AUSTRALIAN HALICHONDRIIDAE (PORIFERA: DEMOSPONGIAE):
I. SPECIES FROM THE BEAGLE GULF.
JOHN N.A. HOOPER, STEPHEN D. COOK, LISA J. HOBBS AND JOHN A. KENNEDY
Queensland Museum, PO Box 3300, South Brisbane, Qld 4101, Australia.
ABSTRACT
Twenty species of sponge in seven genera of the family Halichondriidae (order
Halichondrida) are described from the Beagle Gulf, Darwin region, Northern Territory, including eight new species and five new records for Australia. A resume is
provided of generic characters and a survey of the known Indo-west Pacific species in
these genera.
Porifera, Demospongiae, Halichondriidae, Axinyssa, Myrmekioderma,
Topsentia, Ciocalypta, Amorphinopsis, Halichondria, Hymeniacidon, taxonomy, new
KEYWORDS:
species, northwest Australia.
subsequently included in the order (Halichondriidae, Axinellidae, Dictyonellidae and
Desmoxyidae), although there is not yet complete agreement about the generic composition
of each of these (see Hooper and Levi 1993b).
In its initial conception, Halichondriidae contained 27 very disparate genera allegedly related to each other by their possession of "fusiform
or pin-shaped spicules variously fasciculated
together" (Gray 1867: 503). Apart from Halichondria and Ciocalypta, all the other genera
included here by Gray (1867) were subsequently assigned by contemporary authors to Poecilosclerida, Hadromerida or Haplosclerida. Nevertheless, the family name remains valid for the
current assemblage of genera containing the
genus Halichondria (Article 40 ICZN; Anon.
1985). Few subsequent authors used the family
name Halichondriidae until it was redefined by
de Laubenfels (1936) in a contemporary classification, after which time it came into general
acceptance (e.g. Levi 1973; Berquist 1978; Hartman 1982). De Laubenfels' (1936: 133) interpretation of the family included eight genera, of
which several were clear synonyms, all with
"exceedingly simple spiculation ... of ... only
oxeas ... smooth ... without microscleres ... [with]
special dermal skeletons, although these are not
conspicuous" (i.e. it specifically excluded sponges like Hymeniacidon with principal styles rath-
INTRODUCTION
Of the 114 families of extant Porifera only
two (Raspailiidae and Microcionidae, order
Poecilosclerida) can so far be considered well
known in the Australian fauna (Hooper 1991,
1996). In this present work we deal with a third
family, Halichondriidae, in the order Halichondrida.
Traditional definitions of Halichondrida were
based on the possession of diactinal megascleres,
a relatively homogenous skeletal structure described as `halichondroid' (criss-cross reticulate), viviparous reproduction and completely
ciliated larvae (Bergquist 1978; Hartman 1982).
Under this interpretation two families were differentiated, Halichondriidae Vosmaer, 1887
(principal megascleres oxeas sometimes also
with accessory styles), and Hymenacidonidae
de Laubenfels, 1934 (principal megascleres styles
sometimes with accessory oxeas), (Bergquist
1978). A recent revision of the order by Van
Soest et al. (1990) allowed for the inclusion of
species with compressed axial skeletons, differentiated axial and extra-axial skeletons (i.e. traditional Axinellidae), an indiscriminate occurrence of styles and oxeas within the skeletons,
as well as 'typical' halichondrids with skeletal
arrangements of styles or oxeas in plumose to
ill-defined, confused tracts. Four families were
In: Hanley, J.R., Caswell, G., Megirian, D. and Larson, H.K. (eds) Proceedings of the Sixth International Marine Biological
Workshop. The marine flora and fauna of Darwin Harbour, Northern Territory, Australia. Museums and Art Galleries of
the Northern Territory and the Australian Marine Sciences Association: Darwin, Australia, 1997: 1-65.
1
J.N.A. Hooperei al.
collected from this region (extending from
Northwest Cape, WA, to the Wessel Islands,
NT; Hooper unpublished data), but so far less
than 300 of these have been named in the literature (Hooper and Wiedenmayer 1994). Many
species are unique to this region, with levels of
endemism in some families thought to be about
70% (Hooper and Levi 1994), although for most
sponge families this fauna is still poorly known.
Habitats within this region are also incredibly
diverse, most with at least some sponge species
represented (ranging from ephemeral pockets
of freshwater, large rivers, euryhaline estuaries
and extensive mangroves, mud flats, fringing
intertidal and subtidal rock and coral reefs,
seagrass and Halimeda beds, artificial substrates,
sand, coral rubble and shell-grit bottoms, deeper
offshore rock reefs, submerged 'active' coral
reefs, as well as pristine emergent platform and
fringing coral reefs).
There have been several major publications on
sponges within the Beagle Gulf but prior to the
present work only 71 species have been described. The earliest authenticated record of this
fauna comes from material collected between
1875 and 1877 by T.J. Sturt, which was sent to
Melbourne and initially described as a 'curiosity'
(Barnard 1879), but subsequently identified by
Carter (1879) as an endemic genus and species,
Axos flabelliformis Carter (Hooper 1986a). In
1881 the British survey ship HMS Alert made
extensive collections of the fauna, mostly dredging shallow waters within Darwin Harbour, Van
Diemen Gulf, Dundas Strait and the Timor Sea.
The sponges from this collection were described
by Ridley (1884), and included 24 species, 21
named and three left uncertain, of which only 17
are currently recognisable as valid species. Nine
of these species have not since been recorded
elsewhere (i.e. apparent endemics).
In the early 1960's a team from the Australian Museum, Sydney, led by Elizabeth Pope,
travelled to Darwin, Cape Don and Port
Essington to survey the intertidal marine biology of this region. This collection is the earliest
comprehensive attempt to document the marine
invertebrates of the northwest Australian coast
(Pope 1967), from which the sponge fauna was
described separately (Bergquist and Tizard
1967). These authors described 19 species of
the larger, more prevalent intertidal sponges
found in this region, belonging to 17 genera and
with three new species.
Since 1967, there have been 37 additional
species in 16 genera described from the Beagle
er than oxeas and a fleshy, gelatinous consistency). De Laubenfels (1936) further characterised
the family in having "subdermal cavities ... so
very enormous that the dermis is merely a roof
supported on slender columns ... above ... the
endosome".
Since its creation, 54 genera have been associated with the Halichondriidae, although the
most recent revision by Van Soest et al. (1990)
(with subsequent refinements by Diaz et al.
(1991, 1993) and Pomponi et al. (1991)), now
recognises only 12 of these (the remainder being junior synonyms or incorrect assignments):
Amoyhinopsis Carter, Axinyssa Lendenfeld,
Ciocalvta Bowerbank, Collocalypta Dendy,
Didiscus Dendy,Epipolasi.s de Laubenfels,Halichondria Fleming, Hymeniacidon Bowerbank,
Myrmekioderma Ehlers, Petromica Topsent,
Spongosorites Topsent and Topsentia Berg. All
these genera have a highly dense choanosomal
spicule skeleton arranged in vague, directionless
(halichondroid) tracts composed of spicules in
confusion (Van Soest et al. 1990), with spicules
being either principal oxeas or styles, sometimes also with smaller accessory styles or oxeas
(i.e. including Hymeniacidonidae and some
Axinellidae). The Halichondriidae now represents one of the few spicule-bearing (`nonkeratose) families of Porifera 'accessible' to
contemporary systematics.
So far only 28 species in eight of the 12
halichondriid genera have previous published
records from Australian territorial waters (including Antarctica) (Hooper and Wiedenmayer
1994), and surprisingly only two of these came
from the northwest Australian region.
In the present work we describe 20 species in
seven genera from the Beagle Gulf, of which
five are new records for Australia and eight are
new species. A second contribution describing
halichondriids from other tropical Australian
provinces is currently in progress (Hooper et al.
in prep.).
DARWIN SPONGE FAUNA
In this forum (i.e. The International Workshop on the Flora and Fauna of Darwin Harbour) it is appropriate to summarise existing
knowledge on the Darwin sponge fauna of the
greater region, namely the Beagle Gulf.
The sponge fauna of northwest Australia is
possibly the most diverse of all Australian marine provinces, with about 800 species already
2
Beagle Gulf halichondriid sponges
Gulf region (Darwin Harbour in particular), as
well as from other parts of the NT and northern
WA coasts, including notes on the ecology, biochemistry and/or biogeographical relationships
of several species (Hooper 1984, I986a, 1986b,
1987, 1991, 1994; Hooper and Bergquist 1992;
Hooper and Levi 1993a, 1993b, 1994; Hooper et
al. 1990, 1991, 1992; Capon et al. 1986; Capon
and MacLeod 1988; Cambie et al. 1988a, 1988b;
Van Soest et al. 1991). Significantly, Darwin
Harbour is the type locality for 22 of these
species, most discovered originally from the
East Point Aquatic Reserve. Nevertheless, this
described fauna represents less than 10% of the
species known to live in the Beagle Gulf, and it
is anticipated that this list of published species
will increase substantially over the next decade.
ate spicules not functionally localised to any
particular region of skeleton. Spicules widely
diverging in size in some taxa, sometimes with
size differences between ectosomal and
choanosomal skeletons. Skeletal structures ranging from disorganised plumoreticulate, crisscrossed "halichondroid skeleton", to distinctly
compressed axis (or basal) skeleton and differentiated extra-axial (radial, plumose or plumoreticulate) skeleton. Spongin fibres usually poorly
developed or absent. Ectosomal skeleton organised into tangential layer of spicules or erect
spicule bundles, with minimal collagen, and
large subectosomal spaces in many taxa.
Microscleres raphides, microxeas, or spined
microxeas with central bend in some species.
Family Halichondriidae Gray
METHODS
Halichondriadae Gray, 1867: 503.
Halichondriidae; Vosmaer, 1887: 336. - de
Laubenfels 1936: 133; - Van Soest et al. 1990:
20.
Ciocalyptidae Hentschel, 1923: 408.
Spongosoritidae Topsent, 1928: 35,157.
Petromicidae Topsent, 1928: 34,105.
Hymeniacidonidae de Laubenfels, 1934: 13.
Diagnosis. Halichondrida with choanosomal
skeleton consisting of high density of spicules
arranged in vague, ill-defined, directionless
tracts and spicules in confusion. Ectosomal skeletons frequently detachable, often composed of
spicules of different size than those in
choanosome. Spicules are relatively large oxeas
and/or styles, sometimes strongyloxeas or intermediate spicules
Methods of collection, preservation, histological preparation, and techniques for light
microscopy and scanning electron microscopy
are given elsewhere (e.g. Hooper 1991). Spicule
measurements are based on 25 measurements of
each category of spicule, and represented as
range (and mean). Abbreviations used in the
text are: AM, Australian Museum, Sydney;
BMNH, The Natural History Museum, London;
CCNT, Conservation Commission of the Northern Territory, Darwin; MNHN, Museum National d'Histoire Naturelle, Paris; NCI AIMS,
Australian Institute of Marine Science (US National Cancer Institute collection contract, 19851991); NCI CRRF, Coral Reef Research Foundation, Chuuk State, Federated States of Micronesia (US National Cancer Institute collection
contract, 1992); NT, Northern Territory; NTM,
Northern Territory Museum, Darwin; QM,
Queensland Museum, Brisbane; SMF, NaturMuseum and Forschungsinstitut Senckenberg,
Frankfurt; WA, Western Australia; ZMB, Museum fur Naturkunde an der HumboldtUniversitat zu Berlin.
Genus Axinyssa Lendenfeld
Axinyssa Lendenfeld, 1897: 116.
Pseudaxinyssa Burton, 1931: 350.
Axinomimus de Laubenfels, 1936: 163.
Type species. Axinyssa topsenti Lendenfeld,
1897, by monotypy.
Diagnosis. Massive, subspherical, lobate or
lobate-digitate growth forms, often with apical
cloacal cavity and fistulose surface processes.
