornamentation.
Ectosome and subectosome. Thick, erect,
plumose brushes of ectosomal auxiliary subtylostyles form dense, continuous palisade on exterior
surface; discrete tracts of larger subectosomal
auxiliary styles form thick, multispicular, ascending or paratangential brushes supporting ectosomal skeleton.
Choanosome. Partially compressed, openmeshed reticulate axial skeleton and radial extraaxial skeleton, becoming plumose in periphery;
skeleton composed of very heavy spongin fibres
of large diameter, up to 250p,m diameter, forming
ovoid meshes, and clearly divided into primary
ascending and secondary connecting elements,
differing substantially in diameter; fibres cored
by multispicular tracts of choanosomal principal
styles occupying most of fibre diameter; spicule
content of fibres increases towards periphery;
secondary fibres less heavily cored; echinating
megascleres absent; mesohyl matrix moderately
heavywith scattered microscleres, but
megascleres largely confined within fibres except
at periphery.
Megascleres. Choanosomal principal styles
thick, slightly curved or rarely straight, fusiform,
with smooth, rounded or very slightly subtylote
bases. Length 130-(318.7)-465p,m, width 4(21.2)-26pm.
Subectosomal auxiliary styles long, thick,
straight, fiisiform, with tapering rounded or very
slightly subtylote bases, usually smooth, occasionally microspined. Length 2404465.0)590p,m, width 10-(12.2)-16p.m.
Ectosomal auxiliary subtylostyles straight or
slightly curved, with microspined subtylote
bases, fusiform points. Length 80-(114.6)165m, width 5-(6.1)-8p,m.
Microscleres. Palmate isochelae with long
lateral alae completely fused to shaft and fused
front ala; chelae clearly differentiated into two
size classes, the smaller often twisted. Length I:
4-(4.6)-8jun; length II: 14-(17.5)-22p.m.
Toxas accolada, usually long, thin, slight
central curvature and straight or only slightly
reflexed points. Length 184220.8 )-55011m,
width 0.541.9)-3.311m.
REMARKS. Hallmann (1920) erected Tenaciella
for this species on the basis that it lacked echinating spicules and had a Thalysias-like ectosomal
skeleton. Simpson (1968a) found no correlation
between these 2 features and histological
evidence to support recognition of this genus
distinct from Clathria. Spicule geometry and the
ectosomal skeleton in C. (A.) canaliculata indicate relationship with Thalysias, whereas skeletal
structure (particularly the radial choanosomal
skeleton), is reminiscent of Raspaila nuda
(Hooper, 1991). The shared skeletal structure in
Axociella and Raspailiidae support the contention
that they are monophyletic, belonging to the suborder Microcionina (Hajdu et al., 1994).
The well developed felt-like ectosomal spiculation and radial subectosomal spicule tracts in the
peripheral skeleton of C. (A.) canaliculata is virtually at the opposite end of a continuum from
encrusting Microciona-like species (with
membraneous dermal skeletons). These differences contrast so greatly with species such as C.
(M.) aceratoobtusa that it is tempting to maintain
generic separation between typical Microciona
and Thalysias-like species, but as demonstrated
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
251
elsewhere in this work there are too many other
microcionid species with intermediate conditions
making it impossible to clearly define a generic
boundary within this continuum.
Clathria(Axociella)canaliculata is most closely related to C. (A.) cylindrica, with similar
growth form, surface characteristics and skeletal
structure. However, this species has slightly compressed open-reticulate axis, radial extra-axis,
and continuous plumose ectosome, whereas C.
(A.) cylindrica has a greatly compressed closemeshed axis, a radial extra-axis and a sparse,
discontinuous, plumose ectosome. Toxa morphology and spicule size also differ substantially
between these two species.
CIathria (Axociella) cylindrica
(Ridley & Dendy, 1886)
(Fig. 120-121)
Esperiopsis cylindrica Ridley & Dendy, 1886: 340;
Ridley & Dendy, 1887: 79-80, p1.19, figs 2a-b.
Axociella cylindrica; Hallmann, 1920: 780-783, p1.37,
figs 2-4, text-fig.2;
Not Axociella cylindrica; Sim & Byeon, 1989: 39-40,
p1.5, figs 1-2.
Clathria cylindrica; Hooper & Wiedenmayer, 1994:
262.
Not Rhaphidophlus cylindricus Kieschnick, 1900: 53,
p1.44, fig.10.
MATERIAL. HOLOTYPE: BMNH1887.5.2.96: Off
Port Jackson, NSW, 33°51'S, 151 ° 16'E, 60-70m depth,
coll. HMS 'Challenger' (dredge). OTHER
MATERIAL: NSW- AMZ1527.
HABITAT DISTRIBUTION. Soft sediments; 40-70m
depth; Port Jackson, Botany Bay (NSW) (Fig. 1200).
DESCRIPTION. Shape. Thin, cylindrical
digitate, branching, whip-like, 130-190mm long,
10-20mm diameter, with thin, dichotomously
branched, cylindrical or slightly flattened, distally tapering branches, up to 130mm long, 16mm
diameter, short stalk and expanded basal attachment.
Colour Grey-brown in ethanol.
Oscules. Not seen.
Texture and surface characteristics. Firm,
flexible, tough consistency; surface even, feltlike, unornamented, prominently hispid.
Ectosome and subectosome. Sparse, plumose
brushes of small ectosomal auxiliary subtylostyles form discrete, discontinuous bundles on surface, arising from ends of radial skeletal columns,
scattered around the larger, protruding subectosomal auxiliary styles which project some distance through surface.
Choanosome. Skeletal structure with clearly differentiated axial and extra-axial components,
markedly compressed in axis and radial in extraaxis; compressed axial skeleton with longitudinal
fibres cored by short choanosomal principal subtylostyles, interconnected by thinner pauci- or
aspicular fibres; axial spongin fibres heavy, and
fibre anastomoses very close-meshed forming
elongate reticulation; mesohyl matrix in axis light
with abundant loose auxiliary styles dispersed
between and congregated around fibres; echinating megascleres absent; radial extra-axial
skeleton with large auxiliary styles perpendicular
to axis, forming pauci- or multispicular radial
(non-plumose) tracts associated with very light,
ascending spongin fibres but very few transverse
uni- or aspicular connecting fibres; mesohyl
matrix in extra-axial region moderately heavy,
with few scattered auxiliary megascleres.
Megascleres. Choanosomal principal subtylostyles coring axial fibres relatively short, slightly
curved at centre, with smooth subtylote or sometimes evenly rounded bases and fusiform points.
Length 215-(311.2)-395pm, width 5-(
Subectosomal auxiliary styles forming extraaxial bundles much longer and thicker than principal spicules, with smooth rounded or tapering
(hastate) bases, and fusiform points. Length 424(559.6)-725p.m, width 15-(21.0)-29p,m.
Ectosomal auxiliary subtylostyles straight,
variable length, with smooth rounded or slightly
subtylote bases and fusiform points. Length 208(361.8)-575p.m, width 6-(8.2)-10iim.
Microscleres. Palmate isochelae unmodified, differentiated into two size classes, with long lateral
alae entirely fused to shaft and completely fused
front ala. Length I: 6-(9.3)-13p.m; length ^19(22.3)-25p.m.
Toxas oxhorn, thick, with wide central curvature and slightly reflexed points. Length 45(86.2)-130pm, width 2.5-(4.3)-6p.m.
REMARKS. The chelae and toxa microscleres
show that this species belongs to the
Microcionidae, whereas skeletal architecture is
most closely related to the Raspailiidae (cf
Raspailia (Syringe/la) and Ectyoplasia). Like C.
(A.) canaliculata, this species lacks echinating
megascleres, and this was the primary reason why
Hallmann (1920) created Axociella. Despite contrary arguments by Van Soest (1984b), Axociella
is considered sufficiently different from
Thalysias in skeletal construction to be differentiated at the supraspecific level (although not for
252zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 118. Clathria (Axociella) canaliculata (Whitelegge) (lectotype AMG4325). A, Choanosomal principal
style. B, Subectosomal auxiliary style. C, Ectosomal auxiliary subtylostyle. D, Accolada toxas. E, Palmate
isochelae. F, Section through peripheral skeleton. G, Australian distribution. FI, Lectotype. I, QMG300460.
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253
FIG. 119. Clathria (Axociella) canaliculata (Whitelegge) (A, lectotype AMG4325; B-F, specimen
QMG300460). A, Choanosomal skeleton. B, Extra-axial fibre characteristics (x144). C, Base of ectosomal
auxiliary subtylostyle. D-E, Palmate isochelae. F, Accolada toxas.
254zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
A
a
FIG. 120. Clathria (Axociella) cylindrica (Ridley & Dendy) (AMZ1527). A, Choanosomal principal subtylostyle
(coring axial fibres). B, Subectosomal auxiliary style (coring extra-axial skeleton). C, Ectosomal auxiliary
subtylostyles. D, Oxhorn toxa. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian
distribution. H, Holotype BMNH1887.5.2.96.
the reason that it lacks echinating acanthostyles;
Simpson, 1968a). Supporting this interpretation
is the fact that a specialised Thalysias ectosomal
skeleton is absent in C. (A.) macropora and C.
(A.) thetidis. AMZ1133 from the F1V 'Thetis'
expedition, although not described by Hallmann
(1912), is very close to C. (A.) cylindrica differing
only in having echinating acanthostyles and long
thin, almost straight rhaphidiform toxas. This
material almost certainly represents a new
species but its antiquated condition is too poor to
serve as the holotype of a new taxon. Clathria
(Axociella) cylindrica also shows some
similarities to C. (Thalysias) procera in growth
form and having well differentiated axial and
extra-axial skeletons, but these species differ in
most other respects.
Axociella cylindrica of Sim & Byeon (1989)
from Korea does not compare to the Australian
population, differing markedly in most features,
and is referred here to Clathria (Microciona)
simae sp. nov.
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255
FIG. 121. Clathria (Axociella) cylindrica (Ridley & Dendy) (holotype BMNH1887.5.2.96). A, Choanosomal
skeleton. B, Fibre characteristics. C, Oxhorn toxas. D-E, Palmate isochelae. F, Cross-section through branch of
AMZ1527.
MEMOIRS OF THE QUEENSLAND MUSEUM
256^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathria (Axociella) nidificata
(Kirkpatrick, 1907)
(Figs 122-123)
Ophlitaspongia nidificata Kirkpatrick, 1907: 274;
Kirkpatrick, 1908b: 25, p1.22, fig.6, p1.24, fig.5;
Hallmann, 1912: 254; Burton, 1929a: 433; Burton,
1932a: 324; de Laubenfels, 1936a: 120; de Laubenfels, 1954: 162.
Axociella nidificata; Burton, 1940: 116; Koltun,
1964a: 70, p1.12,figs 7-11; Koltun, 1976: 155, 190;
Desqueyroux, 1975: 67-68, p1.4, figs 47-50;
Hoshino, 1977a: 45-46, table 1.
Clathria (Axociella) nidificata; Hooper & Wiedenmayer, 1994:263; Mothes & Lerner, 1995: 159-160,
figs 22-27, 55.
Ophlitaspongia flabellata Topsent, 1916: 167; Topsent, 1917: 41, p1.1, fig.4, p1.6, fig.2; Burton, 1929:
433; Burton, 1932: 325; Burton, 1934b: 34.
Axociella flabellata; Koltun, 1964a: 70-71, p1.12, figs
12-14; Hoshino, 1977a: 45-46, table 1.
Axociella rameus Koltun,1964a: 71, text-fig.17;
Hoshino, 1977a: 45-46, table 1.
MATERIAL. HOLOTYPE: BMNH1908.2.5. 131:
Coulman I., Victoria Land, Ross Dependency, Antarctica, 73°30'S, 17°00'E, 200m depth, coll. HMS
'Discovery' (dredge).
HABITAT DISTRIBUTION. On hard substrates; 93540m depth; Antarctica: BANZARE coast; Budd
Coast, Wilkes Land; Kemp Land, Enderby Land;
Mawson Peninsula, Oates Land; Coulman I., Victoria
Land; Prydz Bay, MacKenzie Bay and Mawson Coast,
MacRobertson Land; King Edward Ice Shelf, Enderby
Land; Joinville and Elefante Is; Palmer Archipelago;
Graham Land; also South Georgia, S. Shetland Is (Fig.
122E).
DESCRIPTION. Shape. Erect, claviform-flabellate, narrow stalk.
Colour Yellow-brown in ethanol.
Oscules. Oscules on upper surface of club, I 2mm diameter.
Texture and surface characteristics. Texture
compressible, tough, harsh surface; surface
hispid, conulose, membraneous between conules.
Ectosome and subectosome. Ectosomal skeleton
plumose, composed of discrete bundles of subectosomal auxiliary subtylostyles perched on ends
of protruding principal spicules, corresponding to
surface conules.
Choanosome. Skeleton compressed in axis and
plumose in extra-axis; axial region slightly compressed containing heavy spongin fibres cored by
plumo-reticulate pauci- or multispicular tracts of
choanosomal principal styles; skeletal tracts form
oval meshes, up to 250p.m diameter; extra-axial
tracts plumose, diverging towards periphery, with
pauci- or multispicular tracts of choanosomal
principal styles coring heavy spongin fibres;
styles in peripheral regions of fibres often
protrude from fibres at acute angles, but these are
identical to principal megascleres and cannot be
considered as true echinating spicules; mesohyl
matrix moderately heavy, granular, containing
abundant toxas, often in bundles, and occasional
detritus.
Megascleres. Choanosomal principal styles long,
thick, entirely smooth, slightly curved towards
base, with slightly subtylote evenly rounded or
slightly constricted hastate bases and fusiform
points. Length 595-(914.7)-1140p-m, width 21(43.7)-61p.m.
Subectosomal auxiliary subtylostyles short,
slender, straight, with slightly subtylote
microspined bases, fusiform pointed. Length
331-(405.4)-447p.m, width 6-(9.4)-12p.m.
No echinating spicules.
Microscleres. Chelae absent.
Toxas accolada, thick, variable in length, with
prominent but narrow angular central curvature,
straight arms and straight or slightly reflexed tips;
larger toxas with spined tips, small toxas entirely
smooth. Length 98-(286.2)-546p.m, width 1.5(3.2)-5p.m.
REMARKS. The synonymy given above follows
Koltun (1976). The species appears to vary in
growth form, from bushy, branching, reticulate,
digitate to flabellate, but skeletal architecture and
spicule geometry are relatively consistent.
Several echinating acanthostyles were seen in
spicule preparations made from the holotype
(evenly spined, club-shaped, 94-123x5-9Rm),
but none were seen in section preparations and it
is presumed that these are contaminants, although
it is possible that they are present but rare.
The specialised ectosomal skeleton of discrete
auxiliary spicule brushes perched over larger
principal spicules, and compressed `axinellid'
skeletal construction, are reminiscent of
Raspailiidae (e.g., Aulospongus), and were it not
for the toxas in this species it could be included
in Raspailiidae. This species fits best in Clathria
(Axociella) although it lacks differentiated ectosomal and subectosomal spicules (i.e., it has 1
size class of auxiliary styles). Its toxa and spination should be compared with C. (A.) georgiaensis sp. nov. (see below).
Clathria (Axociella) patula sp. nov.
(Figs 124-125)
MATERIAL. HOLOTYPE: NTMZ2909: Between
North and East Wallabi Is, Houtman Abrolhos, WA,
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
257
FIG. 122. Clathria (Axociella) nidificata (Kirkpatrick) (holotype BMNH1908.2.5.131). A, Choanosomal principal styles. B, Subectosomal auxiliary subtylostyles. C, Accolada toxas. D, Section through peripheral skeleton.
E, Antarctic distribution.
28°17.9'S, 113°47.8'E, 11.vii.1987, 39m depth, coll.
J.N.A. Hooper (beam trawl). PARATYPE:
NTMZ2978 (fragment QMG300209): W. of Carnarvon, WA, 24 ° 55.6'S, 112°50.8'E, 14.vii.1987, 85m
depth, coll. J.N.A. Hooper (beam trawl). OTHER
MATERIAL. WA- QMG304633.
HABITAT DISTRIBUTION. Broken limestone reef,
usually in sand sediments; 39-85m depth; Camarvon
and Wallabi Is, Houtman Abrolhos (WA) (Fig. 124G).
DESCRIPTION. Shape. Thickly flabellate, palmate-digitate fan, 140-475mm long, 180-310mm
^ MEMOIRS OF THE QUEENSLAND MUSEUM
258zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Ectosome and subectosome.