Choanosomal skeleton collagenous (although
spongin fibres poorly developed), with widely
spaced spicule tracts cored by larger oxeas,
largely strewn in confusion or slightly plumose,
dendritic or vaguely radiating. Subectosomal
skeleton more organised than choanosomal region, plumose or plumo-reticulate, with regular
tracts of larger choanosomal oxeote megascleres
SYSTEMATICS
Order Halichondrida Vosmaer
Halichondrida Vosmaer, 1887: 335.
Diagnosis. Demospongiae with plumoreticulate skeletal architecture composed of relatively
large oxeas, styles, strongyloxeas or intermedi3
J.N.A. Hooper et al.
Habitat. Sponge bed, soft mud and shell-grit
substrate; subtidal 18-200 m depth.
Description. Massive, lobate, with lumpy appearance, approximately 55 mm diameter.
Oscules large, up to 4 mm in diameter, discrete,
scattered over entire surface (i.e. not confined to
apex of sponge), flush with surface (i.e. without
raised membraneous lip).
Colour. Pale pinkish alive in air (Mansell
5RP 8/2); ectosome purple-grey in ethanol, with
lighter choanosome.
Texture. Firm, resilient, barely compressible; not easily broken.
Surface. Irregular surface with incorporated
detritus; minutely hispid, with series of small
raised ridges running over surface forming
minute reticulation. Cavernous subectosomal
skeleton visible through transparent surface
membrane.
Ectosomal skeleton. Plumose bundles of
larger spicules from subectosomal region protrude through collagenous layer at surface, forming brushes or singly, with sparse tangential
layer of both smaller and larger spicules, lying
singly or in paucispicular tracts, overlaying ascending subectosomal skeleton.
Choanosomal skeleton. Confused halichondroid reticulation of multispicular tracts in
deeper choanosomal region of skeleton, becoming slightly more organised towards periphery;
multispicular choanosomal skeletal tracts composed 20-50 spicules wide, ascending towards
surface, becoming more plumose in subectosomal region, tracts separated by large subectosomal cavities of approximately same width as
tracts themselves. Spongin fibres not evident.
Collagen abundant in choanosomal mesohyl,
associated predominantly with spicule tracts.
Generally all spicule sizes dispersed throughout
ectosomal and choanosomal skeletons, without
clearly differentiated ectosomal and choanosomal
size classes, although smaller spicules mainly
found scattered between tracts whereas larger
spicules mainly aligned in tracts.
Spicules. Oxeas in two size classes, or more
probably one class with large size range: larger
oxeas slightly curved at centre, hastate relatively abruptly pointed (length 550-(622.4)-744
pm, width 13-(16.8)-21 pm); smaller oxeas
straight or slightly curved, hastate abruptly
pointed, occasionally styloid (length 188(241.4)-290 pm, width 4-(6.7)-11 pm).
Remarks. This species has not previously
been recorded having two size classes of oxeas,
although Dendy (1922) notes that the main
ascending to surface; without fibre component
but with heavy collagenous mesohyl. Ectosomal
skeleton highly collagenous, with mineral skeleton composed of smaller oxeote spicules forming discrete, sparse, regularly distributed surface brushes eventually producing surface
conules; or of larger choanosomal oxeotes protruding through surface and producing regularly spaced surface conules. Megascleres are
one or two size categories of oxeas or modified
(styloid, strongyloid) oxeotes.
Remarks. Axinyssa and the related genus
Collocalypta differ from other halichondriids
in having choanosomal spicule tracts protruding slightly through a largely organic ectosome
(i.e. lacking a continuous ectosomal skeletal
crust), producing a finely conulose surface (Van
Soest et al. 1990). Axinyssa differs from the
latter genus by its plumose or vaguely plumoreticulate skeletal structure (particularly near
the surface), and growth form (the latter being
exclusively fistulose). The emended diagnoses
and discussions given by Van Soest et al. (1990)
and Hooper and Bergquist (1992) clearly differentiate the two genera, whereas the diagnosis
provided by Diaz et al. (1991) did not, mainly
because they did not encounter Collocalypta in
the central Atlantic fauna. Worldwide there are
so far 13 described species of Axinyssa, distributed throughout the tropical oceans in shallow
reefs to deeper coastal waters, but only one of
these have been previously described from IndoAustralian waters: A. aplysinoides (Dendy) described from the Great Barrier Reef (Burton 1934).
Axinyssa aplysinoides (Dendy)
(Figs 1-2)
Halichondria aplysinoides Dendy, 1922: 3940, pl. 3, figs 3-5, p1.12, fig. 9.
Trachyopsis aplysinoides - Burton, 1926: 78;
- Burton 1934: 564.
Axinyssa aplysinoides - Van Soest et al., 1990,
fig. 20; - Hooper and Bergquist 1992: 101.
Type material. HOLOTYPE - BMNH
1921.11.7.31: Cargados Carajos, Indian Ocean,
30 August 1905, 58m depth, coll. HMS Sealark.
Material. QM G303561: Outer region of Shoal
Bay, NT, 12°06.9'S, 130°49.9'E, 18 m depth, 12
October 1993, coll. CCNT stn. 136 (ref. 2176),
dredge.
Distribution. Cargados Carajos, Amirante,
Coetivy, Egmont Reef, Seychelles Is, Indian
Ocean; southeast Indonesia; Satellite Reef, Great
Barrier Reef; Shoal Bay, Timor Sea, NT.
4
Beagle Gulf halichondriid sponges
I
111
Fig. 1. Axinyssa aplysinoides (Dendy). A, larger oxeas; B, smaller oxea. C, peripheral skeletal structure;
B, preserved specimen QM G303561; E, live specimen (on deck).
5
J.N.A. Hooperet al.
Fig. 2. Axinyssa aplysinoides (Dendy). Specimen QM G303561. A, Perpendicular section through skeleton showing large
multispicular tracts in choanosomal skeleton, becoming more plumose in periphery; B, transverse section of ectosomal skeleton
showing outer surface of sponge with tangential uni- and paucispicular tracts of both larger and smaller oxeas; C, light micrograph
of section through peripheral skeleton showing plumo-reticulate subectosomal skeletal tracts and protruding larger spicules;
D, hastate pointed spicule terminations on both larger oxeas (left) and smaller oxeas (right).
megascleres "vary considerably in size", with
the upper size limit of 1000 x 30 pm. In the
Darwin specimen, spicules are indiscriminantly
scattered throughout the skeleton, without any
marked localisation of spicule sizes to either
ectosomal or choanosomal regions (unlike those
of Topsentia or Epipolasis which have their
ectosomal skeletons composed of a distinctly
smaller category of spicules), and it is possible
that smaller megascleres seen here (mostly scattered between the major spicule tracts) are juvenile forms of the main structural spicules.
Dendy (1905) suggests that the most distinctive feature of this species is its resemblance to
an Aplysina (Verongida) in external morphology, in particular to the series of minute, raised
ridges forming a reticulation on the surface.
These surface features, together with the ascending multispicular tracts of oxeas cemented
together by granular collagen, and the spicule
geometry (lacking telescoped points, occasion-
ally styloid ends), differentiate this species from
other Axinyssa. We do not know whether material from the Great Barrier Reef recorded by
Burton (1934) is similar to the Darwin specimen, described above, as Burton did not describe his specimen, but it is assumed that
conspecificity of his specimen was checked
against type material given that he had easy
access to the holotype in the BMNH collection.
Genus Myrmekioderma Ehlers
Myrmekioderma Ehlers, 1870: 28.
Acanthoxifera Dendy, 1905: 156.
Anacanthaea Row, 1911: 329.
Neoprosypa de Laubenfels, 1954:189.
Type species. Alcyonium granulatum Esper,
1830: 71.
Diagnosis. Massive or encrusting growth
forms. Surface hispid, with characteristic excavating meandering, sinuous or straight canals
6
Beagle Gulf halichondriid sponges
and grooves (forming polygonal tuberculate
plates in type species). Choanosomal skeleton
collagenous, with compressed central portion
composed of confused mass of larger spicules
forming irregular, ascending, multispicular tracts
bound together with collagen (spongin fibres
present in some species but not all). Subectosomal skeleton composed of ascending tracts running radially or obliquely to surface. Ectosomal
skeleton dense paratangential layer of smaller
spicules forming closely adjacent brushes perpendicular to surface, with larger choanosomal
spicules also protruding. Megascleres at least
two distinct categories of choanosomal oxeas,
strongyles, or more rarely styles, sometimes sinuous and centrally flexed, and smaller ectosomal
oxeas or acanthoxeas of similar morphology.
Microscleres raphides in trichodragmata, in one
or more categories, larger usually sinuous or
curved.
Remarks. Recent revisions of this genus are
provided by Bergquist (1965: 177), Van Soest
et al. (1990: 31), Diaz et al. (1991) and Hooper
and Levi (1993b: 1448). Myrmekioderma and
Didiscus are apparently related (and both easily
recognisable in the field) by their deeply sculptured sinuous and plate-like grooves on the surface, having an ectosome composed of smaller
spicules forming an erect palisade over tangential larger spicules, and a similar basic
choanosomal skeletal organisation (Van Soest
et al. 1990; Diaz et al. 1991). Myrmekioderma
lacks discorhabd microscleres, characteristic of
Didiscus, but has instead bundles of sinuous
raphides (trichodragmata). The genus has been
traditionally defined with lightly acanthose
ectosomal oxeas (e.g. Bergquist 1965; Hooper
and Levi 1993b), but Diaz et al. (1991) correctly note that these ectosomal spicules are as
frequently smooth as they are acanthose, and
they emend the definition accordingly. Hooper
and Levi (1993b) retained this genus in the
family Desmoxyidae, supporting their decision
with chemotaxonomic evidence based on an
inferred close relationship between Higginsia
and Myrmekioderma (Hooper et al. 1992). However, there is no strong morphological support
for homology between the occasionally acanthose
ectosomal oxeas in Myrmekioderma and the
true ectosomal acanthoxeas in Higginsia. The
overall similarity between the ectosomal structure, choansomal organisation and spicule composition (apart from the possession of microxeas
or discorhabds, respectively) largely supports
the inferred close relationship between
Myrmekioderma and Didiscus and their inclusion in Halichondriidae, as proposed by Van
Soest et al. (1990). However, it must be acknowledged that structural similarities in the
erect palisade of ectosomal spicules is remarkable between Myrmekioderma and Higginsia
(see illustrations in Hooper and Levi 1993b), for
which genetic data might provide a more informed opinion.
Myrmekioderma is widespread in shallow
tropical and subtropical oceans, found predominantly on hard substrates, sand and rubble
substrates. One species, M granulata, has been
recorded previously from northwest Australian
and southern Indonesian waters (Van Soest et
al. 1990; Hooper and Levi 1993b), and is thought
to be one of the truly widespread coral reef
sponges throughout the Indo-west Pacific
(Hooper 1994).
Myrmekioderma granulata
(Figs 3-4)
(Esper, 1830)
Alcyonium granulatum Esper, 1830: 71, pl.
24.
Myrmekioderma granulata; - Ehlers, 1870:28;
- Burton 1938: 39, p1.7, fig. 42; - de Laubenfels
1954: 121, fig. 75; - Levi 1961: 14, fig. 17;
- Bergquist 1965: 177, fig.27a-b; - Van Soest et
al. 1990: 29, fig. 28; - Hooper et al. 1992: 265;
- Hooper and Levi 1993b: 1449, figs 35-36.
Acanthoxifer ceylonensis Dendy, 1905: 157,
pl. 9, fig. 5; - Dendy, 1922: 129.
Myrmekioderma tylota de Laubenfels, 1954:
119, fig. 74.
Neoprosypa atina de Laubenfels, 1954: 190,
fig. 127.
Acanthoxtfer fourmanoiri Levi, 1956: 5.
Material. NTM Z196: Dudley Point reef flat,
East Point, Darwin Harbour, NT, 12°25.0'S,
130°49.01'E, 0-0.5 m depth, 13 September 1981,
coll. J.N.A. Hooper and P.N. Alderslade, by
hand. NTM Z2053: Dudley Point reef slope,
Darwin Harbour, 12°25.0'S, 130°48.40'E, 6-10
m depth, 10 May 1984, coll. J.N.A. Hooper and
P.N. Alderslade, SCUBA. QM G303349: East
Point `bommies', 12°24.5'S, 130°48.8'E, 10 m
depth, 23 September 1993, coll. J.N.A. Hooper,
L.J. Hobbs and B. Alvarez, SCUBA. NTM Z430:
Lee Point, Darwin, 12°19.02'S, 130°53.01'E,
0-0.2 m depth, 13 December 1981, coll. J.N.A.
Hooper, by hand.
Comparative material (Refer to Hooper and
Levi 1993b: 1449 for additional material examined). QMG300819: W of Port Musgrave,
7
J.N.A. Hooper et al.
Gulf of Carpentaria, Qld 12°04.6'S,
139°21.5'E, 54 m depth, 25 November 1991,
coll. S. Cook, RV Southern Surveyor, dredge.
QMG303505: W of Duyfken Point, 12°28.4'S,
141°00.8'E, 53 m depth, 6 November 1993,
coll. S.D. Cook and J.A. Kennedy, RV Southern Surveyor, trawl. QMG303172: NE side
Flinders Reef, N. of Moreton I., SE Qld,
26°58.6'S, 153°29.2'E, 9 m depth, 31 May
1993, coll. J.N.A. Hooper, SCUBA.
QMG303135: Lagoon, Truk Atoll, Caroline Is,
7°24.9'N, 151°44.4'E, 31 m depth, 24 March
1992, coll. NCI (CRRF).
Distribution. Dudley Point, East Point, Lee
Point, Darwin Harbour, NT; Port Essington,
Cobourg Peninsula; Cumberland Strait, Wessel
Is, NT; Parry Shoals, Arafura Sea, NT; Enderbry
I., Direction I., Dampier Archipelago, WA;
Hibernia Reef, Sahul Shelf, Territory of Ashmore
and Cartier; Moreton Bay region, Qld; western
Gulf of Carpentaria, Qld; Madagascar, Aldabra,
Seychelles, Gulf of Manaar, SE Indonesia,
Ponape, Truk (Chuuk), Ebon Atolls, Palau,
Ifaluk, Chesterfield Is (refer to references in
Hooper and Levi 1993b).
Habitat. Shallow-water rock and coral platforms, coral pools, coral rubble, usually heavily
sedimented, or in spur-and-grooves of fringing
reefs; in deeper water found on exposed reef
heads in gravel and mud substrates; intertidal to
subtidal depths of 60 m.
Description. Massive, subspherical or elongate bulbous, up to 1 m long, 1 m wide, 0.5 m
thick; partially burrowing in soft sediment or
excavating coralline substrate. Oscules large,
discrete, conspicuous, usually situated in excavated grooves on apical surface, and each oscule
with raised lip.
Colour. Bright orange to dark orange-brown
exterior (Munsell 7.5YR 7/10-5YR 6/10), often
silt-covered in life; paler orange-yellow interior; orange-brown in ethanol.
Texture. Firm but compressible, harsh with
hispid surface; interior soft, producing mucus.
Surface. Opaque and membranous, covered
by rounded, raised polygonal plates separated
by grooves and canals, producing pineapplelike external surface.
Ectosomal skeleton. Detachable; distinct
crust of smaller ectosomal spicules forming continuous paratangential palisade, associated with
and protruding through heavy collagenous
ectosomal layer, and dispersed amongst larger
choanosomal oxeas and styles, the latter moreor-less erect and ascending to surface. Subecto-
somal region dense, collagenous, cavernous in
places, with tracts of larger choanosomal oxeas
supporting ectosomal crust.
Choanosomal skeleton. Skeletal structure
vaguely halichondroid reticulate, with compressed central portion, oblique subectosomal
skeletal tracts supporting paratangential
ectosomal layer. Choanosomal spicule tracts
composed of up to 50 choanosomal spicules
abreast, ascending to surface, becoming both
sparser and more confused in peripheral skeleton. Thick fibres visible only in some specimens (being fully cored by spicules in others),
with differentiated primary, multispicular, ascending fibres and transverse paucispicular fibres. Fibre meshes up to 300 pm diameter.
Collagen abundant, very granular and pigmented
in choanosome. Choanocyte chambers small,
oval, 40-70 pm diameter.
Spicules. Two size classes of megascleres
recognised: choanosomal and ectosomal; variable megasclere geometry, ranging from fusiform oxeas, strongyloxeas, strongyles or styles;
spicule terminations range from pointed to telescoped, styloid or sometimes with terminal deformities; and spicules may be completely smooth
or lightly and evenly acanthose (Berguist 1965).
In Beagle Gulf populations choanosomal
megascleres predominantly oxeas, less often
strongyloxeas or styloid, usually larger than
ectosomal spicules, often with microspination
(length 495-(615.7)-950 pm, width 8-(18.2)-22
pm). Smaller ectosomal megascleres invariably
oxeote, predominantly smooth but occasionally
evenly microspined (length 305-(502.4)-580 pm,
width 3-(7.5)-10 pm).
Microscleres raphides in bundles
(trichodragmata) (110-155 x 10-15 pm).
Remarks. Live specimens of M. granulata
are distinctive in their characteristic dark orange colouration, mango-like texture, massive
growth form and plate-like surface ornamentation. The species is now relatively well documented and known to be widely distributed
within the Indo-west Pacific region, extending
from Madagascar to NW Australia, Indonesia
and the central west-Pacific islands and atolls
(Esper 1830; Ehlers 1870; Dendy 1905, 1922;
Burton 1938; Levi 1956, 1961; de Laubenfels
1954; Bergquist 1965; Van Soest et al. 1990).
In the southwest Pacific the species is rare, so
far known only from isolated records (Flinders
Reef off Moreton Bay (27°S) and Chesterfield
Islands (21.5°S latitude; Hooper and Levi 1993b
and unpublished data). Surprisingly, the spe8
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Fig. 4. Myrmekioderma granulata (Esper). Specimen NTM Z3249. A, perpendicular section through skeleton, showing
compressed central portion of skeleton, oblique subectosomal spicule tracts and cavities, and paratangential ectosomal skeleton;
B, transverse section of ectosomal skeleton showing outer surface of sponge with palisade of smaller ectosomal oxeas dispersed
amongst larger choanosomal oxeas and styles; C, terminations of choanosomal strongyloxeas; D, termination of ectosomal oxen
and raphide microsclere.
cies has not yet been discovered from the Great
Barrier Reef, despite some intensive recent collections from many reefs, and it is possible that
M. granulata is merely an incursive into the
south Pacific region.
Populations of M. granulata in the central
west-Pacific region have been frequently recorded (refer to records in de Laubenfels (1954)
and Bergquist (1965)), whereas Australian and
southern Pacific populations were only recently
published (Hooper and Levi 1993b; Hooper
1994). Nevertheless, its distribution within Australia is now relatively well understood, extending as far south as 27°S latitude on the eastern
coast (restricted to inshore waters), and to 21.5°S
latitude on the western coast of the continent,
living in both coastal and offshore waters. Within
the Beagle Gulf Marine Park this species has
been seen on many shallow inshore fringing
coral reefs and coral platforms, especially in the
Darwin and Cobourg Peninsula regions (Hooper
and Levi 1993b), and it is also abundant on the
more pristine oceanic coral reefs off the coast of
northwest Australia (Hooper 1994) where it
grows to a much larger size and ranges to greater
depths than do the inshore populations.
Bergquist (1965) provided a detailed analysis
of morphometric variability in M. granulata,
which she correlated to some extent with regional populations of the species. Characters
found to vary between populations included surface sculpturing, live colouration, geometry,
spination and size of megascleres. These features are also shown to vary between the inshore
northwest Australian populations and those from
other localities. Specimens from the oceanic
coral reefs of the Sahul Shelf and those from
New Caledonia have meandering ridges excavating the apical surface (rather than discrete
polygonal plates) (compare Hooper and Levi
1993b: fig. 36A and the present work, Figure
3F-G); spicules are virtually entirely oxeote and
10
Beagle Gulf halichondriid sponges
well as raphide microscleres overlaying larger
choanosomal oxeas, typically showing a strict
localisation of spicule sizes, whereas in Topsentia
there there may or may not be size difference
between the ectosomal and choanosomal
spicules, and if present it is usually not consistent (i.e. spicule size may not be exclusive to any
particular region) (Diaz et al. 1991, 1993). Van
Soest et al. (1990) include Trachyopsis in synonymy with Topsentia, but this only applies to
the type species (others subsequently distributed
amongst Axinyssa and Spongosorites). The genus is found in all tropical and subtropical
oceans, extending from shallow waters to moderate depths. Within the Indo-Australian region
eight species of Topsentia has been previously
recorded: T. solicla (Ridley and Dendy) with
several junior synonyms (Hooper and Wiedenmayer 1994) from Tahiti, New Hebrides
(Vanuatu), Christmas Island (Indian Ocean),
Great Barrier Reef, and Ternate, Indonesia
(Ridley and Dendy 1887; Keller 1891;
Kirkpatrick 1900; Burton 1934), T. glabrata
(Keller) from Ternate, Indonesia (Keller 1891),
T. indica Hentschel from Am Island, Indonesia
(Hentschel 1912), T. dura (Lindgren) from
Gaspar Straits, Java Sea, Indonesia (Lindgren
1897, 1898), T. variabilis (Lindgren) and T.
armata (Lindgren) from Java, South China,
southern Philippines and Arafura Seas (Lindgren
1897, 1898; Hentschel 1912; Wilson 1925); T.
cavernosa (Topsent) from the Moluccas, Indonesia (Topsent 1897; Desqueyroux-Faundez
1981), and T. halichondrioides (Dendy) from
the Gulf of Manaar, southern Philippines, northern Great Barrier Reef, New Zealand (Dendy
1905; Wilson 1925; Burton 1934; Bergquist
1970).
smooth (whereas some inshore specimens have
acanthose ectosomal oxeas and styloid modifications to choanosomal oxeas); and skeletal
structure may be more cavernous (whereas most
inshore samples appear to be more compact)
(Hooper and Levi 1993b and present study). It is
possible that these differences are indicative of
a sibling species relationship (rather than
conspecificity), but this is not detectable solely
on the basis of morphological data.
Genus Topsentia Berg
[Anisoxya] Topsent, 1898: 225 (junior homonym of Anisoxya Mulsant, (Coleoptera)).
Topsentia Berg, 1899: 77.
Trachyopsis Dendy, 1905: 147.
Oxeostilon Ferrer-Hernandez, 1922: 10.
Alloscleria Topsent, 1927: 6.
Coelocalypta Topsent, 1928: 167.
Laminospongia Pulitzer-Finali, 1983: 546.
Type species. Anisoxya glabra Topsent, 1898,
by monotypy.
Diagnosis. Massive, amorphous or lobate
growth forms, with brittle texture. Choanosomal
skeleton completely lacking spongin, with mineral skeleton composed of oxeas in confused,
directionless arrangement closely packed around
aquiferous system, lacking any multispicular
tracts. Subectosomal skeleton sometimes with
subdermal cavities, but never large. Ectosomal
skeleton with compact paratangential or erect
layer of oxeas producing microhispid surface,
typically without definite organisation but often
closely compacted producing radial palisade.
Megascleres oxeas in two or more size classes,
including twisted, sinuous and contort modifications, usually without consistent size differences between ectosomal or choanosomal
spicules.
Remarks. The genera Topsentia, Epipolasis
and Petromica have completely confused halichondroid choanosomal skeletal structure, with
spicules criss-crossing in deeper parts of the
skeleton, they lack spongin within the
choanosomal skeleton, and the only recognisable organisation is found at the surface in the
form of spicule brushes. Species of Topsentia
are most distinctive in the disorganisation of
the choanosomal skeleton and lack of spongin
(hence the brittle consistency), and they are
typically drab in colour (Van Soest et al. 1990;
Diaz et al. 1991). Epipolasis has a tangential
felt-like ectosomal crust of smaller oxeas as
Topsentia halichondrioides (Dendy)
(Figs 5-6)
Trachyopsis halichondrioides Dendy, 1905:
147, p1.10, fig.I0. - Burton 1926: 75, figs 6-7; Burton 1934: 564; - Bergquist 1961: 185, fig.9;
- Bergquist 1970: 34.