Thick plumose or paratangential layer of erect auxiliary
styles, usually in thick bundles,
forming more-or-less continuous palisade on ectosomal
skeleton; subectosomal
skeleton with plumose brushes
of both subectosomal auxiliary
styles and choanosomal principal styles supporting ectosomal palisade, arising
directly from choanosomal
fibres in peripheral skeleton.
Choanosome. Skeletal structure wide-meshed heavily
reticulate, slightly compressed
axial region and differentiated
axial and extra-axial skeletons;
axial skeleton composed of
heavy spongin fibres, 80145p,m diameter, thickest and
bulbous at fibre nodes, 130200p,rn diameter, forming relatively wide reticulation of oval
or rectangular meshes, 180360Rm diameter; axial fibres
cored by both principal styles
and subectosomal styles in
multispicular brushes of 2-5
spicules; principal spicules
also protrude through fibres in
FIG. 123. Clathria (Axociella) nidificata (Kirkpatrick) (holotype plumose bundles, at acute
BMNH1908.2.5.131). A, Plumose extra-axial skeletal column. B, Plumo- angles, although these cannot
reticulate axial skeleton. C, Bundles of toxas.
be considered as echinating
spicules; extra-axial skeleton
with wider fibre meshes than
wide, 6-15mm thick, with even apical margin or axial region, with heavy spongin fibres, 60long, erect flattened digits arising from apex of
12011m diameter, bulbous fibre nodes, 80-140p.m
fan; digits single or planar branching, 40-70mm
diameter, forming elongate or rectangular
wide, bifurcated and tapering, with short, thick
cylindrical stalk, 35-63mm long, 15-32mm meshes, 300-470Rm diameter, cored by ascenddiameter; lateral margins of fans prominently ing multispicular tracts of intermingled
crenellated, apical margins range from slightly choanosomal principal and subectosomal styles,
up to 10 spicules per fibre, and thinner transverse
crenellated to prominently digitate.
fibres cored by only principal spicules in
Colour Dark red, red-brown or orange-brown paucispicular tracts; spicules usually protrude
alive (Munsell 5R 5/10 - 1OR 7/8), grey-brown in through fibres in plumose bundles, particularly in
ethanol.
peripheral skeleton, but true echinating spicules
Oscules. Small oscules, less than 2mm diameter, absent; mesohyl matrix heavy but only lightly
scattered over surface.
pigmented, containing abundant microscleres;
Texture and surface characteristics. Firm, com- choanocyte chambers large, oval, 50-1301.Lm
pressible, flexible; surface uneven, rippled, diameter.
slightly conulose, with few large digitate projec- Megascleres. Choanosomal principal styles varitions arising at oblique angles from surface in fan able in length, predominantly long, thick, slightly
specimens.
curved near basal end, evenly rounded smooth
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
259
FIG. 124. Clathria (Axociella) patula sp.nov. (holotype NTMZ2909). A, Choanosomal principal style. B,
Subectosomal auxiliary styles. C, Ectosomal auxiliary styles. D, Wing-shaped - oxhorn toxa. E, Palmate
isochelae. F, Section through peripheral skeleton. G, Known Australian distribution. H, Holotype. I, Paratype.
MEMOIRS OF THE QUEENSLAND MUSEUM
260^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 125. Clathria (Axociella) panda sp.nov. (holotype NTMZ2978). A, Choanosomal skeleton. B, Fibre
characteristics. C-E, Bases of choanosomal principal, subectosomal auxiliary and ectosomal auxiliary spicules.
F, Palmate isochelae. G, Oxhorn - wing-shaped toxas.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
^
261
bases and fusiform points. Length 2254383.6)370p,m, width 10413.2)-1*m.
Subectosomal auxiliary styles long, thinner
than principal spicules, straight, with rounded
microspined bases, slightly subtylote spined
bases, or subtylote smooth bases, and fusiform
points. Length 1934265.2)-30311m, width 5(7.1)-8p.m.
Ectosomal auxiliary styles short, straight or
slightly curved near base, relatively thick, with
rounded smooth or microspined bases and
fusiform points. Length 104-(147.2)-185Rm,
width 1.5-(3.8)-6Rm.
Echinating spicules absent.
Microscleres. Palmate isochelae small, unmodified, with lateral and front alae of approximately equal length; lateral alae fused
completely to shaft, front ala entire. Length 14(15.8)-19pm.
Toxas basically oxhorn, with some intermediate wing-shaped, short, moderately thick,
rounded or slightly angular central curvature and
slightly reflexed points 57-(69.6)-84Rm, width
141.9)-2.511m.
Woolongong, NSW, 34 ° 30'S, 151 ° 12'E, 98-138m
depth, coll. FIV 'Thetis' (dredge).0THER
MATERIAL: NSW- QMG303752. S. AUSTAMZ4624.
HABITAT DISTRIBUTION. Deeper offshore reefs;
50-142m depth; Sydney and Woolongong (NSW),
SW. Great Australian Bight (SA) (Fig. 126H).
DESCRIPTION. Shape. Erect, arborescent or
digitate, whip-like growth forms, 190-480mm
long, with long thin cylindrical stalk and thinly
cylindrical branches, 15-25mm diameter;
branching planar or in more than one plane, either
irregularly bifurcate or pinnate, with evenly
dichotomous bifurcations arising from larger,
laterally disposed branches; apical margins of
digits taper to fine points.
Colour. Deep red alive (Munsell 2.5R 5/10), greybrown when dry.
Oscules. Very small, less than 1.5mm diameter,
scattered over lateral sides of branches.
Texture and surface characteristics. Firm,
flexible; surface optically even, velvetty, microscopically prominently furry, hispid.
Ectosome and subectosome. Ectosome very
ETYMOLOGY. Latin zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
pat ulus, spread out; for the flat
hi
spid, with discrete, multispicular plumose
growth form.
REMARKS. This species could be included in C. brushes of larger (extra-axial) principal styles
(Thalysias), having a thick tangential ectosomal protruding through surface, with sparsely disskeleton composed of two discrete size classes of persed, tangential or paratangential tracts of subauxiliary styles, but it is most appropriately ectosomal auxiliary styles in between principal
placed in C. (Axociella) given that it has differen- spicules.
tiated axial and extra-axial skeletons and lacks Choanosome. Skeletal architecture reticulate,
any echinating spicules. C. (Axociella)patula has with compressed axis and plumose extra-axial
a flabellate, planar growth form similar to C. (T) regions; strong axial compression composed of
cancellaria, although these species differ sub- irregularly reticulate, heavy, large spongin fibres
stantially in their spicule geometry (the latter producing oval to elongate meshes, 90-220p.,m
having echinating acanthostyles and accolada diameter; axial fibres relatively heavy, thick, irtoxas), different spicule sizes of most categories, regularly anastomosing, with bulbous fibre
and skeletal structures (the latter species with a nodes, cored by comparatively small choanoheavy, evenly reticulate skeleton and abundant somal styles; axial fibres incompletely differentiated into primary, uni- or paucispicular,
echinating acanthostyles).
ascending, long, radial fibres, 70-1801.im
Clathria (Axociella) thetidis (Hallmann, 1920) diameter, and secondary, connecting, mostly
unispicular, sometimes bi- or aspicular, short
(Figs 126-127, Plate 4B)
fibres, 58-102p,m diameter. Axial and extra-axial
skeletons strongly differentiated (cf. Hallmann,
Ophlit aspongia t het idis Hal!mann, 1920: 779.
1920), with fibres becoming plumose and fibre
Axociella t het idis; de Laubenfels, 1936a: 113.
Clat hria t het idis; Hooper & Wiedenmayer, 1994: 263
reticulation more regular towards periphery;
Esperiopsis cylindrica, in part; Whitelegge, 1906: 470,
extra-axial fibres form elongate-eliptical, wider
p1.43, fig.6.
meshes than axis, 155-510Rm diameter; extraNot Esperiopsis cylindrica Ridley & Dendy, 1887: 79. axial fibres diminish in thickness towards
peripheral skeleton, whereas coring spicules inMATERIAL. LECTOTYPE: AMG9199: Off
Woolongong, NSW, 34 ° 25'S, 151 ° 10'E, 104-142m crease in size and density towards surface; principal styles in both primary and secondary fibres
depth, coll. FIV 'Thetis' (dredge). PARALECprotrude through fibres at oblique angles, in both
TOTYPE: AMG9191: Off Wata Mooli, Bulgo,
^ MEMOIRS OF THE QUEENSLAND MUSEUM
262zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 126. Clathria (Axociella) thetidis (Hal'mann) (holotype AMG9199). A, Choanosomal principal style (axial
skeleton). B, Choanosomal principal style (extra-axial skeleton). C, Subectosomal auxiliary subtylostyle. D,
Oxeote toxa. E, Wing-shaped toxas. F, Palmate isochelae. G, Section through peripheral skeleton. H, Australian
distribution. I, Holotype.
the axial and extra-axial regions, but true echinat- 11-(14.2)-20p,m), and larger in extra-axial
ing spicules absent; mesohyl matrix heavy, rela- skeleton (length 377-(608.0)-825iim, width 22tively darkly pigmented; choanocyte chambers (26.4)-35p.m.
paired, oval-elongate, 60-1541,m diameter;
Subectosomal auxiliary styles long, thin,
numerous microscleres and subectosomal fusiform, straight, slightly curved or flexuous,
with subtylote, smooth or lightly rnicrospined
auxiliary subtylostyles scattered between fibres.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGF
bases. Length 158-(379.1)-585p,m, width 3Megascleres. Choanosomal principal styles
divided into two size categories, both fusiform, (5.8)-811m.
thick, long or short, slightly curved, with evenly Microscleres. Palmate isochelae abundant, with
rounded, smooth bases; smaller size found in lateral alae completely attached to shaft and front
axial skeleton (length 1804246.5)-365 pm, width ala entirely fused. Length 9413.4)-1611m.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
263
FIG. 127. Clathria (Axociella) thetidis (Hallmann) (QMG303752). A, Choanosomal skeleton. B, Extra-axial
fibre and spicule skeleton (x63). C, Palmate isochelae. D, Wing-shaped toxas. E, Oxeote toxas.
264zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Toxas divided into two geometrically different
forms: oxeote toxas common, usually found in
dragmata dispersed between fibres, long, thin,
symmetrical or asymmetrical-sinuous, with
slight angular central curvature or completely
straight, fusiform points. Length 175-(774.5)1280p,m, width 1.542.4)-5 p.m; wing-shaped
toxas common, thin, with large rounded central
curvature, reflexed points. Length 224104.4)168p.,m, width 142.1)-3.41,m.
REMARKS. This species is not conspecific with
Esperiopsis cylindrica Ridley & Dendy as supposed by Whitelegge (1906), having different
ectosomal skeletal architecture, spicule
geometries and spicule sizes; Hallmann (1920)
recognised a new species for Whitelegge's
material referring it to Ophlitaspongia (=
Echinoclathria) on the basis that it lacked
echinating megascleres; de Laubenfels (1936a)
assigned it to Axociella for the same reason. De
Laubenfels was correct in this transfer, although
not for his stated reason (that it lacked acanthostyles) but because of its compressed skeletal construction, spicule localisation and spicule
geometry. Hallmann's tentative placement of C.
(A.) thetidis in Echinoclathria was probably also
based on comparisons with species such as E.
nodosa (Carter) and to a lesser extent E. subhispida (Carter) which, unlike most species of
Echinoclathria, have a slightly compressed axial
skeleton and plumose, plumo-reticulate or radial
extra-axial fibres. However, spiculation and
spicule localisation within the skeleton of C. (A.)
thetidis is different from all those species.
Similarly, in C. (A.) thetidis there are two distinctive size categories of principal megascleres,
restricted to either axial or extra-axial fibre
skeletons, whereas Echinoclathria have
homogeneous principal spicules dispersed
throughout all skeletal tracts.
Clathria (Axociella) georgiaensis sp.nov.
(Figs 128-129)
Ophlitaspongia thielei Burton, 1932a: 322, p1.55, fig.8,
text-fig.32; Koltun, 1964a: 70.
Axociella thielei; de Laubenfels, 1936a: 113 [note].
Not Hymeraphia thielei Hentschel, 1912: 377-378.
MATERIAL. HOLOTYPE: BMNH1928.2.15. 219
(fragment AMZ2198): 6.3nm N 89 ° E to 4nm N 39 ° E
off Jason Light, Cumberland Bay, South Georgia, S.
Atlantic, 120-204m depth, RRS 'Discovery', 1928
(otter trawl).
HABITAT DISTRIBUTION. On rocks; 18-236m
depth range; Wilkes Land, Australian Antarctic Territory (Fig. 128F); also South Georgia, SW. Atlantic.
DESCRIPTION. Shape. Subspherical, massive
sponge.
Colour Grey-brown in ethanol.
Oscules. Up to 2mm diameter, scattered over
surface, with slightly raised membraneous lip.
Texture and surface characteristics. Firm, compressible; surface conulose, with meandering
ridges producing a clathrous, convoluted mass.
Ectosome and subectosome. Erect plumose
brushes of sparse choanosomal principal styles
protruding from peripheral skeletal tracts, and
paratangential bundles of subectosomal auxiliary
subtylostyles in variable abundance, heavier on
ends of surface conules, lighter between conules.
Choanosome. Skeletal architecture reticulate,
slightly plumo-reticulate near surface, vaguely
renieroid reticulate at core; skeleton with differentiated primary ascending spongin fibres
cored by multispicular tracts of choanosomal
principal styles, with 3-6 spicules per tract, and
lighter transverse connecting fibres containing
1-2 spicules per tract, together producing a slightly renieroid skeleton; no marked differentiation
between axial or extra-axial regions; fibre
reticulation produces cavernous rectangular or
triangular meshes, up to 550p..m diameter; true
echinating spicules absent although principal
spicules protrude through fibres at obtuse angles;
mesohyl matrix heavy, slightly granular, with
abundant chelae and toxas.
Megascleres. Choanosomal principal styles
moderately long, slender, straight or slightly
curved at centre, rounded smooth bases, fusiform
points. Length 3904446.6)-51811m, width 14(17.4)-2211m.
Subectosomal auxiliary subtylostyles short,
slender, straight, slightly subtylote bases, heavily
microspined with large spines, slightly hastate or
rounded points. Length 216-(285.2)-348itm,
width 5-(7.4)-10Rm.
Echinating spicules absent.
Microscleres. Palmate isochelae abundant, relatively small, with lateral and front alae approximately same length, lateral alae fused to
shaft, front alae nearly completely detached, virtually no curvature of shaft. Length 9-(l3.2)17 p.m.
Toxas wing-shaped, thick, variable in length,
the thicker ones with wide angular central curvature, curved arms, slightly reflexed points, and
terminal spines, thinner ones more sharply curved
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
265
FIG. 128. Clathria (Axociella) georgiensis sp. nov. (holotype BMNH1928.2.15.219). A, Choanosomal principal
styles. B, Subectosomal auxiliary subty losty les. C, Wing-shaped toxas. D, Palmate isochelae. E, Section through
peripheral skeleton. F, Antarctic distribution. G, Peripheral spicule tracts. H, Ectosomal skeleton.
at centre, sharply pointed. Length 284107.7)258p.m, width 0.541.7)-3.511m.
REMARKS. This species requires a new name
since C. (Microciona) thielei (Hentschel, 1912)
has seniority. Previous authors could not agree on
its generic assignment: Burton (1932a, 1938b)
compared C. (A.) georgiaensis with Artemisina,
but the possession of a well-structured, slightly
compressed choanosomal skeleton indicates that
266zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 129. Clathria (Axociella) georgiaensis sp. nov. (AMZ2198). A, Choanosomal skeleton. B, Fibre characteristics. C, Choanosomal principal style. D, Base and apex of principal spicule. E, Base and apex of
subectosomal auxiliary subtylostyle. F, Palmate isochela. G, Wing-shaped toxas. H, Spined toxa point.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
^
267
placement in Clathria is more appropriate. Based
on Burton's (1932a) description this species was
included in Echinoclathria by Hooper & Wiedenmayer (1994), but re-examination of the holotype
showed that skeletal structure was not markedly
renieroid, there was no difference between principal spicule sizes in the axis and those in the
peripheral skeletons, and spicule sizes were magnitudes larger than those found in other
Ecinoclathria (but much closer to those seen in
Axociella). De Laubenfels (1936a) also suggested this species should be included in Axociella although his reasons for doing so are different
(i.e., lack of echinating spicules). Koltun (1964a)
also made a general comparison between C. (A.)
georgiaensis and Ophlitaspongia membranacea
Thiele (the latter referred here to Clathria
(Thalysias)), but this comparison is simply based
on similarities in toxa morphology, whereas C.
(T) membranacea has two size classes of
auxiliary spicules and true echinating spicules
(albiet smooth). Toxa measurements recorded by
Burton (1932a) for this species differ substantially from those actually seen in the holotype, but in
most other characters his description is an accurate representation of the species.
OTHER SPECIES OF CLATHRIA
(AXOCIELLA)
Clathria (Axociella) fauroti (Topsent, 1893)
Axosuberites fauroti Topsent, 1893a: 179-181, fig.3 [Tadjoura. Gulf of Aden]; de Laubenfels, 1936a: 118.
? Rhaphidophlus fauroti; Van Soest, 1984b: 130 [possible
generic synonymy].
MATERIAL. HOLOTYPE: MNHNDT1859. Arabian GulfRed Sea.
Clathria (Axociella) fromontae sp. nov.
Axociella toxitenuis Bergquist & Fromont, 1988, 117-118,
p1.55, figs a-c; Dawson, 1993: 36 [index to fauna].
MATERIAL. HOLOTYPE: NMNZPOR120. New Zealand.
Note: C. toxitenuis Topsent, 1925 has seniority.
Clathria (Axociella) lambei (Koltun, 1955)
Microciona lambei Koltun, 1955a: 49, 67, p1.4, fig.5 [Japan
and Okhotsk Seas]; Koltun, 1958: 66-67, text-fig.22
[Kuriles]; Koltun, 1959: 183, p1.29, fig.2, text-fig.143
[USSR].
MATERIAL. HOLOTYPE: ZIL, fragment
BMNH1932.11.17.67. NW. Pacific, Japan. Note: Koltun
(1955a) attributes this species to Burton (1935c), but it does
not appear in that publication.
Clathria (Axociella) macrotoxa Bergquist &
Clathria (Axociella) multitoxaformis
Bergquist & Fromont, 1988
Axociella multitoxaformis Bergquist & Fromont, 1988: 118119, p1.55, figs d-f, p1.56. fig.a; Dawson, 1993: 36 [index
to fauna].
MATERIAL. HOLOTYPE: NMNZPOR121. New Zealand.
Clathria (Axociella) parva (Levi, 1963)
Clathria parva Levi, 1963: 56-57, text-fig. 64, p1.10D [Cape
of Good Hope, South Africa]; Uriz, 1988a: 84-85, p1.21b,
text-fig.60 [Namibia].
Not Clathria parva; Sim & Byeon, 1989: 39, pI4, figs 3-4
[Korea; dubious conspecificity].
MATERIAL. HOLOTYPE: UCT (fragment
MNHNDCL612). S and SE Africa.
Clathria (Isociella) Hallmann, 1920
Isociella Hallmann, 1920: 784; Bergquist & Fromont,
1988: 114.
DEFINITION. Relatively homogeneous isodictyal (triangular meshes) and/or renieroid (rectangular meshes), wide-meshed, main skeleton with
primary plumose ascending, multispicular tracts
cored by smooth choanosomal styles, interconnected by secondary, uni- or paucispicular tracts
cored by ,same spicules, and choanosomal
spicules sometimes diverging and forming
plumose brushes at surface; echinating
megascleres absent; ectosomal skeleton with
single category of auxiliary spicule, tangential,
paratangential. Microscleres palmate-derived
isochelae and toxas.
TYPE SPECIES. Clathria macropora, Lendenfeld,
1888: 221 (=Phakelliajacksoniana Dendy, 1897:236)
(by monotypy).
REMARKS. Four species of Isociella are known
for the Australian fauna, three from tropical WA,
NT and Qld., and one temperate species from
NSW. Only one other species is known from New
Zealand.
Clathria (Isociella) eccentrica (Burton, 1934)
(Figs 130-131, Table 26, Plate 4A)
Ophlitaspongia eccentrica Burton, 1934a: 560, pls
1,8,9, text-fig.12a.
Axociella eccentrica; de Laubenfels, 1936a: 113.
Isociella eccentrica; Bergquist & Tizard, 1967: 186187, p1.5, fig. 1.
Clathria eccentrica; Hooper & Wiedenmayer, 1994:
265.
Fromont, 1988
MATERIAL. HOLOTYPE: BMNH1930.8.13. 109:
Axociella macrotoxa Bergquist & Fromont, 1988: 117, p1.54,
figs c-f; Dawson, 1993: 36 [index to fauna].
MATERIAL. HOLOTYPE: NMNZPOR119. New Zealand.
Crab Spit, Low Isles, Great Barrier Reef, Qld, 16 °23'S,
145°34'E, intertidal, 5.iv.1929, coll. Great Barrier Reef
Expedition (dredge). OTHER MATERIAL: GREAT
^ MEMOIRS OF THE QUEENSLAND MUSEUM
268zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
A
0
FIG. 130. Clathria (Isociella) eccentrica (Burton) (NTMZ2170). A, Choanosomal principal styles. B-C, Subectosomal
auxiliary styles. D, Wing-shaped toxas. E, Palmate isochelae. F, Section through peripheral skeleton. G, Australian
distribution. H, Holotype BMNH1930.8.13.109. I, NTMZ2139.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
269
FIG. 131. Clathria (Isociella) eccentrica (Burton) (QMG303266). A, Choanosomal skeleton. B, Fibre characteristics (x135). C-D, Bases of larger and smaller subectosomal auxiliary styles. E, Wing-shaped toxas. F,
Palmate isochelae.
270zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 26. Comparison between present and published descriptions of Texture and surface characClaihria (Isociella) eccentrica.Measurements in p.m (N=25). teristics. Rubbery, easily com-
pressible, mucusy, but tough,
difficult to tear; surface
SPICULE
porous, membraneous,
(GBR,Queensland)
minutely microconulose with
2324361.7)-540 276-(343.1)-450
Choanosomal
318-464 x 8-22 288-440 x 9.1-18
protruding fibre nodes from
styles
x7-(17.4)-30
x 11-(19.2)-29
peripheral skeleton, with
2574335.6)-440 284-(373.3)-455
Subectosomal
298-457 x 3-9.5
300-470 x 5-8
small ridges and depressions
styles I
x 3-(6.8)-13
x 4-(6.3)-9
connecting adjacent conules;
141-(191.8)-255
85-(185.0)-255
Subectosomal
85-286 x 1.5-5.2
208-220 x 4.0
surface usually silt covered.
styles II
x 1-(3.5)-7
x 2-(3.7)-6
Ectosome and subectosome.
15-19
16-17.5
8-(17.4)-23
13-(17.2)-21
Chelae
Ectosomal skeleton micros64-(161.2)-292
16-(148.0)-283
10-334 x 1.5-10 76-270x 2.5-6.5
Toxas
copically hispid, with
x 1-(5.4)-11
x 2-(6.0)-11
choanosomal principal styles
protruding up to 300p,m from
BARRIER REEF, QLD - QMG304398, QMG304400,
surface, at regular intervals 150-400p.m apart,
QMG304401. DARWIN REGION, NT - AMZ3109,
NTMZ0268, NTMZ0288, NTMZ0158, NTMZ0159, singly or in paucispicular bundles of 2-4 spicules,
with peripheral spongin fibres forming projecNTMZ2116, NTMZ2139, NTMZ2205, NTMZ2210,
QMG300147 (fragment NTMZ2224), QMG300509 tions and enclosing at least basal portion of
(fragment NTMZ2235), NTMZ2416, NTMZ2540, protruding spicules; subectosomal auxiliary
QMG303315, NTMZ2549, NTMZ2557, NTMZ1100,
styles form tangential or paratangential tracts at
NTMZ0386, NTMZ2170, QMG303266, base of peripheral skeleton, interdispersed beNTMZ0045, NTMZ1396, NTMZ3274. AMG4291.
tween choanosomal principal styles of subrenieroid skeleton; auxiliary spicules sparsely
HABITAT DISTRIBUTION. Usually on dead or pardispersed, rarely protruding through surface,
tially dead faviid coral heads, coral rubble, sand and composed of 2 sizes of auxiliary styles without
Halimeda substrates; most specimens partially sheltered under coral rubble or in crevices; shallow sublit- regional localisation; peripheral skeleton undiftoral distribution; 0-18m depth range; Lizard I. (FNQ), ferentiated from choanosomal skeleton, and
Darwin Harbour, Bynoe Harbour, Trepang Bay, Port choanosomal fibres immediately subdermal.
Essington, Cobourg Peninsula (NT) (Fig. 1306); also Choanosome. Irregular subisodictyal, or in places
SE. Indonesia (unpublished data).
more regular isodictyal, sometimes renieroid
reticulation of moderately light spongin fibres,
DESCRIPTION. Shape. Bulbous-digitate, mas- 30-110v1,m diameter, without any obvious or consive or semi-encrusting, with irregularly anas- sistent differentiation between primary or secontomosing, erect or stoloniferous branches; dary elements; fibres cored by 1-10 rows of
branches irregularly cylindrical, flattened or bul- choanosomal principal styles entirely enclosed
bous; specimens range from thick encrustations within fibres, occupying entire fibre diameter;
45mm high, 20-30mm diameter, with few bul- spicules diverge only slightly towards periphery
bous branches on surface, to massive branching whereas at core of skeleton spicules more evenly
growth forms up to 130mm high, 300mm wide, dispersed within fibres; echinating megascleres
with branch diameter between 15-40mm.
absent; fibre anastomoses form irregular, elonColour Dark red (Munsell 5R 3/8), orange-red gate-oval, triangular or sometimes regularly rec(5R 6/10), or less commonly bright orange (10R tangular meshes, 250-850Rm diameter; mesohyl
6/10); pigmentation water miscible, associated matrix light, granular, with abundant
with mucous, confined to the ectosomal and sub- microscleres and auxiliary spicules; collagen
ectosomal regions; pigment washed from sponge; heaviest at fibre nodes but sparse elswhere;
subectosomal colouration light brown or beige; choanocyte chambers oval or elongate, 60150p,m diameter.
grey-brown in ethanol.
Megascleres
(Table 26). Choanosomal principal
Oscules. Large, 4-15mm in diameter, on apex of
bulbous digits, usually at extremities of branches; styles straight or slightly curved near base, with
oscules with slightly raised membraneous lips, smooth rounded or very slightly subtylote bases,
collapsing upon dessication and preservation; fusiform points.
numerous inhalant pores, 0.5-1.0mm diameter,
Subectosomal auxiliary subtylostyles thin,
scattered over entire surface.
usually straight, either with slightly subtylote or
Holotype (BMNH
1930.8.13.109)
Bergquist &
Tizard (1967)
(Danvin,NT)
(19=19)
(Darwin,NT)
(N=3) (Cobourg
Pen.,NT)
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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271
rounded bases, and microspined or smooth bases
in approximately equal proportions, and with
fusiform points; 2 size categories recognised, although undifferentiated in distribution.
Echinating spicules absent.
Microscleres (Table 26). Palmate isochelae abundant, relatively large, unmodified, with long
lateral alae completely fused to shaft and entirely
fused front ala.
Wing-shaped toxas variable length, relatively
thick, with moderate, rounded central curvature,
slightly reflexed or straight points; some oxeote
toxas also present but rare; toxas frequently occur
in dragmata within mesohyl.
Associations. Sometimes smothering live faviid
coral heads, produced localised bleaching and
necrosis of coral tissues at the point of contact
with sponge; probable that this species involved
in chemical bioerosion of coralline substrate;
most specimens (81%) contained scyllid
polychaete worms.
Morphological variation. In growth form, live
colouration, surface features, skeletal construction and spiculate geometry this species shows
very little apparent variation. Some variation observed for: Detritus incorporated into ectosomal
skeleton: abundant (33%), lightly dispersed particles (24%), entirely clear of detritus (43% of
specimens). Ectosomal skeleton: strictly tangential (43%), disorganised paratangential (33%), or
with both structures on different parts of the surface (24% of specimens). Choanosomal skeleton:
characteristically subisodictyal (82%), regularly
isodictyal (9%), or regular renieroid construction
(9% of specimens). Fibre meshes: cavernous
(86%), or close-meshed skeletal reticulation
(14% of specimens). Mesohyl matrix: relatively
heavy, darkly pigmented (19%), heavy, lightly
pigmented (57%), or very light, unpigmented
(24% of specimens). Detritus incorporated into
mesohyl: seen in only 14% of specimens. Abundance of microscleres: very abundant in tracts or
irregularly dispersed throughout the mesohyl
(67%), or uncommon (33% of specimens).
Auxiliary styles dispersed between fibres within
mesohyl: numerous (43%), moderate (19%) or
sparse extra-fibre tracts (38% of specimens).
Spicule dimensions: spicule dimensions varied
considerably between specimens, but this variation was not explained by either geographic distribution or seasonality of collections.
REMARKS. This species is a distinctive component of the tropical Australian intertidal fauna,
although it is only known from two disjunct loca-
tions (the 'Top End' of the NT and Cairns region,
Qld.). No intermediate populations have yet been
discovered despite major collections undertaken
recently in that region.
This species is well characterised by its
haplosclerid-like, predominantly isodictyal
skeletal construction, relatively poor development of the extra-fibre skeleton (including the
cavernous fibre meshes and relatively light
mesohyl matrix), and spicule geometries. The
species is also distinctive in the field, mainly by
its predominantly bright red colouration and
abundant mucus. It has the ability to survive
extensive periods of exposure to direct sunlight
and air, for up to six hours duration (several times
each week during ELWS tides in the Darwin
region), and to water temperatures in rockpools
which may exceed 40°C. Under such conditions
C. (I.) eccentrica produces copious quantities of
mucus, literally dripping its red pigmentation.
Nevertheless, individuals appear to survive these
harsh conditions, and necrotic areas of the surface
are relatively quickly regenerated or recolonised
within several weeks. Despite extensive monitoring of individuals in the Darwin region over many
seasons, reproductive products have not yet been
recorded, and it is possible that its propogation in
the tropics is predominantly clonal.
Clathria (Isociella) macropora Lendenfeld, 1886
(Figs 132-133, Table 27)
Phakelhaflabellata Ridley & Dendy, 1886: 478; Rid-
ley & Dendy, 1887: 171, p1.34, figs 2-3, p1.40, fig.6
[preocc.1.
Not Phakelliaflabellata Carter, 1885f: 363.
Clathria macropora, in part, Lendenfeld, 1888: 221;
Hallmann, 1920: 768; Bergquist & Fromont, 1988:
110; Hooper & Wiedenmayer, 1994: 265.
Not Plectispa macropora Lendenfeld, 1888: 226;
Hallmann, 1912: 203, 205, 242.
Not Clathria macropora; Whitelegge, 1901: 91.
Not Wilsonellamacropora; Hallmann, 1912: 203, 240,
242, 205.
Not Plumohalichondria australis Whitelegge, 1901:
90, p1.11, fig.14.
Isociella flabellata; Hallmann, 1920: 784-789, p1.39,
figs 1-2, p1.40, fig.1, text-fig.3.
Phakelliajacksoniana Dendy, 1897: 236; Whitelegge,
1907: 507.
Isociella jacksoniana; Bergquist & Tizard, 1967: 187.
MATERIAL. HOLOTYPE: AMZ466: Port Stephens,
NSW, 32° 42'S, 152 °06'E, no other details known
(label 'Clathria macropora Lend., type'). LECTOTYPE of P. jacksoniana: BMNH1887.5.2.9: Port
Jackson, NSW, 33 ° 51'S, 151 ° 16'E, 54-90m depth, coll.
HMS 'Challenger' (dredge). PARALECTOTYPE of
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zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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FIG. 132. Clathria (Isociella)macropora Lendenfeld (holotype AMZ466). A, Choanosomal principal styles. B,
Subectosomal auxiliary subtylostyles. C, Modified palmate isochelae. D, Section through peripheral skeleton.
E, Australian distribution. F, Holotype. G, Paralectotype of Phakellia jacksoniana BMNH1887.5.2.8.
^
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273
FIG. 133. Clathria (lsociella) macropora Lendenfeld (holotype AMZ466). A, Choanosomal skeleton. B,
Semi-renieroid fibres. C, Bases of subectosomal auxiliary styles. D, Modified palmate isochelae.
274zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
P. jacksoniana: BMNH1887.5.2.8: same locality.
OTHER MATERIAL: VIC - AMZ771.
HABITAT DISTRIBUTION. Subtidal rock reef to
deeper offshore reefs, in soft sediments; 12-90m depth
known only from Australia: Port Stephens, Port Jackson (NSW), Port Phillip (Vic) (Fig. 132E).
TABLE 27. Comparison between present and published records of Clathria (lsociella) macropora (Lendenfeld). Measurements in p,m, denoted as range
(and mean) of spicule length x spicule width (N=25).
.
DESCRIPTION. Shape. Irregularly flabellatedigitate or flabellate, planar, 85-170nrun long,
55-105mm maximum breadth, with short
cylindrical stalk 15-25mm long, 7-15mm
diameter, one or more thinly lobate, bifurcated
branches, up to 8mm thick, either free or fused to
adjacent branches, with rounded, digitate, uneven
or shaggy margins.
Colour Yellow-grey or brown in ethanol.
Oscules. Oscules small, 1-3mm diameter, in special areas (sieve-plates) scattered over surface of
branches, with series of stellate subdermal
drainage canals surrounding each osculum.
Texture and surface characteristics. Firm,
flexible; branches with separate inhalant and exhalant faces; one surface porous, rugose, with
irregular longitudinal ridges, microconules or irregular striations; other surface relatively
smooth, membraneous.
Ectosome and subectosome. Ectosome
membraneous, hispid, with points of
choanosomal principal styles protruding through
surface, individually or in multispicular plumose
bundles; surface skeleton with relatively sparse
tangential, paratangential or sometimes plumose
erect skeleton of small subectosomal auxiliary
styles projecting between principal spicules,
sometimes surrounding (in proximity to) principal spicules reminiscent of Raspailiidae.
Choanosome. Choanosomal skeleton with very
slightly compressed axis and plumose sub-isodictyal, sometimes renieroid extra-axis. Axial region
with moderately heavy spongin fibres, forming
tight irregularly reticulate meshes cored by
paucispicular tracts of choanosomal principal
styles; axis (corresponding to central lamellae
and basal stalk) has few ascending, primary
tracts, forming multispicular, halichondroid
structures, 250-400p,m diameter, producing few
multispicular, dendritic tracts running from basal
stalk to periphery, 50-150Rm diameter; extraaxial skeleton sub-isodictyal with plumose
spicule tracts bound by collagen (without fibre
component), ascending to surface, cored by uni-,
pauci- or less frequently multispicular tracts of
choanosomal principal styles; primary ascending
extra-axial spicule tracts (up to 5 spicules sideby-side) arise perpendicular to axis, intercon-
Para1ectotype of
SPICULE
Choanosomal
principal
styles
P. jacksoniana
Holotype
(AMZ466)
AMZ77I
(BMNH
3694446.4y
552x
21 - (26.6) - 35
1887.5.2.8)
441-(494.7)558 x
19 - (28.8) - 38
350(462.3)548 x
11 - (24.5) - 36
155(216.7)
384 x 2.5
(5.9)-9
Subectosomal 191 (287.0)
162 (197.1)
auxiliary
424 x 4 - (6.4) - 9 226 x 4 - (6.7) -9
styles
-
Echinating
acanthostyles
-
-
absent
absent
Chelae
8-(12.9)-16
9412.0)-14
Toxas
absent
absent
-
-
-
absent
9-(12.5)-15.5
—
absent
nected by more or less transversely orientated,
-
-
smaller secondary uni- or paucispicular tracts
producing predominantly subisodictyal structure; peripheral spicule tracts more plumose than
deeper choanosomal tracts; meshes produced by
spicule-fibre anastomoses in extra-axial region
triangular or rectangular in shape, 280-560Rm
diameter; echinating acanthostyles absent;
mesohyl matrix contains abundant relatively
heavily pigmented spongin, with moderate quantities of auxiliary spicules, and choanocyte chambers oval, 50-19811m diameter.
Megascleres (Table 27). Choanosomal principal
styles long or short, thick, slightly curved at
centre, less often straight, with rounded or slightly tapering, smooth bases, varying from fusiform
to hastate points.
Subectosomal auxiliary styles variable in
length, thin, straight or very slightly curved, with
basal terminations varying from evenly rounded,
tapering hastate, quasi-diactinal mucronate or
slightly subtylote, and with hastate points.
Echinating spicules absent.
Microscleres (Table 27). Palmate isochelae with
highly modified, relatively small alae bearing
wing-shaped fluted processes; lateral alae entirely fused to shaft; front ala complete or bifurcated
with medial tooth; chelae frequently twisted or
occasionally anisochelate.
Toxas absent.
REMARKS. The synonymy of C. macropora and
P. jacksoniana is obvious on type material, but
virtually impossible to tell from published
descriptions, so the synonymy was overlooked by
Hooper & Wiedenmayer (1994). In nearly all
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
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275
features type material of both species is identical
although auxiliary spicules in macropora are
relatively larger.
The name macropora is the senior name. Ridley & Dendy's (1886) flabellata is a junior
homonym of Carter's (18850 species, and
Dendy's (1897) replacement jacksoniana now
considered to be redundant. Unfortunately macropora requires further clarification given that it
has been misused and misinterpreted. My interpretation of macropora is based solely on the
holotype because despite Hallmann's (1912: 166)
arguments in defense of Lendenfeld's systematics, there is no doubt that Lendenfeld was
dealing mainly with specimens which did not
belong to the Microcionidae. Clathria macropora
is nothing like Plectispa macropora (sensu Lendenfeld, 1888; holotype AMG9159), which has
smooth echinating spicules, identical to those
coring fibres, and is referred here to Holopsamma. Clathria macropora Lendenfeld (1888)
(AMZ959) from Port Jackson, NSW and C. macropora from Port Chalmers, Qld (BMNH1950.2.12.60) are specimens of Crella incrustans
(Crellidae). Three other specimens in the AM
collections bearing the name macropora': one
specimen from Nelson Bay, New Zealand collected by Arthur Dendy (AM unreg.) is a Crella
with an ectosomal crust similar to C. levis var.
digitata (AMZ454) (= C. incrustans); another
(AMZ4035) collected from Dee Why, Sydney
(RRIMPFN1338) and the third (AMZ4187
(RRIMPFN1428)) from 'Tumbledown', Jibbon
Head, NSW, both from the Roche Collection, are
haplosclerid sponges with three dimensional ectosomal skeletons, probably related to Amphimedon (Niphatidae).
This taxon was not among several hundreds of
specimens from subtidal and deeper water in Port
Jackson, Port Stephens and the adjacent coastline
(collected by the NSW Environmental Protection
Authority and Sydney Water Board benthic
monitoring surveys). It is also doubtful that it
occurs in New Zealand, as Bergquist & Fromont
(1988) questioned Lendenfeld's (1888) record
given that it was not subsequently rediscovered
during their substantial contemporary collections
of the NZ fauna. This is confirmed here from
re-examination of Lendenfeld's voucher
specimen from Nelson Bay, NZ (mentioned
above), which belongs to Crella.
Clathria (Isociella) macropora has distinctive
fluting on the teeth isochelae. This feature is
barely visible under light microscopy, and therefore it is possible that it may also occur in other
species of Clathria, which have not yet been
studied using SEM, but apparently it is unique to
the family. The species also has a slightly compressed axial skeleton partially offset from the
diverging, plumose, subisodictyal reticulate
extra-axial skeleton, showing vague structural
similarities to Ceratopsion and Raspailia (Syringella) (Raspailiidae). This skeletal structure
could also justify its inclusion in C. (Axociella),
but it is considered here that the subisodictyal
reticulation dominates the skeleton and is more
characteristic of lsociella than Axociella.
Clathria (Isociella) selachia sp. nov.
(Figs 134-135, Plate 3F)
MATERIAL. HOLOTYPE: NTMZ2946: E. side of
Steep Point Lighthouse, South Passage, Dirk Hartog I.,
Shark Bay, WA, 26°08.5'S, 113 ° 10.3'E, 13.vii.1987,
7m depth, coll. J.N.A. Hooper (SCUBA).
PARATYPE: QMG300562: same data.
HABITAT DISTRIBUTION. Encrusting on excavated limestone plates, in surge zone; 7m depth;
central W coast (WA) (Fig. 134E).
DESCRIPTION. Shape. Massively encrusting,
bulbous lobate-digitate, holotype 155mm wide,
105mm high, paratype 65mm wide, 60mm high,
with thickly flabellate, slightly flattened lobes, up
to 55mm thick, 110mm long, mostly fused to
adjacent lobes, together forming a bulbous mass.
Colour Bright red-orange alive (Munsell lOR
6/10), grey-brown in ethanol.
Oscules. Large, up to 8mm diameter, clustered on
tops of bulbous lobes or on margins of flattened
lobes, slightly raised above surface with
membraneous lip.
Texture and surface characteristics. Firm, compressible, moderately difficult to tear; surface
fleshy, bulbous, slightly microconulose,
membraneous in situ, with membrane collapsing
upon preservation producing fibrous, reticulate,
porous, prominently conulose, shaggy surface.
Ectosome and subectosome. Ectosome
membraneous, slightly hispid from protruding
choanosomal spicule tracts, with relatively thick
paratangential or tangential skeleton of subectosomal auxiliary subtylostyles in multispicular
tracts; ectosomal skeleton thickest at apex of
surface microconules, perched on ends of ascending choanosomal skeletal tracts.
Choanosome. Choanosomal skeleton plumoreticulate, subisodictyal and subrenieroid, with
differentiated primary ascending and secondary
transverse spongin fibres and spicule tracts;
primary ascending fibres well developed, 65-
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c
FIG. 134. Clathria (Isociella) selachia sp.nov. (holotype NTMZ2946). A, Choanosomal principal subtylostyles.
B, Subectosomal auxiliary subtylostyles. C, Anchorate-like isochela. D, Section through peripheral skeleton.
E, Australian distribution. F, Holotype.
1201J,m diameter, cored by multispicular tracts of
larger choanosomal principal subtylostyles, 3-15
spicules per tracts; spicules not occupying entire
fibre diameter; primary fibres bifurcate repeatedly, but anastomose only occasionally, producing
prominent plumose structure most noticeable at
periphery; principal subtylostyles mainly form
axial core of spicules but sometimes they
protrude through primary fibres producing
plumose brushes; primary fibres interconnected
at more-or-less regular intervals by well
developed uni- or paucispicular spongin fibres,
20-451im diameter, cored by principal subtylostyles, forming triangular, rectangular or oval
meshes, 90-180p.m diameter; skeleton slightly
more cavernous near periphery, more plumose in
structure; fibres heaviest in axis, more subisodictyal in structure; echinating megascleres absent;
mesohyl matrix light, nearly unpigmented, with
many smaller auxiliary subtylostyles and
isochelae dispersed between fibres; generally
spicules dispersed between fibres more slender,
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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277
Microscleres. Isochelae
anchorate -like, with slightly to
greatly curved shaft, variable
alae development from vestigial, tooth-like producing unguiferous chelae, to spatulate,
fused forming lobate alae;
lateral alae detached from
shaft; front alae incompletely
fused to adjacent ala; distal
portions of shaft with lateral
ridge. Length 26-(31.2)-34p.m.
Toxas absent.
ETYMOLOGY. Greek selachos,
shark, for Shark Bay, WA.
REMARKS. This species
belongs to Clathria, based on
its spiculation, choanosomal
skeletal structure, ectosomal
skeleton and fibre characteristics. It is assigned here to
C. (Isociella) given its prominent
subisodictyal skeleton, most
obvious in the axial region, and
lack of echinating spicules. It
differs from other Isociella in
having a predominently
plumo-reticulate skeleton in
the peripheral region (although
subisodictyal in the axis) and in
its spicule dimensions.
When this species was first
examined it was considered
that the anchorate-like isochelae described above might
be contaminants from another
FIG. 135. Clathria (Isociella) selachia sp.nov. (holotype NTMZ2946). A, sponge or from the substrate
upon which it grew. But
Choanosomal skeleton. B, Fibre characteristics. C, Anchorate isochelae.
numerous histological preparations made from the holotype,
from various regions within the
sinuous (probably juvenile) than in ectosomal
sponge,
repeatedly
turned up these chelae. Furskeleton; choanocyte chambers large, oval, 220thermore, examination of chelae in situ found
27011m diameter.
Megascleres. Choanosomal principal subtylos- them to be scattered both within the surface
tyles long, thick, straight or slightly curved skeleton, intermingled with the paratangential
towards base, with slightly subtylote or bundles of auxiliary spicules, and also surroundprominently subtylote bases, tapering fusiform or ing choanocyte chambers within the choanotelescoped points. Length 231-(260.6)-303Rm, somal mesohyl. They are native to this species,
although this is at odds with the current hypothesis
width 8-(11.6)-16vim.
Subectosomal auxiliary subtylostyles relative- concerning the derivation and evolutionary sigly long, slender, usually straight, occasionally nificance of these spicules (Hajdu et al., 1994).
The holotype and paratype were found growing
sinuous, with prominently subtylote bases, tapering fusiform points. Length 125-(213.9)-294Rm, side by side and are probably clones of the same
individual.
width 1-(3.4)-6Rm.
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FIG. 136. Clathria (lsociella) skia sp.nov. (holotype QMG300449). A, Auxiliary subtylostyle (coring fibres). B,
Ectosomal auxiliary subtylostyle. C, Palmate isochela. D, Raphidiform toxas. E, Section through peripheral
skeleton. F, Known Australian distribution. G, Holotype.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
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279
35mm diameter, with irregularly
and closely anastomosing digits,
rounded and irregular margins,
branches 8-18rrim diameter.
Colour. Live colouration unknown, dark brown in ethanol.
Oscules. Not observed.
Texture and surface characteristics. Firm, compressible; surface macroscopically even,
microscopically conulose, hispid,
with subectosomal striations.
Ectosome and subectosome. Ectosome with specialised skeleton
composed of smaller auxiliary subtylostyles, forming sparse multispicular bundles on surface,
arising directly from heavy, darkly
pigmented peripheral fibres; tips of
primary spongin fibres in
peripheral region protrude through
surface, producing surface
microconules up to 300pLm long;
majority of peripheral fibres nearly
tangential, forming irregular
reticulate meshes, cored by multispicular tracts of larger auxiliary
spicules (6-10 spicules per tract).
Choanosome. Skeletal architecture is irregularly isodictyal and
renieroid reticulate, with clearly
differentiated primary, ascending
fibres, 60-125Rm diameter, cored
by pauci- or multispicular fibres of
larger auxiliary subtylostyles, up
FIG. 137. Clathria (Isociella) skia sp.nov. (holotype QMG300449). A, to 8 spicules abreast, interconEctosomal skeleton. B, Fibre characteristics. C, Skeletal structure. D,
nected by secondary, transverse,
Palmate isochela. E, Raphidiform toxas.
regular or irregular, uni- or
paucispicular fibres (22-48p.m
Clathria (Isociella) skia sp. nov.
diameter); spongin fibres heavy; echinating
(Figs 136-137, Table 28)
spicules absent, although coring megascleres
may protrude through fibres at oblique angles,
MATERIAL. HOLOTYPE: QMG300449 (fragment becoming more plumose towards periphery; fibre
NTMZ1522): W. of Sudbury Reef, Cairns region, anastomoses produce triangular, rectangular or
Great Barrier Reef, Qld, 17 °03'S, 146 °07.8'E, 33-36m
depth, 28.i.1981, coll. A. Kay (trawl). PARATYPE: oval meshes, 190-425Rm in diameter, becoming
AMG5043: Masthead I. lagoon, Capricorn-Bunker more regular near periphery; mesohyl matrix
Group, Great Barrier Reef, Qld, 23 ° 32'S, 151 °43'E, heavy, darkly pigmented, and many scattered
auxiliary subtylostyles dispersed between fibres;
40m depth, no other details known.
choanocyte chambers small, oval, 40-85 p.m
HABITAT DISTRIBUTION. Attached to shell frag- diameter.
ments or coral rubble, in soft sediments; 33-40m depth;
Me gascleres (Table 28). Principal spicules abCairns region (NEQ), Gladstone region (MEQ) (Fig.
sent.
136F).
Auxiliary subtylostyles (coring fibres) long,
DESCRIPTION. Shape. Erect, bulbous-lobate slender, invariably straight, with prominent,
digitate, clathrous sponge, 45-95mm long, 18- smooth subtylote bases and fusiform points.
280zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Ectosomal auxiliary subtylostyles significantly smaller than coring spicules but otherwise of
identical geometry.
Echinating spicules absent.
Microscleres (Table 28). Palmate isochelae uncommon, unmodified, with long lateral alae entirely fused to shaft and front ala completely
fused, slightly spatulate.
Toxas raphidiform, extremely thin, slightly
curved at centre, sometimes nearly straight or
bow shaped, rarely asymmetrical.
ETYMOLOGY. Greek skias , dark, dim.
REMARKS. This species is a greatly reduced
Clathria, lacking true principal spicules, echinating spicules and having containing relatively
sparse microscleres. Conversely its spongin fibre
system is well developed and fibres are heavy. It
is assigned to the Isociella group on the basis that
its skeletal structure is essentially subisodictyal,
and it lacks echinating spicules, but it is admitted
that its reduced characteristics make its true affinities speculative.
The sparsely developed ectosomal skeleton,
composed of plumose bundles of smaller
auxiliary styles, also indicates affinities to the
Thalysias group, but in most respects (spicule
geometry, skeletal structure, fibre development
and growth form) it differs from all other species
of either lsociella and Thalysias. Clathria
(Isociella) skia was initially thought to be Amphilectus hispidulus Ridley, from Torres Strait
(FNQ). From both published descriptions of A.
hispidulus (Ridley, 1884a; Hentschel, 1911) it
apparently lacked echinating megascleres, lacked
principal spicules and had well developed subisodictyal skeletal structure, but re-examination
of the holotype showed that it does have echinating acanthostyles, and differentiated principal
and auxiliary megascleres (thus more appropriately included in Clathria (Clathria); see
above), and it also has substantially different fibre
characteristics from C. (I.) skia indicating that
they are not conspecific.
OTHER SPECIES OF CLATHRIA
(ISOCIELLA)
TABLE 28. Comparison between of Clathria (lsociella) skia sp.nov. Measurements in p.m, denoted as
range (and mean) of spicule length x spicule width
(N=25).
Paratype
(AMG5043)
Holotype
(QMG300449)
SPI CULE
Choanosomal
principal styles
absent
absent
zyxwvutsrqponmlkjihgfedcbaZYXWVUTS
Auxiliary
(coring) styles
I 62-(248.4)-368x
2.5- ( 4.5) - 8
1544242.1)-318 x
244.1) - 6
Auxiliary
(ectosomal) styles
97-(111.9)-129 x
l.542.4)-4
87-(110.2)-132 x
1.5- ( 2.4) - 4
absent
absent
Echinating
acanthostyles
Chelae
Toxas
9-(15.3)-18
14-(16.3)-I 9
724104.4)-145 x
65-(81.1)-108 x
0.2-(0.5)-0.8
0.240.4)-0.8
Clathria (Thalysias)
Duchassaing & Michelotti, 1864
Thalysias Duchassaing & Michelotti, 1864: 82.
Rhaphidophlus Ehlers, 1870: 19.
Tenacia Schmidt, 1870: 56.
Echinonema Carter, 1881a: 378.
? Thalassodendron Lendenfeld, 1888: 222.
StylotellopsisThiele, 1905:456; de Laubenfels, 1936a:
112.
Colloclathria Dendy, 1922: 74.
Damoseni de Laubenfels, 1936a: 110.
DEFINITION. Specialised ectosomal skeleton
composed of two size classes of auxiliary (subtylo)styles, with smaller ectosomal spicules
usually overlaying larger subectosomal ones
forming a continuous palisade, or discrete
bundles, mainly erect, sometimes paratangential,
or rarely tangential to surface; choanosomal
skeleton without any marked differentiation between axial and extra-axial regions; echinating
acanthostyles usually present.
TYPE SPECIES. Spongia juniperina Lamarck, 1814:
444 (by synonymy).
REMARKS. Of 137 named species described in,
or referred to Thalysias or one of its synonyms,
93 are thought to be valid of which 37 are
recorded here from Australian waters including
10 new species.
Clathria (Isociella) incrustans (Bergquist, 1961)
Isociella incrustans Bergquist, 1961a: 42-43, text-figs
Clathria (Thalysias) abietina (Lamarck, 1814)
15a-b [Ahipara Bay, N. New Zealand; originally
(Figs 138-141, Tables 29-30, Plate 4C-D)
assigned to the Suberitidae, Hadromerida].
Bergquist & Fromont, 1988: 114-116, p1.53, figs e-f,
Spongia abietina Lamarck, 1814: 450, 377.
p1.54, figs a-b; Dawson, 1993: 36.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Clathriaabietina; de Laubenfels, 1954: 141-142, textMATERI AL. HOLOTYPE: NMNZ unregistered. NZ.
fig.90; Hooper & Wiedenmayer, 1994: 267.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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281
Clathria aculeata Ridley, 1884a: 443-444, p1.40, fig. 1,
p1.42, fig.k; Ridley & Dendy, 1887: 147, 246, 254;
Topsent, 1897b: 447; Burton, 1934a: 558; Burton,
1959a: 243; Levi & Levi, 1989: 80-81, p1.7, fig.3.
Rhaphidophlus aculeatus; Topsent, 1932: 115, p1.4,
fig. 10; Vacelet & Vasseur, 1977: 114; Vacelet et al.,
1976: 73-74.
Rhaphidophlus cervicornis, in part; Vacelet & Vasseur, 1971: 73.
cf. Microciona prolifera; Vosmaer, 1935a: 610, 633,
664.
MATERIAL. LECTOTYPE: MNHNDT634: precise
locality unknown (suspected to be 'Australia'; Topsent
(1932: 115)). PARALECTOTYPES: MNHNDT3342,
3343: same details. HOLOTYPE of C. aculeata:
BMNH1882.2. 23.258: Torres Strait, Qld, 9°41'S,
142°17'E, 6-8m depth, coll. HMS 'Alert' (dredge).
OTHER MATERIAL. MICRONESIA - USNM
22808, USNM23090. QLD - BMNH1887.5.2.110,
QMG300791. DARWIN HARBOUR, NT NTMZ226, NTMZ426, NT MZ458, NTMZ468,
NTMZ498, NTMZ512, NTMZ515, NTMZ886,
NTMZ903, NTMZ928, NTMZ955, NTMZ2050,
NTMZ2078, NTMZ 2079, NTMZ2085, NTMZ2258,
NTMZ2390, NTMZ2391, NTMZ2395, NTMZ2399,
NTMZ 2611, NTMZ2642, QMG300169, NTMZ2646,
QMG304077, NTMZ I 943, NTMZ1958, NTMZ 1963,
QMG303373, QMG303382, NTMZ2089, NTMZ2161, NTMZ2163, NTMZ2186, NTMZ2191,
NTMZ2194, NTMZ2195, NTMZ820, NTMZ835,
QMG300414. BYNOE HARBOUR, NT - NTMZ1073,
NTMZ2106, QMG303447, QMG303534. SHOAL
BAY, NT - QMG303539, QMG303571. TIMOR SEA,
NT - NTMZ3090. PORT ESSINGTON, COBOURG
PENINSULA, NT - NTMZ68, NTMZ69, NTMZ90,
NTMZ1393, NTMZ3304, NTMZ577, NTMZ 1328,
NTMZ1329, NTMZ1330, NTMZ1331, NTMZ1332,
NTMZ1333, NTMZ1334, NTMZ 1343, NTMZ2500,
NTMZ2501, NTMZ2509, NTMZ2510, NTMZ3245,
NTMZ3255, NTMZ 3258, NTMZ3260, NTMZ3268,
NTMZ3272, NTMZ3277, NTMZ3278, NTMZ3284,
NTMZ 3289, NTMZ3295, QMG300386, NTMZ1352.
ARAFURA SEA, NT - NTMZ2521, NTMZ 2522,
NTMZ2523, NTMZ129, NTMZ130, NTMZ138.
WESSEL ISLANDS, NT - NTMZ 3902, QMG300764
(NCIQ66C-4692-Q), NTMZ3921, NTMZ3930,
QMG300757 (NCIQ 66C-4773-F), QMG300508
(NCIQ66C-4772-C), NTMZ3947, QMG300765
(NCIQ66C-4808-R). NORTHWEST SHELF
REGION, WA - NTMZ1036, NTMZ1209, NTMZ1217, NTMZ 1244, NTMZ1272, NTMZ1314,
NTMZ1411, NTMZ1423, WAM151-82, WAM15582 (fragments NTMZ1731, NTMZ1732), NTMZ1770,
NTMZ1801, NTMZ1820, NTMZ1824, NTMZ 1852,
NTMZ2272, NTMZ2329, NTMZ2349, NTMZ2486,
NTMZ3017, NTMZ3030, NTMZ 3031, NTMZ3032,
NTMZ3033, NTMZ3396, QMG300448 (NCIQ66C1517-P), QMG300117 (NCIQ66C1518-Q) (fragments
NTMZ3488, NTMZ3489), PIB0004-595 (fragment
QMG300051)
HABITAT DISTRIBUTION. Shallow-water0-25m
depth, predominantly on sides and tops of rock and
dead coral heads, invariably exposed to currents.
Deeper-water specimens (26-86m depth) mostly associated with exposed rock substrates in gravel, silt or
shell-grit substrates; central SW Pacific Ocean (Low
Isles, Great Barrier Reef (Burton, 1934a), Torres Strait
(Ridley, 1884a; Ridley & Dendy, 1887)); E Indian
Ocean (Arafura Sea, Timor Sea, mid-WA coast
(present study)); also tropical Indo-west Pacific:
central NW. Pacific Ocean (Marshall and Caroline Is
(de Laubenfels, 1954), Philippines (Levi & Levi,
1989)); W. Indian Ocean (Madagascar (Vacelet et al.,
1976, 1977), Red Sea (Burton, 1959a)). Within
Australian waters this species extends across the N and
NW coasts, from the Cairns region, Torres Strait and
Gulf of Carpentaria, Qld, to the Exmouth Gulf region,
WA (Fig. 138J). It is only rarely encountered on the E
Qld. coast and must be considered a predominantly
Indian Ocean species.
DESCRIPTION. Shape. Arborescent; thickly
cylindrical digitate branches on long or short,
thick stalk (3-21mm basal diameter) with expanded basal attachment, never rhizomous;
branches rarely anastomosing, branching mostly
irregular, bushy, occasionally regular growing in
I plane, sometimes flagelliform, occasionally expanded, club-shaped; total length and branch
diameter variable (50-460mm; 2-22mm, respectively).
Colour. Live colouration highly variable, ranging
from maroon, red, orange, yellow, brown to grey;
colour in ethanol ranges from black, brown to
grey (Munsell values given below); pigments
oxidise in air; maroon colouration is highly alcohol soluble; yellow pigmentation more stable
in ethanol but easily scratched from surface of
living sponge, leaving a maroon or mauve pigmentation beneath; pigmentation extends into
periphery of choanosomal mesohyl.
Oscules. Exhalent pores usually very small, 0.62.0mm diameter, barely visible optically, dispersed on distal ends of branches surrounded by
surface conules, occasionally scattered along entire lateral margins of branches.
Texture and surface characteristics. Firm, barely
compressible; branches rubbery whereas stalk
more rigid; surface dense, entirely opaque in life,
with characteristic regularly scattered surface
conules; conules tapering, distally rounded or
blunt, usually absent from basal and distal extremities of branches, 2-6mm maximum basal
diameter.
Ectosome and subectosome. Very dense, continuous palisade of small ectosomal auxiliary
subtylostyles forming discrete brushes overlay-
282zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 138. Clathria (Thalysias) abietina (Lamarck) (holotype MNHNDT634). A, Choanosomal principal subtylostyle. B, Ectosomal auxiliary subtylostyle. C, Subectosomal auxiliary subtylostyle. D-E, Echinating
acanthostyles. F, Accolada toxa. G, Wing-shaped toxas. H, Palmate isochelae. I, Section through peripheral
skeleton. J, Australian distribution.
ing thickly plumose tracts of larger subectosomal secondary elements but forming irregular, closeauxiliary subtylostyles in peripheral skeleton (the meshed reticulation; fibre meshes oval to eliptilatter often obscuring ectosomal brushes); subec- cal, l00-350ttm diameter; fibres cored by
tosomal spicule tracts arise from ultimate multispicular tracts of larger auxiliary subtyloschoanosomal fibres, intermingled with tufts of tyles occupying 60-80% of fibre diameter;
principal choanosomal subtylostyles echinating
choanosomal principal subtylostyle uncommon
peripheral fibres; principal spicules not extending
into ectosomal skeleton; mesohyl matrix heavy within fibres, mainly found at fibre nodes
and pigmented in both ectosomal and subec- protruding through fibres individually or in
bundles; fibres also echinated by acanthostyles
tosomal regions.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
especially on peripheral fibres; mesohyl matrix
Choanosom e. Skeleton irregularly reticulate;
heavy spongin fibres, 50-125 t.i.m diameter, heavi- heavy but only lightly pigmented, slightly
ly anastomosing, not divided into primary or granular; choanocyte chambers oval or eliptical,
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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283
TABLE 29. Comparison in range of spicule dimensions between present and (more abundant in
published records of Clathria (Thalysias) abietina. All measurements are given in smaller form). Length I:
p.m and denoted as length x width (N=25).
7.9412.1)-19.9pm (lec-
totype 11 -(12 .5 )14.5p,m), length II: 0.5Principal
172-254x 12.7
165-258 x 9-16
230
172-254
(5.8)-10Rm (lectotype
270-320 x 6-15
styles
4-(5)-6p,m).
Subectosomal 157-343 x 5-10
350 x 8.5
149-276 x 2-5
120-250 x 2-3 127-387 x 1-20
Toxas relatively unstyles
common
with 2
Ectosomal
45-230x 1-10
62-132 x 2-5
81-117 x 2-4
geometric forms; (i)
styles
wing-shaped, short,
41-109 x 1-19
53-74 x 4-6
50-70 x 7-8
90 x 7.9
Acanthostyles 61-94 x 4-10
thin, generously curved
11-15
12.7
10-14
10-15
8-20
Chelae I
at central with reflexed
4-6
1-10
Chelae II
points; (ii) accolada
15-180
63
91-210
45-55
3-345
Toxas
toxas long, thin almost
Material:
straight with small anholotype MNHN DT634.
1. Spongia abietina zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
gular central curvature
2. Clathria aculeata Ridley (1884a: 443).
3. De Laubenfels' (1954:141) material USNM 22808, 23090.
and straight points.
4. Rhaphidophlus cervicornis, in part, (Vacelet & Vasseur, 1971:96).
Length I: 5424.9)5. Present material (N=25).
75pm, width 1.8-(2.4)4.1pm (lectotype
40-170p.m diameter; larger auxiliary styles also 15-(30.8)-61 x 2.0-(2.9)-3.5p.,m). Length II: 58dispersed between fibres in disorganised tracts.
(184.6)-345p,m, width 0.3-(0.8)-1.5pm (lecMegascleres. Choanosomal principal styles-sub- totype 65-(120.8)-180 x 0.540.9)-1.211m).
tylostyles usually uncommon, short, stout, slightly curved at centre or near base, occasionally Larvae. Parenchymella larvae observed in
straight, with slightly subtylote bases or evenly peripheral choanosomal skeleton of about 20
rounded bases; spicules usually completely specimens, predominantly during the tropical wet
smooth, exceptionally with minutely season (February-April) and less so during the
pre-dry season (May-July). Larval incubation
microspined bases, tapering to abruptly (hastate)
sharp points. Length 1214202 .7)-300.5p,m, was distinctly seasonal and occurred more-orwidth 5-(14.4)-24pm (lectotype 1654215.5)- less evenly throughout the population (i.e., irrespective of depth of collection; Fig. 141).
258pm x 9.5-(14.3)-15.5pm).
Larvae were 600-950pm diameter, elongate-oval
Subectosomal auxiliary subtylosytles straight
in shape and many contained larval megascleres;
or slightly curved near base, with prominent subcilia
were not observed (preserved material).
tylote, usually microspined bases, tapering to
3% of specimens had filamentous
Associations.
fusiform points. Length 127-(258.4)-386.9p,m,
width 1.148.2)-20.1 p.m (lectotype 1574274.4)- algae coring fibres in addition to longer auxiliary
megascleres.
343pm x 5-(6.7)-10p,m).