Type material. HOLOTYPE - BMNH
1907.2.1.44: Galle, Periya Paar, Gulf of Manaar,
Ceylon (Sri Lanka).
Other Material. NTM Z3262: SW side of
cliff face, Table Head, Port Essington, Cobourg
Peninsula, NT, 11°13.5'S, 132°10.5'E, 5 m
depth, 11 September 1986, coll. J.N.A. Hooper
and C. Johnson, SCUBA. QM G303442: W
11
J.N.A. Hooper et al.
Fig. 6. Topsentia halichondrioides (Dendy). Specimen QM G303442. A, perpendicular section through skeleton showing
disorganised ascending halichondroid choanosomal skeleton supporting the plumose subectosomal skeleton of larger oxeas, with
a thin tangential ectosomal layer of smaller oxeas just below surface; B, transverse section of ectosomal skeleton showing outer
surface of sponge with erect palisade of larger oxeas, with smaller oxeas lying below and tangential to this layer; C, light
micrograph of section through peripheral skeleton showing distinct demarkation between the halichondroid reticulate choanosomal
skeleton and the nearly plumose subectosomal skeleton; D, larger oxea (left) with faintly telescoped point, and sharply smaller
oxea (right).
in dimensions; T. dura is greenish alive, olivebrown when preserved, stony consistency, small
sunken oscules, two sizes of oxeas up to 600 x
20 pm and 260 x 6 pm, respectively, and an
erect, continuous ectosomal palisade; T. armata
is tuberculate, fibrous consistency, minute
oscules, very large oxeas up to 1700 x 48 pm,
and a sparse ectosomal skeleton; T. variahilis is
massive flattened, grey-brown preserved, fibrous
consistency, with prominent surface microconules, minute oscules, two sizes of oxeas, up
to 770-1160 x 16-30 pm and 520-650 x 10-24
pm respectively, with an oblique, plumose
ectosomal skeleton composed of discrete (not
continuous) brushes, and multispicular tracts in
the choanosomal skeleton; T indica is gray preserved, fibrous consistency, small oscules with
slightly raised margins, oxeas in two sizes, up
to 800 x 23-55 pm and 380 x 18 pm, respectively, ectosomal skeleton radial, well devel-
oped; T. cavernosa is massive, has a hard texture, moderately well developed erect ectosomal
skeleton composed of only marginally smaller
oxeas than in the choanosomal skeleton, and
oxeas are 600-1000 x 17-20 pm.
Topsentia dura (Lindgren)
(Figs 7-8)
Halichondria dura Lindgren, 1897: 480. Lindgren 1898: 286-287, pl. 17, fig. 2, pl. 19,
fig. 3.
Type material. Fragment of holotype BMNH 1929.11.26.41: Selat Kelasa (Gaspar
Straits), Java Sea, Indonesia, coll. C. Aurivillius,
1891.
Other Material. NTM Z3195: East Point
`bommies', Darwin, NT, 12°24.5'S, 130°48.8'E,
9 m depth, 16 September 1987, coll. N. Smit,
14
Beagle Gulf halichondriid sponges
■
I
MOW
Fig. 7. Topsentia dura (Lindgren). A, larger (predominantly) choanosomal oxea; B, smaller (predominantly) ectosomal oxea;
C, peripheral skeletal structure; D, preserved specimen NTM Z3I95; E, specimen NTM ZI442.
Colour. Yellow-green to dark green in life
(Munsell 5Y 8/4 - 2.5GY 5/4), white in ethanol.
Texture. Very firm, incompressible, brittle,
easily broken.
Surface. Optically smooth but uneven, lumpy;
encrusted in places with detritus; hispid.
Ectosomal skeleton. Continuous erect palisade of protruding single spicules or paucispicular brushes, composed mainly of larger
oxeas, standing erect on ascending peripheral
choanosomal tracts and protruding through surface only slightly beyond collagenous membrane;
sparse tangential layer of smaller oxeas at base
of protruding larger spicules, presumably lying
on ectosomal membrane.
Choanosomal skeleton. Dense halichondroid
disorganised mass of single spicules, mostly
larger oxeas, organised only around lacunae
SCUBA. NTM Z3178: Dudley Point Reef, East
Point, Darwin, NT, 12°25.0'S, 130°49.1'E, 0.5
m depth, 10 September 1987, coll. N. Smit, by
hand. NTM Z1442: Blue Hole, Gunn Point,
NT, 12°09.0'S, 131°00.5'E, 25 m depth, 19
August 1983, coll. J.N.A. Hooper and P.N.
Alderslade, SCUBA.
Distribution. Java Sea, Indonesia; East Point
Aquatic Reserve and Gunn Point, Darwin Harbour, NT.
Habitat. Laterite rock and fringing coral reef,
sandy substrate; intertidal-subtidal depths.
Description. Massive to lobate, irregularly
anastamosing lobes, up to 110 mm long, 40 mm
diameter. Oscules small, less than 2 mm diameter, discrete, scattered over surface of lobes,
sunken in slight depressions on surface, without
visible oscular rim.
15
J.N.A. Hooperet al.
Fig. 8. Topsentia dura (Lindgren). Specimen NTM Z3195. A, perpendicular section through skeleton of branch showing
halichondroid reticulate choanosome and plumose subectosomal skeletons; B, transverse section of ectosomal skeleton showing
outer surface of sponge with a plumose ectosomal skeleton composed of protruding larger oxeas and a sparse tangential skeleton
of smaller oxeas at their base; C, light micrograph of section through peripheral skeleton showing faint demarkation between the
halichondroid reticulate choanosomal skeleton and the nearly plumose subectosomal skeleton; D, sharply pointed fusiform larger
oxea (left) and smaller oxea (right).
and in peripheral skeleton, becoming plumose
in periphery. Choanosomal and subectosomal
regions equally cavernous, with occasionally
large lacunae (>500 pm diameter) but mostly
smaller oval meshes (<230 pm diameter). Only
faint demarkation visible between choanosomal
and subectosomal regions. Collagen in
choanosome present but lightly pigmented.
Spicules. Oxeas in two sizes; larger predominantly choanosomal oxeas moderately or slightly
curved at centre, fusiform, sharply pointed
(length 344-(447.0)-585 pm, width 10-(12.3)16 pm); smaller predominantly ectosomal oxeas
straight or slightly curved, fusiform, sharply
pointed (length 190-(211.5)-262 pm, width 1(3.8)-6 pm)
Remarks. Topsentia dura is also characteristically stony in texture, has a massive lobate
growth form, and has a well developed continuous palisade of erect ectosomal oxeas on the
surface, similar to T. halichondrioides. It differs from T. halichondrioides in its live
colouration (greenish), oscule morphology
(small, sunken in slight depressions, without an
oscular rim), spicule dimensions (mean of 447
x 12 pm and 211 x 4 pm for larger and smaller
oxeas, respectively) (although only one size class
of oxeas was originally recorded by Lindgren
1897, 1898), sharply pointed spicule terminations, and a less extensive ectosomal palisade.
By comparison, T. halichondrioides has a distinctive purple-mauve live colouration, its
oscules are large, discrete, lying on the apex of
bulbous projections, with a membraneous lip,
and oxeas differ in both dimensions (562 x 23
pm and 185 x 8 pm, respectively) and geometry
(the larger ones commonly with telescoped
points). Refer to Remarks for T. halichondrioides for further comparisons between species.
As in T. halichondrioides, his species has some
16
Beagle Gulf halichondriid sponges
degree of localisation between spicule sizes forming various portions of the skeleton, with smaller
oxeas predominantly found in the ectosome and
larger oxeas predominantly in the choanosomal
skeleton, reminiscent of Epipolasis. But this
localisation is not completely strict, with both
sizes also found throughout the skeleton, and
the value of this character in Topsentia is questionable.
Genus Ciocalypta
these differences will eventually be considered
artificial, especially considering that echinating
ectosomal styles in Amorphinopsis can only be
detected reliably using SEM techniques (i.e. a
character of limited value to field biology), but
any proposed merger between these genera requires more rigorous evaluation of a greater
suite of species than presented here.
Species of Ciocalypta are widely distributed
in temperate, subtropical and tropical oceans,
mostly confined to shallow waters on muddy or
sandy bottoms, frequently burrowing, at least
partially, into the soft sediments. Within the
Indo-Australian region eight species are known:
C. massalis (Carter) from Bass Strait, Vic., and
Fremantle, WA (Carter 1883); C. polymastia
(Lendenfeld) from an unspecified locality, east
coast of Australia (Lendenfeld 1888; Hallmann
1914); C. occulata (Kieschnick) and C. simplex
Thiele from Ternate, Moluccas, Indonesia (Kieschnick 1896; Thiele 1900), and C. subaceratus
(Ridley and Dendy) from southern Mindanao,
Philippines, and Ternate, Indonesia (Ridley and
Dendy 1886; Thiele 1900); C. fenestrata (Ridley)
from the Arafura Sea, Great Barrier Reef, and
South China Sea (Ridley 1884; Lindgren 1897,
1898; Burton 1934); C. gracilis (Hentschel)
and C. heterostyla Hentschel from the Arafura
Sea (Hentschel 1912), both synonymised in Halichondria ryleri (Bowerbank) by Burton (1959;
see Hooper and Wiedenmayer 1994), but this
synonymy is rejected.
Bowerbank
Ciocalypta Bowerbank, 1863: 1105.
Leucophloeus Carter, 1883: 323.
Apatospongia Marshall, 1892: 16.
Type species. Ciocalypta penicillus Bowerbank, 1864, by monotypy.
Diagnosis. Growth form exclusively fistulose,
with base burrowing into substrate. Surface usually semi-transparent, detachable, parchmentlike. Choanosomal skeleton composed of styles
in confused, directionless arrangement, spicule
tracts unispicular, and spongin virtually absent
in massive portion of sponge, whereas fistules
supported by multispicular columns of styles
confined to central portion of fistules. Ectosomal
skeleton tangential layer of smaller styles, usually in bundles, supported by widely spaced
multispicular tracts of styles, perpendicular to
central choanosomal tracts, producing large
subclermal cavities (lacunae) in peripheral region. Megascleres predominantly styles, sometimes accompanied by fewer oxeas.
Remarks. The revised concept of Ciocalypta
is now restricted to fistulose species (Van Soest
et al. 1990) which have exclusively stylote
megascleres including a second, smaller category of ectosomal styles. Although fistulose
growth forms seen in Ciocalypta are also known
for some species of Topsentia, Halichondria
and Petromica (i.e. this is likely to be an independently acquired ecological adaptation to living in sand substrates whereby fistules prevent
smothering), these other genera differ from
Ciocalypta substantially in their spicule geometry (oxeas instead of styles) and ectosomal characteristics (with erect bundles of spicules).
Ciocalypta is most similar to Amorphinopsis,
both having smaller ectosomal styles and larger
stylote or oxeote choanosomal megascleres,
whereas in Ciocalypta these ectosomal spicules
are simply styles overlaying the choanosomal
skeleton, in Amorphinopsis the ectosomal skeleton is composed of bundles of oxeas which are
echinated by smaller styles. It is possible that
Ciocalypta fenestrata (Ridley)
(Figs 9-10)
Leucophloeus fenestratus Ridley, 1884: 464,
pl. 42, fig. s. - Burton, 1934: 564.
Leucophloeus fenestratus var. [unnamed]
Ridley, 1884: 464, pl. 43, fig. g.
Not Leucophloeus fenestratus - Dendy, 1922:
124.
Hymeniacidon fenestratus - Lindgren, 1897:
483; - Lindgren, 1898: 312-313.
Axinyssa fenestratus - Van Soest et al., 1990:
27; - Hooper and Bergquist, 1992: 102.
Type material. HOLOTYPE - BMNH
1882.2.23.255: Port Darwin, NT, 16-24 m depth,
October 1881, HMS Alert, bottom mud and
sand. Holotype of variety - BMNH 1882.2.23.195:
Arafura Sea, NT, 64-72 m depth, 18 October
1881, coll. HMS Alert, bottom mud, sand, shells.