Ectosomal auxiliary subtylostyles short, thin, Variation. Highly variable in live colouration,
invariably straight, with prominently subtylote, non-fibre skeletal development, megasclere size,
typically microspined bases, tapering to fusiform relatively consistent in growth form, surface feapoints. Length 44.8-(114.6)-230p,m, width 1.1- tures, fibre skeleton, spicule geometry. Gross
(3.8)-10.1pm (lectotype 62-(85.5)-132 x 243.4)- morphology: stalked, bushy, branching in more
than 1 plane, branches bifurcate, occasionally
5p,m).
Echinating acanthostyles moderately long, anastomosing (46%), planar branching (33%),
thick, straight, with prominently subtylote, dendritic planar branching (fans) (14%), or single
spined base, virtually aspinose 'neck' (proximal digits with no or few bifurcations (7% of
to base), and lightly spined shaft; spines relatively specimens). Atypical growth forms (few surface
large. Length 40.6-(82.4)-109pm, width 1.1- conules, few branches, thin branching) found
(7.5)-19pm (lectotype 61-(74.4)-94 x 446.5)- predominently in deeper offshore coastal waters
(40m depth). Live colouration: highly variable,
1011,m).
Microscleres. Palmate isochelae with long lateral no particular pigment considered to be typical,
alae completely fused to shaft, shorter entire front ranging from (i) maroon, evenly pigmentation
alae; two size categories present, both relatively (Munsell 5R4/10-2.5R4/8-10), (ii) red-maroon,
abundant, both with examples of contort shafts even (5R518-10), (iii) bright red-orange, even
SPICULE
1
-
2
3
4
5
MEMOIRS OF THE QUEENSLAND MUSEUM
284zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
Clat hr ia ( Thaly sias) abiet ina (Lamarck) typical growth forms. A, Lectotype of C. aculeat a
FIG. 139. zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
BMNH1882.2.23.258. B, BMNH1887.5.2.110. C, Lectotype MNHNDT634. D, Paralectotype MNHNDT3343.
E, Paralectotype MNHNDT3342. F-G, Deeper water specimens NTMZ1820, NTMZ1314. H, Shallow-water
specimen NTMZ68.
(5R5/12), (iv) bright orange, even (10R5112-
10R6/10-12-10R7/10), (v) orange-brown, even
(10R418-10), (vi) orange-yellow, even
(7.5YR7112), (vii) pale brown-pink, even
(10R7/4), (viii) light brown, with pink conules
(10R7/4, 5R8/4), (ix) light brown, with dark greybrown conules (2.5YR714-5YR712-6, 2.5YR6165 YR3-5/2), (x) pale brown, even
(5YR616-5YR7/4-6-5YR8/4), (xi) muddy grey,
even (7.5YR7/2-8/2) (xii) yellowish grey, even
(2.5Y816), (xiii) yellow, even (2.5Y7/8), (xiv)
yellow, with pink conules (2.5Y8/6, 10R7/6),
(xv) lime-yellow, with dark brown conules
(2.5Y7/10, 2.5Y5/2), (xvi) yellow-green mottle
(5Y8/12). No obvious relationship between live
colouration and depth or substrate type from
present data; moreover, specimens with widely
different pigmentation observed growing sideby-side (Plate 4D). Subectosomal skeletal
development: correlation between branch
diameter and extent of development of peripheral
skeleton, with 8% of specimens (all thinly
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
285
TABLE 30. Summary of results from one-way ANOV's (Model I), testing for variability in spicule lengths and
widths between locality, bathymetric and seasonal distributions of Clathria (Thalysias) abietina.
LOCALITY
SPICULE
(N)
DEPTH2
F
Prob.
F
SEASON3
Prob.
(N)
F
Prob.
(775)
1.67
P>0.05
(775)
0.26
P>0.05
P<0.0005
(775)
2.46
P>0.05
4.04
P<0.01
(775)
2.08
P>0.05
5.48
P<0.001
(775)
4.15
P<0.01
Choanosomal
styles L
(1950)
3.31
P<0.05
2.11
P>0.05
W
(1950)
0.54
P>0.05
4.44
P<0.005
Subectosomal
(1950)
6.21
P<0.001
10.61
W
(1950)
1.89
P>0.05
Ectosomal
styles L
(1950)
0.85
P>0.05
W
(1950)
0.77
P>0.05
1.13
P>0.05
(775)
1.99
P>0.05
Acanthostyles
L
(1950)
4.74
P<0.01
5.95
P<0.0005
(775)
1.22
P>0.05
W
(1950)
3.17
P<0.05
4.64
P<0.005
(775)
5.17
P<0.0025
Chelae I L
(1925)
0.48
P>0.05
1.21
P>0.05
(775)
4.04
P<0.01
Chelae II L
Toxas L
(1925)
0.14
P>0.05
2.04
P>0.05
(775)
0.88
P>0.05
(1925)
0.39
P>0.05
2.72
P<0.05
(750)
1.17
P>0.05
W
(1925)
0.42
P>0.05
5.49
P<0.001
(750)
1.33
P>0.05
styles L
Number of groups:
1. 3 locality groups (NWS, DAR, CP localities).
2.4 depth groups (0-4m, 4-10m, I0-40m, 40m depth).
3. 4 seasonal groups (Darwin region only: wet (FMA), pre-dry (MJJ), dry (ASO), pre-wet (NDJ)).
branching) having peripheral choanosomal fibres
lying immediately below ectosomal crust, thin
paratangential subectosomal region, and acanthostyles echinating peripheral fibres piercing ectosomal skeleton. Development of extra-fibre
skeleton: 4% with very abundant juvenile
auxiliary subtylostyles distributed throughout
mesohyl, 90% with at least some interstitial
auxiliary spicules, and 6% of specimens without
any interstitial auxiliary spicules dispersed between fibres. Megasclere geometry: Principal
spicules range from relatively common (21%),
uncommon (61%), or very rare (18% of
specimens). Bases of all principal spicules
smooth (79%), or up to one-quarter of principal
spicules with microspined bases (21% of
specimens). Larger auxiliary spicules
predominantly subtylote with minutely
microspined bases, but 0-74% of spicules may be
smooth in any particular specimen. Acanthostyle
geometry relatively consistent, although in 5% of
specimens two size categories were recognised
(although smaller category probably juvenile
form of larger and subsequently lumped together
in analyses), in 4% they were significantly thinner, and 1% had significantly shorter and stouter
acanthostyles than typical forms. Acanthostyle
spination slightly variable, from scattered robust,
recurved spines (65%), spines arranged in regular
longitudinal rows (5%), or minutely microspined
(12% of specimens). Microsclere geometry:
Proportion of contort to normal morphs of palmate isochelae varied from 0-44% for smaller
category, 0-20% for larger. Chelae typically
abundant, 7% of specimens isochelae of both
classes very rare, 3% smaller category rare but
larger abundant, 4% larger category rare but
smaller, 1% of specimens isochelae absent entirely. Toxas of both categories very abundant (12%),
uncommon (70%), rare (17%), or absent entirely
(1% of specimens).
Variability in spicule dimensions: Some spicules
(choanosomal principal styles, larger auxiliary
subtylostyles, acanthostyles) showed significant
variations in dimensions between samples collected from different localities (Tables 29-30),
although statistical significance was never high,
and no obvious patterns were apparent when
groups of specimens from the same localities
were compared with other groups. Some spicule
categories varied between samples collected
from different depths, in some cases with high
levels of statistical significance (P<0.001) (e.g.,
larger auxiliary subtylostyles), but no obvious
trends apparent. It is probable that effects of
differential geographical and bathymetric distributions of specimens are linked due to the
preponderance of deeper water samples from the
Northwest Shelf region and shallow water
samples from the Darwin region, making it im-
286zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 140. Clathria (Thalysias) abietina (Lamarck) (A-B, NTMZ2642; C-I, QMG303447). A, Choanosomal
skeleton. B, Fibre characteristics. C, Echinating acanthostyles. D, Acanthostyle spines. E, Base of subectosomal
auxiliary subtylostyle. F-G, Palmate isochelae. H, Wing-shaped toxas. I, Accolada toxas.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
^
287
SEASON
TOTAL SAMPLES
SAMPLES WITH
LARVAE
WET
15
PREDRY
17
9
3
DRY
PREWET
19
0
17
0
DEPTH
0-4m
5
0
4-10m
30
7
10-40m
23
5
>40m
11
0
FIG. 141. Clathria (Thalysias) abietina (Lamarck) Incidence of incubated larvae within sampled populations from NW Australia.
possible to separate the influences of either factors on this observed variability. Variability of
spicule size between samples collected during
different seasons were analysed for Darwin
samples only (Table 30), with significant differences observed for lengths of ectosomal
auxiliary subtylostyles and larger isochelae, and
widths of acanthostyles. Data also indicate a
higher level of variability in sizes of larger
isochelae during the wet season (February-April)
than during other seasons, although this result is
of uncertain biological significance.
REMARKS. Notwithstanding its considerable
morphological variability C. (T.) abietina is easily recognised in the field with distinctive stalked
digitate growth form and prominent surface conules. The most similar species in growth form is
C. (T) cervicornis but this has a much thinner,
stoloniferous branching morphology, lacks principal megascleres completely, fibres have less
spongin, toxas are different in morphology and
size, and spicule dimensions are different. Skeletal structure is unusual where larger auxiliary
subtylostyles are found in three locations in the
skeleton: 1) subectosomal skeletal tracts forming
organised plumose tracts supporting the ectosomal skeleton; 2) dispersed between fibres in
disorganised tracts; 3) and coring all spongin
fibres to the virtual exclusion of principal
spicules. Principal styles mainly form plumose
brushes protruding from fibres in plumose
bundles, functionally representing a second
category of echinating spicule.
There are some minor differences between type
material and other specimens examined, including a higher proportion of choanosomal principal
spicules found echinating fibres, the absence of
contort isochelae, and specific details in some
spicule dimensions (Table 29). But given the
large range of variation in some characters, the
relatively large sample sizes from widely dispersed populations, and the antiquity of the dried
holotype this variability is insignificant. Re-examination of Ridley's (1884a) holotype of C.
aculeata confirmed that it is a synonym of C. (T)
abietina, with shape, texture, colour in spirit,
spiculation and skeletal architecture virtually
identical. Conversely, C. (T) coralliophila (see
below) has different spicule geometry and skeletal architecture, and Burton's (1959a: 243)
proposed merger of C. coralliophila into C.
aculeata is rejected.
Specimens described by de Laubenfels (1954)
from the central west Pacific differ from
Australian populations in that they have a more
restricted size range of isochelae (i.e., one size
category), relatively small acanthostyles (Table
29), and an ectosomal (peripheral) skeleton almost completely covered by subectosomal
spicule brushes (as opposed to ectosomal spicule
brushes). In fact de Laubenfels (1954) completely
overlooked the presence of ectosomal
megascleres, not differentiating between spicules
coring fibres from those forming the peripheral
skeleton.
Clathria (Thalysias) aphylla sp. nov.
(Figs 142-143, Plate 5D)
MATERIAL. HOLOTYPE - QMG300477 (NCIQ66C-4640-K): NW. of E. Passage, Easter Group, Houtman Abrolhos, WA, 28°40'S, 113°50'E, 20m depth,
17.ix.1990, coll. NCI (SCUBA).
HABITAT DISTRIBUTION. Staghom and plate coral
fringing reef; 20m depth; known only from the type
locality, Houtman Abrolhos (WA) (Fig. 142H).
DESCRIPTION. Shape. Thin, leaf-like,
foliaceous, convoluted, basically frondose
bundles of lamellae covering coral substrate; individual fronds attached directly to substrate,
sometimes completely enveloping staghorn
corals, or attached via small basal stalk, up to
24mm long, 5mm diameter, or attached to adjacent lamellae; individual lamellae usually flat,
elongate, oval or eliptical, up to 65mm long, 2nun
thick, with rounded or sinuous, convoluted margins, superficially resembling a Padina algae, or
palmate-digitate margins, or sometimes curled in
vasiform growth forms.
Colour. Dull yellow alive (Munsell 2.5Y 8/8),
pale brown in ethanol.
Oscules. Small, on upper surface, less than 2rnm
diameter, flush with surface.
288zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
Texture and surface characteristics. Soft, compressible but difficult to tear; upper osculiferous
surface slightly concave, smooth, usually even,
occasionally concentrically striated, or producing
short fronds of folds on surface; lower porous
surface slightly convex, even or slightly striated,
with parasitic zoanthids covering parts of surface.
Ectosome and subectosome. Erect or paratangential brushes of ectosomal auxiliary styles forming
thin surface layer, with choanosomal principal
styles protruding through ectosome perpendicular to surface, singly or several together, extending up to 150p.m through surface;
choanosomal fibres immediately below ectosome, with reduced subectosomal skeleton;
subectosomal auxiliary styles tangential or
paratangential, lying immediately below surface
underlying ectosomal spicule brushes; mesohyl
matrix in peripheral skeleton heavy, darkly pigmented.
Choanosome. Regularly reticulate, slightly
renieroid skeleton; without compression or major
differences between peripheral skeleton and core;
spongin fibres heavy, 20-70p,m diameter, not obviously differentiated into primary or secondary
elements, cored by multispicular (up to 15
spicules abreast) or paucispicular (2 or more
spicules abreast) tracts of principal choanosomal
styles; at core of skeleton principal styles confined entirely within fibres whereas in peripheral
fibres, principal spicules erect, perpendicular to
fibres, protruding through surface in plumose
bundles; fibres moderately lightly echinated by
acanthostyles, mainly at fibre nodes; fibre meshes
mainly rectangular (fibres oval), 90-170iim
diameter, with some triangular meshes, relatively
even throughout skeleton; mesohyl matrix light,
with few scattered megascleres; choanocyte
chambers oval to eliptical, 25-40Rm diameter,
lined by abundant, minute isochelae.
Megascleres. Choanosomal principal styles long,
thick, straight or slightly curved towards basal
end, rounded or slightly tapering bases, invariably smooth bases, fusiform points. Length 152(205.1)-252pm, width 4.5-(8.4)-11p.m.
Subectosomal auxiliary styles long, slender,
straight, with rounded or faintly subtylote smooth
bases, fusiform points. Length 208-(258.l)297 p.m, width 2-(2.7)-4.5 p.m.
Ectosomal auxiliary styles short, slender,
straight, rounded or slightly subtylote bases,
smooth or microspined (or mucronate) bases,
fusiform points. Length 162-(178.2)-204Rm,
width 1-(1.6)-2.5p.m.
Echinating acanthostyles relatively long,
slender, subtylote, sharply pointed, evenly spined
although fewer spines in 'neck' region proximal
to base (but not aspinose); spines small, sharp,
recurved. Length 45-(86.3)-102Rm, width 3(5.5)-81.1.m.
Microscleres. Palmate isochelae very abundant,
minute, with lateral alae completely fused to shaft
and partially fused to front ala. Length 445.1)6p.m.
Toxas not abundant, predominantly v-shaped,
variable in size, with angular central curvature
and arms bent at approximately right angles to
each other, non-reflexed arms, occasionally toxas
forceps shaped with pinched central curve and
nearly parallel arms. Length 34-(62.3)-106p.m,
width 0.8-(1.3)-2.5pan.
ETYMOLOGY. Greek phyllon , leaf; for the growth
form.
REMARKS. This species is a sister of C. (C.)
angulifera (Vic. and S Qld), having similar live
colour, slightly renieroid skeletal structure (mainly rectangular meshes), similar toxa morphology
and chelae size. In particular both species have
large v-shaped toxas, and a skeletal architecture
that verges on isodictyal (some triangular
meshes). However, they are clearly different
species showing both obvious and subtle differences in a number of characters. Clathria (T)
aphylla has a very thin, leaf-like growth form
(whereas C. (C.) angulifera is thickly lamellate,
lobate); very thick, well developed, evenly
spaced fibres cored by multispicular tracts of
large principal styles (versus widely spaced, vestigial fibres with light spongin, cored by much
smaller principal spicules in uni- or paucispicular
tracts); a specialised ectosomal skeleton composed of two size classes of auxiliary spicules
forming brushes and principal spicules protruding through the surface forming plumose brushes
and piercing the ectosome (versus a single
category of auxiliary spicule forming a tangential
or occasionally paratangential ectosomal
skeleton, without participation of principal
spicules in the ectosomal skeleton); clearly differentiated principal and auxiliary spicule
geometry (versus more subtle differences, mainly
in thickness and basal termination); rounded
bases of principal and auxiliary megascleres with
the smaller microspined (versus subtylote and
completely smooth); and substantially larger
dimensions for most megascleres. Another
species with a similar, slightly renieroid skeleton
reminiscent of C. (T.) aphylla is C. (C.) hispidula
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
289
FIG. 142. Clathria (Thalysias) aphylla sp.nov. (holotype QMG300477). A, Choanosomal principal styles. B,
Subectosomal auxiliary style. C, Ectosomal auxiliary styles. D, Echinating acanthostyles. E, Palmate isochela.