Other Material. NTM Z2018: West side
Weed Reef, Darwin, NT, 12°29.2'S, 130°47.1'E,
17
J.N.A. Hooper et al.
C
C
Fig. 9. Ciocalypta fenestrata (Ridley). A, larger choanosomal styles; B, smaller ectosomal styles;
C, peripheral skeletal structure (surface at right); D, preserved specimen NTM Z2018.
18
Beagle Gulf halichondriid sponges
8 m depth, 11 May 1984, coll. J.N.A. Hooper,
P.J. Homer and P.N. Alderslade, SCUBA.
Distribution. Darwin Harbour, Arafura Sea,
Shoal Bay, NT; Low Isles, Mangrove Park, Great
Barrier Reef, Qid.; Providence Reef, Amirante,
Indian Ocean; Java and Lesser Sunda Islands,
Indonesia (Ridley 1884; Lindgren 1897, 1898;
Dendy 1922).
Habitat. Burrowing in sand, mud, coral rubble, shell grit substrates; 4-72 m depth range.
Description. Partially burrowing; subspherical, massive, buttressed base (50mm diameter,
100 mm high), with small tapering fistules on
apex (up to 30 mm long, 10 mm basal diameter) with basal portion buried beneath sediment and fistules protruding through sand.
Oscules large, up to 8mm diameter, on apex of
fistules.
Colour. Yellow-brown in life (Mansell 7.5YR
7/8), pale brown in ethanol.
Texture. Compressible, harsh, easily torn;
sand and coral rubble embedded on under surface and outer surface of base.
Surface. Irregular, transparent and hispid.
Ectosomal skeleton. Multispicular tracts of
larger choanosomal styles, up to three spicules
abreast, lying tangential to surface, and irregular bundles of smaller ectosomal styles arranged
mostly paratangentially to surface as plumose
brushes. Collagen present but light in ectosomal
region.
Choanosomal skeleton. In deeper regions of
sponge, skeleton disorganised halichondroid
criss-cross of both unispicular and multispicular
tracts, containing 5-10 spicules abreast, composed of larger choanosomal styles; in subectosomal region tracts becoming more wide-meshed,
paratangential, producing cavernous subectosomal region; in fistules choanosomal tracts more
compressed in central region, cavernous towards
Fig. 10. Ciocalypta fenestrata (Ridley). Specimen NTM Z2018. A, perpendicular section through skeleton of fistule, showing
multispicular halichondroid reticulate skeleton, large subectosomal lacunae, and paratangential subectosomal skeleton;
B, transverse section of ectosomal skeleton showing outer surface of sponge with a tangential multispicular bundles of larger
styles and paratangential bundles of smaller styles; C, terminations on choanosomal styles; D, terminations on ectosomal styles.
19
J.N.A. Hooperet al.
also substantially larger (670-920 x 20-30 pm,
200-400 x 7-10 pm, respectively), and growth
form consists of a thinner crust, but apparently
not burrowing, with fairly large fistules up to 30
mm long and 15 mm diameter. Similarly, C.
massalis is massive and buttressed, with a prominently rugose surface but apparently lacking
fistules (probably destroyed in the unregistered
BMNH dry type material, re-examined by the
senior author in the BMNH `Bowerbank Collection'). Ciocalypta massalis also has a relatively distinctive skeletal structure (consisting
of a clearly differentiated compressed central
region and radiating peripheral tracts of styles
which become plumose closer to the surface,
supporting a paratangential ectosomal crust composed of smaller styles), and spicule dimensions
differ from those of C. fenestrata with only
marginal size differences between the choanosomal and ectosomal styles (475 x 15 pm, 225 x
8 pm, respectively). It is possible that these four
species are conspecific, representing substantial
morphometric variability across a large latitudinal gradient, but this explanation is unacceptable under current interpretations of sponge morphometrics, and it is suggested that these species
are at most close siblings.
periphery; cavities at least as wide as spicule
bundles, up to 400 pm in basal region, 1.5 mm
in fistular region. Choanosomal spicule tracts
predominantly composed of larger styles,
whereas smaller (ectosomal) styles scattered
throughout mesohyl. Collagen moderately abundant in central region of choanosome, sparse in
peripheral skeleton; no spongin fibres observed.
Spicules. Larger choanosomal styles variable
in size, straight or slightly curved at centre,
fusiform with sharp points or very slightly telescoped points, tapering to hastate rounded base
smaller in diameter than centre of spicule (length
355-(521.6)-775 pm, width 8-(19.2)-32 pm).
Smaller ectosomal styles fusiform, sharply
pointed with evenly rounded bases (length 153(246.5)-330 pm, width 6-(9.1)-12 pm).
Remarks. Comparison between the known
specimens of this species, including type material, confirm their conspecificity, and the species is referred here to Ciocalypta based on the
possession of a specialised tangential-paratangential ectosomal skeleton. In re-evaluating the
genera Pseudaxinyssa and Axinyssa, both Van
Soest et al. (1990) and Hooper and Bergquist
(1992) considered that this species might be
included in Axinyssa, (based on its published
skeletal structure and surface characters; the
ectosomal skeleton of type material was in relatively poor condition). However, examination
of fresh material described above discovered
that smaller styles are mostly confined to a
specialised ectosomal layer, not scattered within the choanosomal skeleton as presumed previously, indicating that Ciocalypta is a more appropriate generic assignment.
Ciocalypta fenestrata belongs to a species
group of Ciocalypta characterised in having
`subacerate' larger choanosomal styles, whereby the thickest part of the spicule occurs in the
apical third and the basal end is substantially
narrower than the centre of the spicule. Included in this group are C. oculata (Kieschnick,
1896) from Temate, Indonesia, C. oculata var.
maxima Hentschel, 1912, from Aru Is, Indonesia, C. subacerata (Ridley and Dendy, 1886)
from the Mindinao, Philippines, and Ternate,
Indonesia, and C. massalis (Carter, 1883) from
southwest Australia. The growth form of C.
subacerata consists of massive anastomosing
branches (`trabeculae') forming a cavernous construction, and megascleres are substantially larger than in C. fenestrata (1200 x 31 pm, 200 x 6
pm for both categories of styles). Spicule dimensions in C. oculata and its subspecies are
Ciocalypta oscitans sp. nov.
(Figs 11-12)
Type material. HOLOTYPE - QM G303560:
4 km E of Fish Reef, Bynoe Harbour, NT,
12°24.8'S, 130°28.9'E, 17 m depth, 6 October
1993, coll. CCNT stn. 92 (ref. 1128), dredge.
Distribution. Bynoe Harbour, NT.
Habitat. Coarse sand and gravel sediment;
subtidal.
Description. Massive, subspherical, 110 mm
diameter, excavated, partially hollow interior,
buttressed exterior, partially burrowing into soft
sediments. Oscules very large (10-15 mm diameter) on upper surface, with vestigial fistules up
to 10 mm long.
Colour. Pale mauve alive on deck (Munsell
5RP 8/2), white in ethanol.
Texture. Firm, compressible, harsh and easily torn.
Surface. Optically smooth, unornamented,
with subectosomal cavities visible through translucent ectosomal membrane.
Ectosomal skeleton. Detachable, dense tangential layer, composed of both smaller and
larger styles arranged mostly tangential to surface, in thick multispicular tracts, supported by
20
Beagle Gulf halichondri id sponges
O
O
Fig. 11. Ciocalypta oscitans sp. nov. A, larger choanosomal style; B, smaller ectosomal
styles; C, peripheral skeletal structure (surface at right); D, preserved holotype QM
G303560; E, live holotype (on deck).
21
J.N.A. Hooper et al.
Fig. 12. Ciocalypta oscitans sp. nov. Holotype QM G303560. A, perpendicular section through skeleton showing disorganised
halichondroid reticulation and cavernous meshes in central choanosomal region, more organised towards periphery; B, transverse
section of ectosomal skeleton showing outer surface of sponge with a thin tangential layer of intermixed smaller and larger styles;
C, hastate point (left) and rounded base of smaller ectosomal style (right); D, slightly telescoped point (left) and evenly rounded
base of larger choanosomal style (right).
paratangential spicule tracts from subectosomal
skeleton.
Choanosomal skeleton. Disorganised halichondroid reticulation of distinct multispicular
tracts in centre of choanosome, composed mainly
of larger choanosomal styles, with only slightly
cavernous meshes throughout choanosomal and
subectosomal regions, up to 500 pm diameter in
choanosomal region but much smaller in peripheral skeleton; subectosomal tracts of larger
styles ascending perpendicular to surface, producing nearly regular, open reticulate (crisscross) arrangement of multispicular tracts, with
large subdermal cavities visible close to surface;
smaller styles also scattered throughout skeleton, mostly paratangential to major spicule
tracts, these largely confined to ectosomal skeleton. Fibres absent; collagen poorly developed
in mesohyl.
Spicules. Styles in two size classes: larger
choanosomal styles straight or slightly curved
near basal end, fusiform, sharply pointed or
slightly telescoped points, with evenly rounded
or slightly tapering hastate bases (length 515(776.1)-944 pm, width 11-(16.7)-20 pm);
smaller ectosomal styles hastate pointed, with
evenly rounded bases (122-(175.4)-216 pm,
width 5-(6.3)-8 pm).
Etymology. Named for the large oscules, from
oscitans (L.), yawning.
Remarks. This species has a choanosomal
skeleton notably more compact (less cavernous)
than other known species of Ciocalypta. It also
has a more dense tangential ectosomal skeleton,
distinctly detachable from the subectosomal region, whereas in most other species the tangential layer of smaller ectosomal styles is relatively
sparse and forms mainly uni- or paucispicular
tracts. These characters are perhaps better developed than in most other species examined, but
this is a subjective interpretation which is difficult to substantiate amongst the relatively homo22
Beagle Gulf halichondriid sponges
geneous skeletal structures seen in this group of
halichondriids (i.e. they are not particularly discriminatory between species). Ciocalypta
oscitans is a typically 'amorphous' burrowing
sponge, similar in growth form to many other
halichondriids, such as C. confossa sp. nov., but
it has distinctive, very large oscules raised on
small (vestigial) fistules. Certainly in comparison
with C. confossa the present species differs substantially in all skeletal structures, whereas
spicule dimensions are comparable between the
species (although spicule geometries are not).
Spicule dimensions in both these burrowing species, C. oscitans and C. confossa, are also equivalent to those seen in C. heterostyla Hentschel,
from Aru Island, Indonesia, although growth
form is substantially different (the latter compris-
ing erect fistulose ridges, distinct ectosomal
`skin' and oscules on the apex of ridges), and
skeletal structures are markedly more cavernous
in the Indonesian species.
Ciocalypta confossa sp. nov.
(Figs 13-14)
Type material. HOLOTYPE - NTM Z3106
(fragment NCI Q66C-0559-W): Parry Shoals,
Arafura Sea, NT, 11°12.53'S, 129°42.08'E, 20
m depth, 15 August 1987, coll. A.M. Mussig
and NCI AIMS, SCUBA.
Other Material. QM G303287: S of South
Shell I., East Arm, Darwin, NT, 12°29.8'S,
130°53.5'E, intertidal, 19 September 1993, coll.
J.N.A. Hooper and L.J. Hobbs, by hand. QM
■ IN e • ■ III 1
Fig. 13. Ciocalypta confossa sp. nov. A, larger choanosomal style; B, smaller ectosomal styles; C, peripheral skeletal structure;
D, preserved specimen QM G303287; E, live intertidal specimen QM G303287, burrowing under substrate.
23
J.N.A. Hooper et al.
G303541: W of South West Vernon I., NT,
12°06.9'S, 131°04.8'E, 13 m depth, 10 October
1993, coll. CCNT stn. 140 (ref. 1664), dredge.
QM G303558: NW of Cape Hotham, NT,
12°01.3'S, 131°13.9'E, 34 m depth, 9 October
1993, coll. CCNT stn. 159 (ref. 1498), dredge.
Distribution. East Arm, Darwin Harbour;
Vernon Islands, Cape Hotham, Parry Shoals,
Arafura Sea.
Habitat. Coral reef, rubble, coarse sand and
shell grit; intertidal-subtidal depths.
Description. Completely burrowed into soft
muddy substrate, with small fistules and small
apical oscules barely visible above sediment;
massive, up to 120 mm diameter, subspherical
or tubulo-digitate, cavernous insubstantial construction (excavated throughout); fistules rudimentary, up to 18 mm long, 12 mm diameter,
with moderately small oscules up to 5 mm diameter conspicuous on upper surface.
Colour. Yellow or yellow-brown in life
(Munseli 2.5Y 8/8 - 7/10), pale brown in ethanol.
Texture. Firm, compressible, stringy, friable,
easily torn.
Surface. Opaque, membranous, roughened,
fibrous, usually with some detritus, often mudcovered in situ.
Ectosomal skeleton. Thin, detachable, tangential layer of small styles, scattered in uni- or
paucispicular directionless tracts over surface,
usually associated with granular collagen.
Choanosomal skeleton. Disorganised halichondroid reticulation of both multispicular
directionless tracts and single spicules;
multispicular tracts consisting of spicule bundles composed mainly of larger styles, 5-20
spicules abreast, without apparent organisation
but forming vaguely halichondroid-reticulate
skeleton; single spicules scattered throughout
mesohyl in between major tracts; meshes large,
oval, cavernous, up to 700 pm diameter, found
throughout skeleton (not necessarily confined
to subectosomal region). Fibres absent; poorly
developed collagen in choanosomal mesohyl.
Fig. 14. Ciocalypta confossa sp. nov. Specimen QM G303287. A, perpendicular section through skeleton showing cavernous
halichondroid reticulate skeleton supporting a thin ectosomal peel; B, transverse section of ectosomal skeleton showing outer
surface of sponge with a thin tangential layer of smaller styles; C, smaller ectosomal style with fusiform point (left) and slightly
subtylote base (right); D, larger choanosomal style with fusiform point (left) and evenly rounded base (right).
24
Beagle Gulf halichondriid sponges
Spicules. Styles in two size classes; larger
choanosomal styles straight or slightly curved
near pointed end, fusiform, sharply pointed with
evenly rounded bases or sometimes anisotylote
(length 4254661.6)-940 pm, width 8-(13.4)-20
pm); smaller ectosomal styles slightly curved at
centre or near basal end, fusiform, sharply
pointed, with evenly rounded or slightly subtylote
bases, occasionally anisotylote or centrotylote
(length 1304183.3)-275 pm, width 3-(5.1)-7
pm).
Etymology. Named for the burrowing, excavated habit, from confossus (L.), perforated, full
of holes.
Remarks. Although both are burrowing species, with vestigial fistules and similar ranges of
spicule sizes, C. confossa differs from C. oscitans
substantially in both its field characteristics and
skeletal stuctures. It is nearly completely submerged below the substrate, with rudimentary
fistules barely protruding above the surface, having much smaller oscules placed on top of these
fistules, its ectosomal skeleton is sparse, with
poorly developed paratangential single ectosomal
styles scattered over the surface, overlaying a
relatively cavernous subectosomal skeleton,
spicule geometry of the smaller ectosomal styles
in particular differs, and choanosomal skeletal
structure is much less compact than in C.
oscitans.
Genus Amorphinopsis Carter
Amorphinopsis Carter, 1887: 77.
Migas Sollas, 1908: 395 (junior homonym of
Migas Koch (Arachnida)).
Prostylissa Topsent, 1925: 208.
Tumata de Laubenfels, 1936: 77.
Milne de Laubenfels, 1954: 116.
Type species. Amorphinopsis excavans
Carter, 1887, by monotypy.
Diagnosis. Growth forms encrusting, excavating or ramose. Choanosomal skeleton with
widely spaced halichondroid criss-cross of larger
spicules, producing relatively large meshes
bounded by mostly multispicular tracts.
Ectosomal skeleton composed of thick bundles
of larger oxeas lying tangential to surface
echinated by smaller styles or oxeotes.
Megascleres include smaller styles and/or oxeas,
and larger choanosomal spicules predominantly
oxeas or strongyloxeas, occasionally stylote.
Remarks. The possession of smaller styles
forming the ectosomal skeleton, overlaying
larger choanosomal megascleres, is shared with
25
the sister genus Ciocalypta. In Amorphinopsis,
however, these smaller styles echinate the tracts
of larger choanosomal oxeas that lie tangential
to the ectosome (which is unique amongst the
Halichondriidae). The few known species appear to be restricted to the Indo-west Pacific
region, confined to shallow waters (Van Soest
et al. 1990).
Four species of Amorphinopsis have been previously described in the Indo-west Pacific: A.
excavans (Carter) from Torres Strait, Java Sea
and Arafura Sea (Lindgren 1897; Burton 1934),
A. megarrhaphea (Lendenfeld) from the south
coast of NSW (Lendenfeld 1888), A. foetida
(Dendy) and A. sacctformis (Thiele) from the
Moluccas, Indonesia, and Papua New Guinea
(Topsent 1897; Thiele 1900; Kelly-Borges and
Bergquist 1988).
Amorphinopsis excavans Carter
(Figs 15-16)
Amorphinopsis excavans Carter, 1887: 77,
p1.5, figs 12-15. - Annandale 1915: 467, fig.
4A; - Burton 1959: 257; - Van Soest et al. 1990:
45, fig. 61.
Amorphinopsis excavans digitifera Annandale, 1915: 469, figs 4B, 5.
Halichondria armata Lindgren, 1897: 480. Lindgren 1898: 285-286, pl. 17, fig. 1, pl. 19,
fig. 2.
Halichondria panicea hemisphaerica Dendy,
1905: 146-147.
Amorphinopsis kempi Kumar, 1925: 226,
fig. 7.
Prostylissa siamensis Topsent, 1925: 208, figs
1-2. - Van Soest et al. 1990: 45, fig. 64.
Type material. HOLOTYPE - IM 6597/7
ZEV: Mergui Archipelago, Burma, Andaman
Sea. holotype fragment of H. armata - BMNH
1929.11.26.31 (slide): Batan Island, Philippines.
Syntype of H. panicea hemisphaerica - BMNH
1907.2.1.43: N. of Galle, Gulf of Manaar, Sri
Lanka. Holotype of A. excavans digitifera - IM
501/7 ZEV: Fisher Bay, Tavoy Island, off
Tenasserim, Burma, Andaman Sea. Holotype of
A. kempi - IM P199/1: Waltair, Bay of Bengal,
India.
Other Material. QM G303658: Vernon Islands, South of Marsh Shoals, NT, 12°07.0'S,
130°56.1'E, 16 m depth, 11 October 1993, coll.
CCNT stn. 138, dredge (ref. CCNT 1997).
Distribution. Widespread in coral reefs of
the Indo-west Pacific - Gulf of Manaar, Sri
Lanka; Bay of Bengal, India; Andaman Sea,
J.N.A. Hooper et al.
B
00
Fig. 15. Amorphinopsis excavans Carter. A, larger choanosomal oxea; B, smaller ectosomal style;
C, peripheral skeletal structure (surface at right); D, preserved specimen QM G303658;
E, live specimen (on deck) encrusting coral.
26
Beagle Gulf halichondri id sponges
Burma; South China Sea, Vietnam; Sulu Sea,
Philippines; Gaspar Strait, Java Sea, Banda Sea
and Arafura Sea, Indonesia; Vernon Islands,
NT; Torres Strait, Old.
Habitat. Gravel substrate; intertidal to 54 m
depth.
Description. Thickly encrusting over
Pocillopora coral fronds and large fragments of
siliceous detritus, up to 15 mm thick, with convoluted surface profile. Oscules small, discrete,
1-2 mm diameter, scattered over surface, with
slighlty raised membraneous lip.
Colour. Yellow-brown in life (Mansell 7.5YR
7/10), beige-white in ethanol.
Texture. Firm but compressible, easily torn.
Surface. Opaque, optically smooth, porous
convoluted surface.
Ectosomal skeleton. Ectosome a thin peel,
detachable, composed of tangential layer of
larger oxeas, mostly in bundles lying on surface, with small styles oriented paratangential
to ("echinating") and protruding through surface. Ectosome with sparse collagen and detritus.
Choanosomal skeleton. Tightly packed confused halichondroid reticulate skeleton of larger
oxeote spicules, mostly formed by multispicular
bundles bound together with very little collagenous material, producing small elongate cavities (lacunae) and oval meshes throughout skeleton. Towards periphery, spicule tracts becoming more paratangential, subplumose, producing larger lacunae free of spicules. Sparse collagen dispersed throughout mesohyl.
Spicules. Larger choanosomal oxeas fusiform,
slightly curved at centre, tapering to sharp points
(length 175-(506.6)-642 pm, width 8-(12.8)-18
pm). Smaller ectosomal styles slightly curved at
Fig. 16. Amorphinopsis excavans Carter. Specimen QM G303658. A, perpendicular section through skeleton, showing
halichondroid reticulate skeleton of mostly multispicular tracts and spaces (lacunae) within choansomal skeleton, paratangential
subplumose subectosomal tracts and thin ectosomal peel; B, transverse section of ectosomal skeleton showing outer surface of
sponge with a tangential layer of larger oxeas, in bundles, echinated by small styles; C, closer view of tangential ectosomal
skeleton; D, sharply pointed terminations on choanosomal oxeas (left) and ectosomal styles (right).
27
J.N.A. Hooperet al.
Amorphinopsis foetida - Topsent, 1897: 445,
p1. 18, fig. 8; - Desqueyroux-Faundez 1981:
757.
Ciocalypta foetida - Lindgren, 1897: 483; Lindgren 1898: 313-314; - Thiele 1900: 75, pl.
3 fig. 26; - Hentschel 1912: 427-428.
Leucophloeus foetida - Dendy, 1905: 197,
234.
Material. QMG303677: Shoal Bay, middle
of bay, NT, 12°13.1'S, 130°56.0'E, 17 m depth,
12 October 1993, coll. CCNT stn. 122, dredge
(ref. CCNT 2104).
Distribution. Tuticorin, Gulf of Manaar, Sri
Lanka; S. coast of Vietnam, South China Sea;
Ambon and Ternate, Moluccas, Indonesia;
Arafura Sea, Indonesia; Shoal Bay, Darwin region, NT.
Habitat. Sandy mud substrate with sparse
seagrass cover.
Description. Enlarged base burrowing into
substrate with two subconical, low volcanoshaped fistules protruding above sediment.
Larger oscules, up to 8 mm diameter, grouped
on a terminal sieve-plate at the apex of each
fistule.
Colour. Yellow-beige on deck (Munsell
7.5YR 8/2), white in ethanol.
Texture. Soft, compressible and easily torn.
Surface. Transparent, membranous, irregular, wrinkled and hispid.
Ectosomal skeleton. Dense layer of larger
choanosomal styles lying tangential to the surface, mostly in irregular multispicular bundles
of 2-10 spicules, with smaller ectosomal styles
producing tufts (`echinating') around spicule
bundles. Spicule bundles bound together with
small amounts of collagen.
Choanosomal skeleton. Architecture cavernous, reticulate, with distinct structural differences between central and peripheral skeletons.
Deeper choanosomal skeleton with large
multispicular tracts composed of larger
choanosomal styles, 20-50 spicules wide, mainly
confined to central region of sponge (particularly in fistules). Peripheral skeleton with widely
spaced multispicular radial tracts of styles arising perpendicular to central skeleton, becoming
plumose in near surface, supporting tangential
ectosomal skeleton; spicule tracts forming widemeshed reticulate skeleton with very large cavernous meshes up to 2 mm diameter, particularly in fistular region; some collagenous material associated with spicule tracts but true fibres
absent.
centre, with rounded bases and tapering fusiform points (length 164-(190.3)-244 pm, width
2-(4.1)-8 pm).