F, V-shaped toxas. G, Section through peripheral skeleton. H, Australian distribution. I, Holotype. J, Protruding
choanosomal spicules in ectosomal skeleton.
(from Torres Strait and Shark Bay), which has
only one category of auxiliary spicule and different ectosomal structure, wing-shaped toxas,
different acanthostyle geometry and specific differences in spicule sizes. All three species
probably have sister species relationship based on
skeletal architecture, here referred to the
`anguliferd group.
A few other Clathria species also have Vshaped toxas (C. (T) juniperina) and vaguely
isodictyal skeletal structure (e.g., C. (T)hirsuta),
but these differ from C. (T) aphylla in most other
respects and are not considered here to be closely
related.
Clathria (Thalysias) arborescens
(Ridley, 1884) (Figs 144-145)
Rhaphidophlus arborescens Ridley, 1884a: 450-451,
p1.40, fig.L, p1.42, figs n-n'; Thiele, 1903a: 958.
MEMOIRS OF THE QUEENSLAND MUSEUM
290zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^
FIG. 143. Clathria (Thalysias) aphylla sp.nov. (holotype QMG300477). A, Choanosomal skeleton. B, Fibre
characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E-F, Base of subectosomal and ectosomal
auxiliary styles. G, Palmate isochela. H, V-shaped toxas.
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTS
^
291
Clathria arborescens; Hooper & Wiedenmayer, 1994:
268.
cf. Microciona prolifera tropus spinosa; Vosmaer,
1935a: 610, 634.
MATERIAL. HOLOTYPE - BMNH1881.10.21.272:
Friday I., Torres Strait, Qld, 10°36'S, 142°10'E,
1.v.1881, coll. HMS 'Alert' (dredge).
HABITAT DISTRIBUTION. Ecology unknown; Torres Strait, Qld (Fig. 14411).
DESCRIPTION. Shape. Branching, arborescent,
125rrun long, 60mm maximum width of branching, with short stalk, 18mm long, 8mm diameter,
no basal attachment (presumed not collected),
proliferous bushy, slightly flattened cylindrical
branches, up to 12mm diameter, with individual
branches up to 55mm long, mostly anastomosing
with adjacent branches; branching mostly planar.
Colour Live colouration unknown, grey-brown
in dry state.
Oscules. Not observed.
Texture and surface characteristics. Texture
harsh in dry state; surface peel thick, darker than
choanosomal skeleton, lightly hispid, with
numerous irregular subectosomal striations, irregular conules and low ridges.
Ectosome and subectosome. Ectosomal skeleton
with very thick crust of discrete spicule brushes
composed of ectosomal auxiliary subtylostyles,
occasionally paratangential or tangential to surface, supported below by thick, vaguely plumose,
slightly disorganised bundles of larger subectosomal auxiliary subtylostyles arising from terminal choanosomal fibres; subectosomal
skeleton lacking any spongin fibres and only light
mesohyl matrix; together ectosomal and subectosomal bundles extend up to 800Rm from
peripheral fibre skeleton, and comprise up to 25%
of branch diameter.
Choanosome. Choanosomal skeleton irregularly
close-meshed reticulate. Spongin fibres very
large, heavy dark yellow-brown, 110-17011,m
diameter, forming tight oval or elongate meshes,
130-350Rm diameter; fibre reticulation without
any clear distinction between primary or secondary components, although ascending fibres marginally more abundantly echinated than
transverse connecting fibres, and some smaller
fibres between major meshes with uni- or
paucispicular core; fibres generally all cored by
multispicular tracts of choanosomal principal
subtylostyles together with subectosomal
auxiliary subtylostyles, forming a dense axial
core within fibre but occupying only about 50%
of fibre diameter; fibres abundantly echinated by
relatively large acanthostyles, more-or-less evenly dispersed throughout skeleton (not congregated in periphery as reported by Ridley);
choanocyte chambers not observed (dry
material).
Megascleres. Choanosomal principal subtylostyles with similar geometry to larger auxiliary
spicules, long, straight, usually with subtylote,
smooth or microspined bases, rarely rounded
bases, with tapering fusiform points. Length 205(225.3)-262m, width 8-(9.7)-12p.m.
Subectosomal auxiliary subtylostyles more
slender but longer than principal spicules, straight
or slightly curved towards base, with smooth,
slightly constricted bases, fusiform points.
Length 210-(251.4)-282p,m, width 4-(5.4)-7p.m.
Ectosomal auxiliary subtylostyles relatively
long, straight or slightly curved towards base,
prominently subtylote, with smooth or
microspined bases, fusiform points. Length 82(132.7)-147p.m, width 2-(3.3)-4pLm.
Echinating acanthostyles long, robust, subtylote, large conical (not recurved) spines, scattered mainly over base and midsection of shaft,
frequently with bare point and 'neck'. Length
68-(76.3)-88p.m, width 6-(8.2)-11p,m.
Microscleres. Palmate isochelae small, frequently contort, single size category, with long lateral
alae completely fused to shaft and entire front ala
of equal length. Length 6-(10.8)-14p.m.
Toxas short, robust, wing-shaped, with
generous angular central curvature, recurved and
slightly reflexed points Length 27-(41.8)-58p.,m,
width 1.5-(1.8)-2.0p.m.
REMARKS. This species is relatively poorly
known, recorded so far only from the holotype.
But several important characters were seen in the
holotype using SEM (Fig. 145), not described by
Ridley (1884a). These include: presence of differentiated principal and auxiliary spicules,
presence of toxas, spination pattern on acanthostyles, and his spicule dimensions were innaccurate. These characters now provide better clues
as to its affinities.
Ridley (1884a) considered this species similar
to C. (T) cactiformis in growth form and skeletal
structure. They also both have principal and
auxiliary megascleres very similar in geometry
(i.e., on first appearances fibres seem to be cored
only by auxiliary spicules), but they have many
morphological differences. Clathria (T)arborescens has a substantially better developed ectosomal peel occupying a significant proportion
of branch diameter, a close-meshed reticulate
MEMOIRS OF THE QUEENSLAND MUSEUM
292^
zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 144. Clathria (Thalysias) arborescens (Ridley) (holotype BMNH1881.10.21.272). A, Choanosomal
principal subtylostyles and bases. B, Subectosomal auxiliary subtylostyle. C, Ectosomal auxiliary subtylostyles.
D, Echinating acanthostyle. E, Wing-shaped toxa. F, Palmate isochelae. G, Section through peripheral skeleton.
H, Australian distribution. I, Holotype.
choanosomal skeleton, very thick fibres all cored
by megascleres (C. (T) cactifonnis has a relatively thin ectosomal skeleton, cavernous subectosomal region, wide-meshed choanosomal
reticulation, substantially thinner fibres with only
primary ones cored). Also, echinating acanthostyles are much larger than those of cactiformis,
they have erect conical spines rather than hooklike recurved ones, and acanthostyles are found
throughout the skeleton rather than mainly on
peripheral fibres (although both species have
spines absent from both "necks" and points of
acanthostyles). Toxa morphology differs markedly between the two, with arborescens having only
thicker, wing-shaped toxas, and cactiforniis with
mainly very thin, raphidiform, sinuous or accolada toxas but occasionally also with small
wing-shaped forms. Spicule dimensions also differ considerably between the two species. Both
species belong to the juniperina' species complex (refer to discussion below under C. (T)
cactifonnis and C. (T.) hirsuta).
Dendy (1922), Burton & Rao (1932) and Burton (1938a) merged C. arborescens with C. (T)
procera on the basis that choanosomal principal
megascleres were supposedly excluded from the
axis of fibres, but this is not entirely correct (in
any case the character is also known to occur in
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
293
FIG. 145. Clathria (Thalysias) arborescens (Ridley) (holotype BMNH1881.10.21.272). A, Choanosomai
skeleton. B, Fibre characteristics. C, Echinating acanthostyle. D, Acanthostyle spines. E, Base of ectosomal
auxiliary subtylostyle. F, Wing-shaped toxas. G, Palmate isochelae.
294zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
at least eight other species (see below under C.
cactiformis)). Thiele (1903a) also compared C.
(T) arborescens and C. (T) cervicornis, for the
same reason (i.e., supposed absence of principal
spicules), but C. procera does have principal
spicules (exclusively echinating peripheral
fibres), and those in C. (T) arborescens are intermingled with auxiliary subtylostyles within
fibres. Skeletal structure and growth forms also
differ considerably between these two taxa.
Clathria (Thalysias) cactiformis (Lamarck, 1814)
(Figs 146-148, Table 31, Plate 4E-F)
Spongia cactifortnis Lamarck, 1814: 440, 370
Clathria cactiformis; Rudman & Avern, 1989: 335;
Hooper & Wiedenmayer, 1994: 268.
Spongia appendiculata, in part, Lamarck, 1814: 383,
362.
Wilsonella appendiculata; Topsent, 1930: 46, p1.2,
fig.3; Topsent, 1933: 22.
Echinonema typicum Carter, 1875: 195 [nomen
nudum]; Carter, 1881: 362, 377, 388, 384.
Thalassodendron typica; Lendenfeld, 1888: 223;
Hallmann, 1912: 242; Hal!mann, 1914a: 267.
Not Thalassodendron typica; Whitelegge, 1901: 86.
Clathria typica; Vosmaer, 1885b: 357; Ridley &
Dendy, 1887: 153; Dendy, 1889c: 11;[in part]
Dendy, 1896: 32; Whitelegge, 1901: 80, 117;
Hentschel, 1911: 280, 364-367, text-fig. 43.
Not Clathria ypica var. porrecta Hentschel, 1912:
298, 359-360.
Not Clathria typica; Kirkpatrick, 1903: 248-249;
Kirkpatrick, 1904: 148; Vacelet & Vasseur, 1971:
94-95, p1.3, fig.5, text-fig. 48.
Wilsonella typica; Hallmann, 1912: 203, 242.
Not Tenacia typica var. porrecta Hallmann, 1920: 771.
Rhaphidophlus typicus; Hallmann, 1912: 176-177,
184-203, 234, 299, 300, p1.27, p1.28, figs 1-4, p1.29,
text-figs 38-42 [et yard; Shaw, 1927:424-425; Topsent, 1932:88, p1.4, fig.4; Burton, 1938a: 12; Guiler,
1950: 8; Vacelet & Vasseur, 1977: 115; Vacelet et
al., 1976: 73; Liaaen-Jensen et al., 1982: 170-172.
Echinonema anchoratum Carter, 1881a: 362, 379, 380;
Lendenfeld, 1888: 219-220.
Echinonema anchoratum var. dura Whitelegge, 1901:
81.
Echinonema anchoratum var. lamellosa; Lendenfeld,
1888: 219-220.
Rhaphidophlus anchoratum; Bergquist & Fromont,
1988: 112; Dawson, 1993: 39.
Wilsonella anchoratum var. lamellosa; Hall mann,
1912: 211 [imperfectly known].
Not Echinonema anchorata var. lamellosa;
Whitelegge, 1901: 82.
Echinonema flabelhformis Carter, 1885f: 352.
Echinonema pectiniformis Carter, 1885f: 353.
Phakelliaventilabrum var. australiensis Carter, 1886g:
379.
Thalassodendron brevispina Lendenfeld, 1888: 225;
Whitelegge, 1901: 87.
Clathria favosa Whitelegge, 1907: 498-499.
Clathria indurata Hallmann, 1912: 299.
Clathria acanthodes Hentschel, 1911: 367-370, text-
fig.44; Hentschel, 1923: 380, fig.349.
Rhaphidophlus acanthodes; Hallmann, 1912: 177.
Rhaphidophlus sp. 2; Vacelet & Vasseur, 1971: 97-98,
text-fig. 52.
cf. Microciona proltfera; Vosmaer, 1935a: 609, 628629, 647, 667.
MATERIAL. LECTOTYPE: MNHNDT580:
'Southern Seas', Peron & Lesueur collection, no other
details known. PARALECTOTYPE: MNHNDT3360:
same details (= R. typicus var. cactiformis; Topsent,
1932: 88). LECTOTYPE of S. appendiculata:
MNHNDT526 (fragment BMNH1953.4.9.83) (dry).
PARALECTOTYPE of S. appendiculata: MNHNDT3394: King George Sound (Albany), WA. LECTOTYPE of T. brevispina: AMZ931: Port Jackson,
NSW, 33°48'S, 151°20'E, coll. R. von Lendenfeld
(dredge). SYNTYPE of E. typicum: BMNH1877.5.21.149: Fremantle, WA, 32°03'S, 115°38'E,
coll. E. Clifton (dredge). HOLOTYPE of C. favosa:
AMZ944: Off Port Jackson, NSW, 33°50'S, 151°30'E,
98-100m depth, coll. FIV 'Thetis' (dredge). LECTOTYPE of R. typicus var. proximus: AMZ930: Henley Beach, St. Vincent Gulf, SA, 34 455, 137°57'E,
coll. T. Worsnop (dredge). PARATYPE of R. typicus
var. obesus: AMZ937: Tuggerah Beach, NSW,
33°18'S, 151°30'E, coll. NSW Fish Commission
(trawl). HOLOTYPE of R. typicus var. geminus:
AMZ928: specific locality unknown, WA, no other
details known. HOLOTYPE of R. typicus var. stellifer
AME648: E. coast of Hinders I., Bass Strait, Tas,
40°01'S, 148°02'E, no other details known. SYNTYPE of E. anchoratum: BMNH1886.12.15.423: Port
Phillip, Vic, 38°09'S, 144°52'E, no other details
known. SYNTYPE of E. anchoratum var. dura and
LECTOTYPE of C. indurata: AMG9113: specific
locality unknown, WA, no other details known.
PARALECTOTYPE of C. indurata: AM unreg.:
specific locality unknown, WA, no other details
known. HOLOTYPE of E. pectiniformis: BMNH1886.12.15.141 (fragment AMG2777): Port Phillip,
Vic, 38°09'S, 144°52'E, 40m depth, no other details
known. Fragment of HOLOTYPE C. acanthodes:
ZMB4443: Surf Point, outer bar exit to South Passage,
Shark Bay, WA, 26°08'S, 113°08.5'E, 0.5-3.5m depth,
coll. W. Michaelsen & R. Hartmeyer (dredge). OTHER
MATERIAL. NSW - QMG301403. TAS - AME624,
AMZ1415, AMZ2203, QMG311412 (NCIQ66C3713-A) (fragment NTMZ3811). VICTORIA AMZ1430, AMZ1158, NMV RN359, NMVRN43 I,
NMVRN436, NMVRN 438, NMVRN551, NMVRN677, NMVRN797, NMVRN840, NMVRN900,
NM V RN959, AMZ3921 (RRIMPFN3527/000/02),
AMZ4277 (RRIMPFN1906), NTMZ2832. S AUST AMZ41, SAM unreg. (fragment NTMZ1657), SAM
unreg. (fragment NTMZ 1693), SAM unreg. (fragment
NTMZI628), SW COAST, WA - MNHN unregistered
0
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUT
295zyxwvutsrq
A^BzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 146. Clathria (Thalysias) cactiformis (Lamarck) (lectotype MNHNDT580). A, Choanosomal principal
styles. B, Subectosomal auxiliary subtylostyles. C, Ectosomal auxiliary subtylostyles. D, Echinating acanthostyles. E, Sinuous and wing-shaped accolada toxas. F, Palmate isochelae. G, Section through peripheral skeleton.
H, Australian distribution.
(Peron & Lesueur collection), WAM656-81(1),
WAM628-81(1), WAM631-81(1), WAM655-81(1).
HOUTMAN ABROLHOS, WA - QMG300731
(NCIQ66C-4213-U), QMG3046 60, AMZ4331
(RRIMP-ZW62), WAM639-81(1), WAM661-81(1),
W A M660-81, QMG300736 (NCIQ66C -4189-T),
QMG300733 (NCIQ66C-4197-C), QMG300730
(NCIQ66C-4214-V). NORTHWEST SHELF, WA WAM133-82, NTMZ1128, NTMZ3018, NCIQ66C1460-C (fragment NTMZ3347). ERITREA, RED SEA
- PIBOCB20-125 (fragment QMG300055).