Remarks. Amorphinopsis excavans is a
widely distributed Indo-west Pacific species commonly associated with coral reefs and coral rubble. Burton (1959) included it under the name
A. megalorhaphis (Carter), but this synonymy
cannot be corroborated (the type material being
lost; Hooper and Wiedenmayer 1994) and the
name excavans' has seniority. Similarly, many
of the synonyms given by Burton (1959) are
questionable, and we include only those species
for which conspecificity has been established
through comparison of relevant type material
(listed above). In particular, two species mentioned in synonymy with A. excavans, Halichondria dura Lindgren and H. variabilis
Lindgren (both from Indonesia and both belonging to Topsentia), are specifically excluded
from comparison of fragments of type material
(BMNFI 1929.11.26.41 and BMNH 1929.11.26.1,
respectively). Significant differences in spicule
sizes, geometries and skeletal structures justify
their separation contrary to the opinion of Burton
(1959), and followed subsequently by Hooper
and Wiedenmayer (1994). Amorphinopsis
excavans is most closely related to A. foetida in
skeletal structure (both species having relatively
cavernous choanosomal skeletons), differing
from the latter species in spicule geometry and
spicule dimensions.
The specimen described above from the
Vernon Islands is a typical example of this
widely dispersed species, showing close similarities to the type material in its excavating
growth form, and ectosomal and choanosomal
skeletal structure (tangential `echinating' smaller
styles; relatively wide-meshed choanosomal halichondroid reticulation, respectively). Conversely,
it is atypical in having relatively consistent
spicule geometry (most recorded populations of
A. excavans have interchangeable smaller oxeas
and styles composing the tangential ectosomal
skeleton) and in the specific dimensions of
spicules (these are relatively smaller than most
described specimens). Nevertheless, present material clearly belongs to this species.
Amorphinopsis foetida (Dendy)
(Figs 17-18)
Hymeniacidon (?) foetida Dendy, 1889: 8788, pl . 4 fig. 5.
28
Beagle Gulf halichondriid sponges
Fig. 17. Amorphinopsis foelida (Dendy). A, larger choanosomal style;
B, smaller ectosomal style; C, peripheral skeletal structure (surface at left and top);
D, preserved specimen QM G303677; E, live specimen (on deck).
29
J.N.A. Hooper et al.
Fig. 18. Amorphinopsis foetida (Dendy). Specimen QM G303677. A, perpendicular section through skeleton in fistule region,
showing cavernous reticulate construction; B, transverse section of ectosomal skeleton showing outer surface of sponge with a
tangential layer of larger styles echinated by small styles; C, terminations on choanosomal styles; D, terminations on ectosomal
styles.
spicules predominantly supporting the central
portion of fistules, producing a wide-meshed
cavernous reticulation supports the close relationship between the two genera.
This species is now relatively well known in
the literature from several populations in the
Indo-west Pacific (Sri Lanka, South China Sea,
Moluccas and Arafura Sea), probably being a
dominant sponge of the soft, shallow-water
benthos. The specimen described above, the first
record for Australia, agrees closely with Dendy's
(1889) original description, including the characteristic low, volcano-like fistules on the upper
surface, skeletal structure and spicule sizes,
whereas spicule dimensions cited in the Indonesian records are larger than those reported in
other populations. Dendy (1889) also records
that the species has a strong fetid smell (hence
the specific name) but this characteristic has not
been subsequently recorded.
Spicules. Choanosomal styles long, slender,
straight or slightly curved at centre, nearly
hastate (abruptly pointed), occasionally slightly
telescoped points, with evenly rounded bases
(length 448-(609.8)-794 pm, width 8-(16.4)-23
pm). Smaller ectosomal styles slender, straight,
fusiform, with sharp points and evenly rounded
bases (length 67-(106.8)-252 pm, width 3-(5.4)-8
pm).
Remarks. The possession of fistules and styloid spicules in this species suggests it could be
placed in Ciocalypta. However, the fistules in
A. foetida (short, squat, volcano-like) are unlike
those characteristic of Ciocalypta (long, slender, pointed). Moreover, this species has an
ectosomal skeleton with `echinating' smaller
styles over tangential tracts of larger choanosomal styles (whereas in Ciocalypta the smaller
spicules are in bundles and also placed tangentially). Nevertheless, the possession of a core of
30
Beagle Gulf halichondriid sponges
Fig. 19. Amorphinopsis sacciformis (Thiele). Specimen NTM Z22 I 5. A, perpendicular section through skeleton, showing
halichondroid reticulate choanosomal skeleton, oblique tracts in peripheral skeleton and tangential ectosomal layer (bottom of
photo); B, transverse section of ectosomal skeleton showing outer surface of sponge with a tangential layer of larger oxeas
echinated by small styles; C, closer view of tangential ectosomal skeleton; D, sharply pointed terminations on choanosomal oxeas
(left) and ectosomal styles (right).
Amorphinopsis sacciformis (Thiele)
Description. Thinly encrusting under laterite
rock boulders and covering bivalve shell, up to
8 mm thick following contours of substrate.
Oscules not visible.
Colour. Yellow-brown in life (Munsell 7.5
YR 6/8), brown in ethanol.
Texture. Firm, compressible, harsh.
Surface. Hispid, not opaque.
Ectosomal skeleton. Indistinct, very thin surface peel with larger choanosomal oxeas lying
tangential to surface and pointed ends of smaller
ectosomal styles arranged paratangentially
(`echinating') over larger spicules.
Choanosomal skeleton. Confused halichondroid reticulation of larger choanosomal oxeas,
packed densely and bound together by small
amount of collagen, in uni- and paucispicular
bundles. Skeletal meshes oval, usually compressed, and only few larger spaces (lacunae)
(Figs 19-20)
Ciocalypta sacciformis Thiele, 1900: 76, pl.
3, fig. 28 - Hentschel 1912: 125-126; - KellyBorges and Bergquist 1988: 132, pl. 2a.
Ciocalapata sacciformis - de Laubenfels,
1936: 134; - de Laubenfels 1954: 183-184, fig.
122.
Material. NTM Z2215: Vestey's Beach,
Bullocky Point, Darwin, NT, 12°26.2'S,
130°49.9'E, intertidal, 21 January 1985, coll.
J.N.A. Hooper, by hand.
Distribution. Moluccas, Arafura Sea; Palau
Is., central west Pacific; Motupore Island, Papua
New Guinea; and Darwin Harbour, 0-23 m
depth.
Habitat. Intertidal rock and dead coral reef,
frequently mud covered.
31
J.N.A. Hooper et al.
Fig. 20. Amorphinopsis saccifbrmis (Thiele). A, larger choanosomal oxea; B, smaller ectosomal styles;
C, peripheral skeletal structure; D, live intertidal specimen (encrusting under coral boulder and bivalves);
E, preserved specimen NTM Z22I5 on bivalve.
32
Beagle Gulf halichondriid sponges
visible in choanosomal or subectosomal regions.
Subectosomal skeleton with more obliquely directed tracts supporting tangential ectosomal
skeleton.
Spicules. Larger choanosomal oxeas straight
or slightly curved at centre, with long tapering
fusiform sharply pointed ends, in two size classes
(length I 2254318.5)-420 pm, width 5-(7.9)-10
pm; length 11 540-(781.1)-1105 pm, width 11(20.3)-35 pm)). Styles short, slender, fusiform,
sharply pointed, with rounded or occasionally
slightly subtylote bases (length 118-(213.4)-264
pm, width 3-(5.3)-9 pm).
Remarks. This species is atypical of
Amorphinopsis in having a more compressed
choanosomal skeleton than most other species,
and only few small meshes between multispicular
tracts, but this is considered here to be a function of its thinly encrusting (juvenile) growth
form. The possession of smaller `echinating'
styles in the ectosomal skeleton indicates its
affinities with Amorphinopsis rather than with
Ciocalypta as suggested by previous authors.
Amorphinopsis sacciformis is readily differentiated from other species of the genus in having
two size categories of choanosomal oxeas, and
both spicule dimensions and skeletal structures
agree with those originally described for the
species, although architecture is more noticeably wide-meshed in the Indonesian material
(Thiele 1900; Hentschel 1912).
skeleton (or vice versa). Cavities (lacunae) often occuring in choanosomal and subectosomal
regions, with ascending choanosomal spicule
tracts supporting ectosomal crust. Ectosomal
skeleton invariably tangential to surface, usually easily peeled, composed of thin detachable
crust of single spicules or multispicular tracts
forming halichondroid criss-cross reticulation.
Megascleres include predominantly oxeas of
wide size ranges, commonly sharply pointed,
but sometimes strongylote, stylote or rarely true
styloid.
Remarks. There is a wide range of variation
in the development of choanosomal spicule tracts
amongst species of Halichondria, most having
the major tracts obliquely directed or parallel to
the surface. Similarly, development of the
ectosomal skeleton varies from discrete tangential bundles to largely collagenous with fewer
scattered tangential spicules (Van Soest et al.
1990). Halichondria, like Hymeniacidon, is
characterised by the possession of a thin detachable ectosomal membrane bearing a unispicular,
tangential, halichondroid criss-cross of oxeas,
or sometimes vague bundles of spicule tracts
lying tangential to the surface. The two genera
are differentiated by their spicule geometry, having predominantly oxeas or exclusively styles,
respectively.
Many species of Halichondria have been described from all oceans, although the genus has
diversified to a larger extent in colder seas,
including polar regions, and found predominantly in shallower waters (Diaz et al. 1991).
Within the Indo-Australian region, 15 species
have been described so far: H. aciculata (Carter),
H. arenacea Dendy, H. claviformis Carter, H.
nigrocutis (Carter) and H. schmidti (Dendy)
from Bass Strait, Vic. (Carter 1885, 1886; Dendy
1895; Ayling et al. 1982); H. latrunculioides
Ridley and Dendy and H. prostata Thiele from
the Australian Antarctic Territory (Ridley and
Dendy 1887; Thiele 1905; Hentschel 1914;
Koltun 1964, 1976); H. lycopodium (Esper) and
H. cartilaginea (Esper) from unspecified localities in northern Australia and Southern Indonesia (Topsent 1933); H. mollissima (Lendenfeld)
from central NSW (Lendenfeld 1888); H.
phakellioides Dendy and Frederick from the
Houtman Abrolhos and Perth regions, WA
(Dendy and Frederick 1924); H. tyleri (Bowerbank) widely distributed in northern Australia
and Indonesia (Marshall 1892; Hentschel 1912;
Burton 1934); H. mertoni (Hentschel) and H.
Genus Halichondria Fleming
Halichondria Fleming, 1828: 520.
Raspaigella Schmidt, 1868: 25.
Eumastia Schmidt, 1870: 42.
Spuma Miklucho-Maclay, 1870: 13.
Menanetia Topsent, 1896: 115.
Pyloderma Kirkpatrick, 1908: 51.
Halichondriella Burton, 1931: 137.
Trachyopsilla Burton, 1931: 138.
Ciocalapata de Laubenfels, 1936: 134.
Ciareamastia de Laubenfels, 1942: 265.
Type species. Spongia panicea Pallas, 1766,
by original designation.
Diagnosis. Massive amorphous growth forms.
Choanosomal skeleton composed of poorly defined directionless spicule tracts; often two components recognisable: vaguely reticulate
multispicular tracts and halichondroid crisscrossed single spicules dispersed between the
major tracts, but major tracts may be sustantially
reduced in favour of unispicular halichondroid
33
J.N.A. Hooper et al.
(Hentschel) from the Am Islands,
Indonesia (Hentschel 1912); and H. melichlora
(Sollas) from Straits of Malacca and Arafura
Sea (Sollas 1902; Hentschel 1912) .
Description. Flattened or massive burrowing
base extending below substrate for up to 30 mm,
with long, thick, tapering fistules protruding
above substrate, up to 125 mm long, 15 mm
thick at base. Oscules surmounted on apex of
most fistules, although some fistules blind and
merely sharply pointed.
Colour. Live colouration mottled mauvebrown (Munsell 2.5R 5/6) and mauve-green
(5Y 5/4) fistules and beige base (5Y 8/4); mauvebeige in ethanol.
Texture. Compressible, flexible, relatively
easily torn.
Surface. Opaque and membranous with drainage channels running down sides of fistules,
visible through ectosome. External surface even,
slightly conulose and minutely hispid.