SOMALIA, W. INDIAN OCEAN - PIBOCB12-388
(fragment QMG300060). SEYCHELLES - PIB0001140 (fragment QMG300063).
HABITAT DISTRIBUTION. Colonises a wide variety
of substrates, including rock reefs, coral-rubble, soft
sand, mud sediments, algal beds; shallow subtidal to
100m depth; widely distributed in temperate waters of
eastern and western Australian coasts, with incursions
into subtropical waters extending to the Port Hedland
region on the west coast and Byron Bay on the east
coast (Fig. 146H). This species is not yet recorded from
the Indonesian archipelago, contrary to Hentschel
(1912) — his records being misidentifications (see
below) — but it does appear to range across the Indian
Ocean, with confirmed records from Madagascar
(Vacelet et al., 1971, 1976, 1977), East Africa and
Seychelles (present study).
DESCRIPTION. zyxwvutsrqponmlkjihgfedcbaZY
Shape. Typically flabellate,
planar, stalked, with long bifurcate, occasionally
anastomosing digitate processes on margin of
fan; gross morphology variable, with or without
basal stalk and expanded basal attachment, varying from globular, cylindrical planar digitate, globular
digitate, planar flabellate, lamellate branching in
more than one plane, or thinly lamellate.
Colour. Live colouration pale orange-brown
(Munsell 5YR8/4) to bright red (5R5110); beige,
^ MEMOIRS OF THE QUEENSLAND MUSEUM
296zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
FIG. 147. Clathria (Thalysias) cactiformis (Lamarck) typical growth forms. A, Lectotype MNHNDT580. B,
Paralectotype MNHNDT3360. C, Lectotype of S. appendiculata MNHNDT526. D, Paralectotype of S.
appendiculata MNHNDT3394. E-F, Variety cactiformis WAM656-81(1), WAM655-81(1). G, Variety stellifera NTMZ1708. H, Variety geminus WAM639-81(1). I, Somalia specimen PIBOCB12-388. J, Syntype of
E. typicum BMNH1877.5.21.149. K, QMG300453. L,Variety obesus with Rostanga feeding (photo W.
Rudman) NTMZ2832.
^
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSR
297
FIG. 148. Clathria (Thalysias) cactiformis (Lamarck) (QMG300225). A, Choanosomal skeleton. B, Fibre
characteristics. C-D, Echinating acanthostyles. E, Acanthostyle spines. F-G, Bases of subectosomal and
ectosomal auxiliary subtylostyles. H-I, Palmate isochelae. J, Accolada toxas. K, Wing-shaped toxas.
298zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
^ MEMOIRS OF THE QUEENSLAND MUSEUM
skeletal components; fibre anastomoses produce
widely spaced rectangular meshes formed by ascending primary fibres, containing pauci- or multispicular bundles of choanosomal principal
styles which occupy only the central portion of
fibres, and uncored secondary fibres; occasionalSpecimens
S.zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Lectotype of zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
Lectotype
SPICULE
ly secondary fibres uni- or paucispicular, usually
(N=-46)
appendiculat a
with an optically diffuse pith reminiscent of dic103.6(199.2)Choanosomal 134(191.2)157-181 x
tyoceratid sponges; spongin fibres charac314 x 3.2232 x 5-(7.6)principal
4.5-6
styles
(9.7)-19.4
11
teristically thin, without marked differences
between primary and secondary fibre diameter,
11534235.7)Subectos°mal 2 044229.8)191-297x
356.6 x 1.6auxiliary
with pigmentation ranging from extremely light,
4-5.5
8 x 4-(5.4)-6
styles
(6.2)-16.9
yellow (specimens from Port Phillip, Vic) to ex73.8(114.7)101(124.3)tremely heavy and dark brown (material from
Ectosomal
85-104 x 2-3.5
162.1 x 1.6148x 1.5auxiliary
Port Hedland, WA); echinating acanthostyles
styles
(4.4)-13.6
(2.5)-3
very heavy, especially on peripheral fibres imme38.2-(61.3)diately
below subectosomal skeleton; larger
68-(74.4)-79
x
Echinating
93.1 x 1.746-57 x 3-4.5
acanthostyles 5.5-(8.8)-11
auxiliary subtylostyles usually abundant between
(6.7)-13.1
fibres, strewn without order within mesohyl;
8.5-(13.9)-22.2
7-8
12-(14.7)-18
Chelae I
mesohyl matrix relatively light, choanocyte
2.5-(7.1)-11
15-17
6-(8.8)-I0
Chelae II
chambers large, oval or elongate, up to 12011m
1134201.7)-296 82-146 x 0.5- 8-(127.8)-355
diameter.
Toxas
x 0.8-(1.3)-2.5
x 0.8-(1.2)-2
1.0
Megascleres (Table 31). Choanosomal principal
styles morphologically similar to larger auxiliary
brown, to dark brown in ethanol; ectosomal crust subtylostyles but relatively shorter, thicker,
usually lighter colour than choanosomal regions. slightly curved near base, usually with tapering,
Oscules. Oscules typically large (up to 3mm dia- evenly rounded smooth bases but sometimes
meter), dispersed over entire sponge surface, or slightly subtylote, and fusiform points.
sometimes clumped in special pore areas; oscules
Subectosomal auxiliary subtylostyles nearly
usually flush with surface, without raised lip.
completely straight, occasionally slightly curved
Texture and surface characteristics. Firm, com- or sinuous, usually with slightly subtylote smooth
pressible, rubbery; surface characteristically bases, less frequently with microspined bases,
fleshy, smooth but uneven, with bumps, ridges, tapering to fusiform points.
depressions and subdermal striations commonly
Ectosomal auxiliary subtylostyles more
dispersed over surface; opaque skin-like covering prominently subtylote, slightly curved at centre,
present stretched between adjacent conules in with microspined or smooth bases and fusiform
live material; microscopic ridges and points.
microconules abundant.
Echinating acanthostyles relatively short,
Ectosome and subectosome. Ectosomal skeleton stout, prominently subtylote, with large spines
predominantly heavy, with a dense crust of mainly dispersed on base and midsection, leaving
smaller auxiliary subtylostyles forming mainly a bare 'neck' (below basal swelling) and bare
erect or less often paratangential brushes; oc- point.
casionally ectosomal skeleton reduced to rela- Microscleres (Table 31). Larger palmate
tively thin tangential or paratangential crust; isochelae variable in abundance, with lateral alae
larger subectosomal auxiliary styles perched on completely fused to shaft, wide gap between
ultimate choanosomal fibre endings, forming lateral and front alae, and fused front ala; usually
plumose tracts underlaying brushes of smaller non-contort. Smaller isochelae common, freectosomal spicules, usually protruding through quently contort.
and obscuring ectosomal bundles; subectosomal
Toxas differentiated into two main morphs,
region cavernous, with well differentiated, wide- larger often found in toxodragmata; accolada
ly spaced plumose tracts of larger auxiliary sub- toxas most common, long, very thin, slight antylostyles.
gular central curvature, straight arms, sometimes
Choanosome. Skeleton irregularly reticulate, sinuous asymmetrical. Small toxas less abundant,
vaguely radial, with clearly differentiated relatively thick, widely curved at centre, slightly
primary ascending and secondary transverse reflexed points.
TABLE 31. Comparison between present and published records of Clathria (Thalysias) cactiformis
(Lamarck). All measurements are given in p.m,
denoted as range, or as range (and mean) of spicule
length x spicule width (N=25).
REVISION OF MICROCIONIDAEzyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQ
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299
Larvae. Reproductive products observed in only
four specimens. Incubated parenchymella larvae
large, oval to eliptical in shape, 220-420Rm
diameter, containing well developed centrally aggregated larval styles.
Associations. Heavy infestations of zoophytic organisms observed on surfaces of several
specimens (both compound ascidians and zooanthid polyps); numerous microalgal cells seen in
sponge mesohyl of few samples; sponge apparently serves as a food source to Rostanga sp.
nudibranchs (W. Rudman, pers. comm.).
Variation. Extremely variable in some features
(e.g., gross morphology, spicule size) but not in
others (e.g., surface features, spicule geometry,
skeletal structure). Gross morphology: Victorian,
Queensland and Indonesian morphs showed
greatest differences in growth form, but all
specimens retained most other characteristics
common to other populations. Ectosomal skeletal
structure: typically heavy, erect plumose brushes,
well differentiated from subectosomal plumose
tracts (52%), moderately developed (34%), or
thick tangential ectosomal peel reminiscent of
Crella (Crellidae) (14% of specimens). Subectosomal skeletal structure: very cavernous, well
differentiated ascending plumose (extra-fibre)
skeletal tracts (36%), paratangential, immediately subdermal tracts (21%), or intermediate of
these conditions (43% of specimens).
Choanosomal skeletal structure: irregularly
reticulate (64%), regular fibre anastomoses
(22%), or predominantly radial to plumose
primary fibre architecture (14% of specimens).
Fibre meshes: wide, rectangular, with uncored
secondary elements (67%), with central pith component (7%), close-meshed, irregularly oval to
eliptical fibre reticulation with uncored secondary fibres (7%), or with both cored primary and
secondary fibre elements (19% of specimens).
Spongin fibres: very light (7%), moderately
heavy, yellow (48%), very heavy, yellow (43%),
or extremely heavily, dark brown in colour (2%
of specimens). Echinating acanthostyles: concentrated on peripheral fibres, more sparsely
dipersed on axial fibres (69%), evenly distributed
throughout fibres (17%), rare (14% of
specimens). Extra-fibre spicules: typically abundant, dispersed without order within mesohyl
(74%), very light or absent entirely (24%), or
organised into ascending extra-fibre tracts (2% of
specimens). Megasclere geometry: Choanosomal
principal styles: predominantly fusiform (76%),
distinctly hastate (abruptly pointed) points (7%),
or mixture of both types of terminations (17% of
specimens); bases of spicules predominantly
smooth (90%), with both smooth and spined
bases (8%), or only microspined bases on
choanosomal styles (2% of specimens); bases of
spicules tapering, non-tylote (46%), rounded,
slightly subtylote (52%), or with both conditions
(2% of specimens); megascleres slightly curved
(83%), exclusively straight (7%), or both conditions (10% of specimens). Subectosomal
auxiliary subtylostyles: tapering, rounded or very
slightly subtylote bases (69%), or distinctly subtylote (31% of specimens); exclusively straight
(69%), predominantly curved (7%), or both conditions (24% of samples). Ectosomal auxiliary
subtylostyles: subtylote (76%), tapering nontylote (10%) or both conditions (14% of
specimens); bases predominantly microspined
(71%), exclusively smooth (3%), or with both
conditions (26% of specimens); ectosomal
spicules slightly curved near base (54%), completely straight (29%), or with examples of both
(17% of specimens); most megascleres with
fusiform points (95%), or also including few hastate spicules (5% of specimens). Echinating acanthostyles: typically short, fusiform (69%),
significantly smaller, stouter (19%), or longer,
thinner (12% of specimens); bases subtylote
(81%), or distinctly non-tylote (19% of
specimens); spines characteristically long, thick,
recurved (74%), or reduced, granular spination
(26% of specimens). Microsclere geometry:
Larger isochelae: common (70%), uncommon
(26%), rare (2%), or common but poorly silicified
(2% of specimens); contort larger isochelae
present but uncommon (10%), present, common
(2%), or absent (86% of specimens). Smaller
isochelae: abundant (7%), common (62%), uncommon (21%), rare (3%) or absent entirely (7%
of specimens); 0-10% of smaller chelae were
contort (2%), 10-30% contort (5%), 30-50% contort (12%), 50-70% contort (24%), 70-90% contort (43%), or 100% contort (7% of specimens).
Toxas: typically very abundant (82%), common
(3%), uncommon (5%), rare (3%), or absent entirely (7% of specimens); only symmetrical toxas
present, with large central curvature and relatively straight points (52%), with both symmetrical
and asymmetrical, sinuous toxas (41%), or with
both these and wing-shaped toxas (7% of
specimens).
Variability in spicule dimensions: With few exceptions, and despite apparent morphological incongruence in many other features, spicule
dimensions were surprisingly closely comparable
between specimens despite that they were col-
300zyxwvutsrqponmlkjihgfedcbaZYXWVUTSRQPONMLKJIHGFEDCBA
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lected from widely separated geographic
localities. Results were not analysed statistically
because bathymetric, habitat and season data
were not available for the majority of material.
However, scatterplot comparisons between
samples found that for each spicule category
there were only few anomolous specimens which
consistently differed from typical forms of the
species. These anomalies are discussed further
below.
REMARKS. Clathria (Thalysias) cactiformis is
predominantly temperate. It is characterised by
its typically flabellate or flabello-digitate growth
form, a smooth surface with a detachable thick
ectosomal peel which is usually paler than the
choanosome, relatively thin spongin fibres heavily echinated but poorly cored, and especially its
short, thick, squat acanthostyles with a bare neck,
bare points and large recurved spines over the
remainder of the spicule. There is considerable
morphological variability between regional
populations (see Variation, above), to some extent vindicating Hallmann's (1912) subdivision
of the species into 'varieties' (subspecies), based
mainly on differences in growth form. In fact
some specimens examined were assigned to this
species with hesitation given their diverse external morphologies, but no other consistent skeletal
or other characters were found to correlate with
these morphotypes. Nevertheless, this taxon is
probably composed of several cryptic sibling
species (morphospecies) which might only be
reliably differentiated using biochemical, genetic
and other non-skeletal data.
NMVRN0436 and RN0551 from Port Phillip
Bay, Vic. have acanthostyles which are long,
evenly spined, spines are sparse and small;
echinating spicules occur in heavy concentrations
and are evenly dispersed throughout the
choanosome, not predominant in the peripheral
skeleton; ectosomal skeleton is very well
developed and well differentiated from subectosomal plumose tracts; the subectosomal region
is noticeably more cavernous than in most typical
forms; the choanosome is distinctly plumoreticulate; mesohyl matrix is very heavy and
fibres are heavily invested with spongin; the
smaller category of isochelae and toxas are absent. In some respects these features correspond
to Spongia appendiculata, perhaps justifying the
retention of that species separately form cactiformis, but other general morphological features
indicate that these specimens cannot be otherwise
reliably separated.
WAM133-82 from the Northwest Shelf of WA
has consistently basally spined choanosomal
styles, and isochelae are completely unmodified.
The ectosome and subectosomal regions are
poorly developed and incompletely differentiated, but the specimen is otherwise comparable
with typical forms.
Some further comment is necessary on the
synonymy above. Spongia appendiculata is included here as a new synonym of C. (T) cacnformis (cf. Hooper & Wiedenmayer, 1994) with
virtually identical growth form, spicule geometry
and skeletal structure as typical forms of this
species. However, there are some notable differences between these nominal species, especially in the markedly smaller dimensions of most
spicules in appendiculata, acanthostyles have a
vestigial granular spination, and they are also
almost entirely incorporated into the fibres themselves (i.e., enveloped by collagen). These differences are not presently considered sufficient to
recognise it as a distinct taxon. Incorporation of
echinating acanthostyles into spongin fibres is
also known for several other species of Clathria
(termed here the `phorbastformis' species complex, including C. (T) phorbasifonnis, C. (D.)
dura, C. (D.) myxilloides, and others). However,
in Spongia appendiculata acanthostyles are inserted perpendicular to fibres, much like typical
echinating megascleres, but they have 'sunk' into
them, whereas in these other species acanthostyles run parallel to fibres and replace or supplement the usual coring spicules.
Clathria acanthodes is considered a synonym
of C. (T) cactifonnis, despite having slightly
different acanthostyle morphology ('neck' often
spined). The characters used by Hentschel (1911)
to differentiate C. acanthodes from other species
(i.e., similar geometry between principal and
auxiliary spicules, presence or absence of
microspines on bases of principal and auxiliary
spicules, presence of two size classes of palmate
isochelae, and possession of rhaphidiform toxas),
are found in most other populations of C. (T)
cactifonnis and are not considered to be of
primary importance in differentiating species.
Clathria typica var. porrecta, also described by
Hentschel (1912) from Indonesia, is a synonym
of Clathria (T) reinwardti not of C. (T.) cactiformis (this syonymy was overlooked by Hooper &
Wiedenmayer, 1994). The type material of
Clathria indurata and Echinonema anchoratwn
var. dura from WA have a uni- or paucispicular
core of choanosomal styles in secondary fibres,
whereas typical populations of C. (T) cacnformis