Ectosomal skeleton. Thick, continuous, dense
spiculous layer, easily peeled from sponge surface, with multispicular tangential skeletal tracts
composed mainly of larger oxeas, overlayed by
unispicular paratangential tracts or plumose tufts
of smaller oxeas. Ectosomal skeleton associated
with moderate quantities of pigmented collagen.
Choanosomal skeleton. Interior of
choanosome a confusion of single spicules,
mostly large, becoming more organised into
radial bundles of spicules towards surface. In
some specimens, medium-sized spicules forming distinct tracts approximately 20-50 spicules
in width, traversing subdermal lacunae perpendicular to surface. In fistules choanosomal
skeleton compressed, confined to central portion of skeleton, with huge subectosomal spaces
(up to 5 mm diameter) and only few large subectosomal multispicular tracts support ectosomal
peel. Collagen present in interior of choanosome
and closely associated with spicule tracts.
Spicules. Two size classes of oxeas: larger
choanosomal oxeas slightly or moderately curved
at centre, tapering fusiform, sharply pointed,
occasionally asymmetrical anisoxeote (length
423-(536.7)-665 pm, width 8-(14.4)-22 pm);
smaller ectosomal oxeas straight or slightly
curved at centre, sharply pointed fusiform (length
150-(205.4)-312 pm, width 3-(4.2)-5 pm).
Remarks. Under the traditional definitions
of halichondrids, this species would have been
included in Ciocalypta, having a fistulose growth
form, plumose ectosomal skeleton of smaller
spicules overlaying a tangential skeleton of larger
choanosomal megascleres, and with supporting
spicules located mainly at the core of fistules (as
stalagmites
Halichondria tyleri (Bowerbank)
(Figs 21-22)
Ciocalypta tyleri
Bowerbank, 1873: 21, pl. 4,
figs 9-12.
Ciocalypta penicillus - Burton 1934: 564;
Ciocalypta penicillus
Burton 1959: 264 (not
Bowerbank, 1864: 180).
Ciocalypta penicillus var. gracilis Topsent,
1897: 445, pl. 18, figs 6-7.
Ciocalypta tyleri var. manaarensis Dendy,
1889: 91.
Apatospongia fallax Marshall, 1892: 16, pl.
8, figs 1-5.
Type material. SYNTYPE - BMNH
1930.7.3.32: Port Elizabeth, South Africa, coll.
Capt. Charles Taylor.
Other Material. - NTM Z592: Table Head,
Port Essington, Cobourg Peninsula, NT,
11°13.5'S, 132°10.5'E, 3 m depth, 04 May 1982,
coll. J.N.A. Hooper, SCUBA. NTM Z3286: SW
side of cliff face, Table Head, Port Essington,
11°13.5'S, 132°10.5'E, 5 m depth, 11 September 1986, coll. J.N.A. Hooper and C. Johnson,
SCUBA. NTM Z1358: Coral Bay, Port
Essington, 11°11.3'S, 132°03.75'E, 6 m depth,
16 May 1983, coll. J.N.A. Hooper, SCUBA.
NTM Z1395: same locality, 11°10.4'S,
132°02.8'E, 2m depth, 19 May 1983, coll. J.N.A.
Hooper, SCUBA. NTM Z941: East Point
`bommies', Darwin, NT 12°24.05'S,
130°48.0'E, 12 m depth, 13 September 1982,
coll. J.N.A. Hooper, SCUBA. NTMZ3133: Parry
Shoals, 11°11.41'S, 129°43.01'E, 18m depth,
13 August 1987, coll. A.M. Mussig and NCI
AIMS, SCUBA. NTM Z3410: 500 m east of
Hermite Island, Monte Bello Islands, WA,
20°27.6'S, 115°32.8'E, 8m depth, 29 August
1988, coll. D. Low Choy and NCI AIMS,
SCUBA.
Distribution. South Africa, Red Sea, Gulf of
Manaar, Indo-Malayan region, northern Australia; Turtle Islands, North Direction Island,
Great Barrier Reef, Q1d.; East Point, Darwin
Harbour; Parry Shoals, Arafura Sea; Port
Essington, Cobourg Peninsula, NT; Monte Bello
Islands, WA.
Habitat. Base of coral reefs, sandy substrate;
intertidal to 81 m depth.
34
Beagle Gulf halichondri id sponges
E
O
Fig. 21. Halichondria tyleri (Bowerbank). A, larger choanosomal oxeas; B, smaller ectosomal oxea; C, peripheral skeletal
structure; D, preserved specimen NTM Z592; E, preserved specimen NTM Z1358.
Topsentia, Ciocalypta, and Petromica, as well
as many other sponge orders.
Burton (1959) synonymised many quite different species, including Ciocalypta tyleri, under the name C. penicillus. Most of this synonymy is now rejected following examination of
relevant type material. Of relevance to the Australian fauna, Hooper and Wiedenmayer (1994)
listed several species in synonymy with H. tyleri,
following Burton (1959), but these can now be
opposed to peripheral supporting spicules found
in Petromica). However, given that all spicules
are oxeas (not styles), this species is referred to
Halichondria under the revised criteria of Van
Soest et al. (1990) and Diaz et al. (1991) which
hypothesises that fistules are an ecological adaptation for colonising soft substrata, and consequently they have been independently developed throughout various genera of Halichondriidae, and are also found in some species of
35
J.N.A. Hooper et al.
Fig. 22. Halichondria tyleri (Bowerbank). Specimen NTM Z94I. A, perpendicular section through skeleton of fistule, showing
choanosomal skeleton restricted to central portion of fistules, extremely cavernous subectosomal skeleton and thick paratangential
ectosomal skeleton; B, transverse section of ectosomal skeleton showing outer surface of sponge with a multispicular and
unispicular tangential skeletons; C, light micrograph of section through choanosomal skeleton showing compressed halichondroid reticulation in central portion of skeleton and halichondroid plumose subectosomal region; D, sharply pointed spicule
terminations on larger oxeas (left) and smaller oxeas (right).
allocated more accurately now that we have had
access to type material. Of Burton's (1959)
synonymised species, Ciocalypta heterostyla
Hentschel (1912) and C. rutila gracilisHentschel
(1912) (which belong to Ciocalypta), and C.
mertoni Hentschel (1912) (which belongs to
Halichondria), are not fistulose, they do not
have exclusively oxeote spicules, and are not
closely related to H. tyleri. Ciocalypta penicillus
aciculata Carter (1885) is fistulose, with tapering "sub-pinlike" spicules (which are subacerate
styles, with the thickest part of the spicule near
the point rather than the base), and lacks the
specialised tangential ectosomal skeleton characteristic of Ciocalypta. It is probably most appropriately referred to Halichondria, although
this needs to be established from recollection of
fresh material. Stylotella polymastia Lendenfeld
(1888) is also a Ciocalypta (Hamann, 1914),
and not closely related to H. tyleri. Sigmaxinella
papillata Brondsted (1923), Axinella colvilli
Brondsted (1924) and Hymeniacidon novaezelandiae Brondsted (1924) have both large
choanosomal styles and small ectosomal oxeas
and a fistulose growth form. Bergquist (1970)
subsequently included these three species from
New Zealand into synonymy with the European
species Ciocalypta penicillus Bowerbank, but
this conspecificity is very unlikely. They probably represent a single, distinct New Zealand
species, possibly best allocated toAmorphinopsis
or Ciocalypta (depending on whether ectosomal
spicules are merely tangential or echinating).
The name C. papillata (Brondsted) is the oldest
available name for this taxon. As a consequence
of rejecting Burton's (1959) synonymy, Halichondria tyleri is restricted now to Indo-west
Pacific populations, extending into the Pacific
36
Beagle Gulf halichondriid sponges
Fig. 23. Halichondria vansoesti sp. nov. Holotype NTM Z2678. A, perpendicular section through skeleton showing distinctive
cavernous, reticulate subectosomal skeleton and thin tangential ectosomal peel; B, transverse section of ectosomal skeleton
showing outer surface of sponge with a mostly uni- or paucispicular tangential layer; C, light micrograph of section through
deeper choanosomal skeleton showing halichondroid reticulation of multispicular tracts; D, faintly telescoped points of larger
oxeas (left) and smaller oxeas (right).
cific attributes distinguish it from other fistulose
species including those mentioned above and H.
vansoesti sp. nov.
only as far as the northern Great Barrier Reef.
Several other species with similar growth forms
but differing in various aspects of their
spiculation and skeletal structures have been
recently collected from central, southern Queensland and New South Wales waters, and these
represent distinct sibling species, some at least
still undescribed. These will be dealt with in a
second part to this present contribution (Hooper
et al., in prep.).
Halichondria tyleri is distinctive in its burrowing fistulose growth form, with sharply
pointed fistules with small terminal oscules,
mauve-brown live colouration, slightly conulose
surface with clearly visible subectosomal drainage canals, cavernous skeletal construction in
the peripheral skeleton (not in the central portion of the skeleton), exclusively oxeote spicules
both of moderate size range (mean 540 x 14 pm
and 205 x 4 pm), and sharply pointed spicule
terminations (no telescoped endings). These spe-
Halichondria vansoesti sp. nov.
(Figs 23-24)
Type material. HOLOTYPE - NTM Z2648:
East Point, Darwin, NT, 12°24.5'S, 130°48.0'E,
12m depth, 3 April 1986, coll. J.N.A. Hooper
and A.M. Mussig, SCUBA. PARATYPE - QM
G303450: W. side Fish Reef, Bynoe Harbour,
NT, 12°26.1'S, 130°26.5'E, 11 m depth, 26
September 1993, coll. J.N.A. Hooper and L.J.
Hobbs, SCUBA.
Material. NTM Z84: Coral Bay, Port
Essington, Cobourg Peninsula, NT, 11°11.5'S,
132°02.0'E, 4 m depth, 18 October 1981, coll.
J.N.A. Hooper and P.N. Alderslade, SCUBA.
NTM Z2678: East Point, Darwin, NT,
12°24.50'S, 130°48.0'E, 12 m depth, 3 April
37
J.N.A. Hooper er al.
O
O
Fig. 24. Halichondria vansoesti sp. nov. A, larger choanosomal oxeas; B, smaller ectosomal oxeas;
C, peripheral skeletal structure; D, preserved specimen NTM Z84; E, live subtidal paratype QM G303450.
38
Beagle Gulf hahchondriid sponges
1986, coll. J.N.A. Hooper and A.M. Mussig,
SCUBA. QM G300815: E of Gove Peninsula,
Gulf of Carpentaria, Qld, 12°37.1'S, 136°49.3'E,
26 m depth, 23 November 1991, coll. S.J. Cook,
RV Southern Surveyor, dredge.
Distribution. East Point, Darwin Harbour;
Fish Reef, Bynoe Harbour; Cobourg Peninsula,
NT; eastern Gulf of Carpentaria.
Habitat. Rocky reef, reef slope, sandy
substrate; subtidal 4-26 m depth.
Description. Flattened base burrowing only
shallow distance into substrate; top of base with
long, tapering, pointed fistules protruding up to
250 mm above substrate; base approximately 25
mm thick, fistules up to 200 mm long, 25 mm
diameter at base. Oscules large, up to 3 mm
diameter in preserved state, located on basal
plate in between fistules.
Colour. Completely unpigmented, white in
life and in ethanol.
Texture. Firm, compressible, fistules flexible
but easily torn and detachable from basal plate.
Surface. Transparent, parchment-like membrane with subectosomal cavities clearly visible
beneath. Surface of fistules regularly conulose,
minutely hispid, with longitudinal striations visible subectosomally (presumably peripheral skeletal tracts running longitudinally from base to
tip of fistule).
Ectosomal skeleton. Detachable relatively
thick peel formed by halichondroid reticulation
of mostly smaller ectosomal oxeas, in uni- or
paucispicular tracts, lying tangential to surface.
Ectosomal skeleton varying greatly in thickness, thicker in region of base than at apex of
fistules. In addition to tangential skeleton, sparse
plumose tufts of smaller spicules occurring at
junction where choanosomal tracts meet
ectosome, standing paratangential to and just
protruding through surface, producing prominent surface conules.
Choanosomal skeleton. Confused halichondroid skeleton of mostly directionless multispicular tracts, 20-50 spicules in width, with
some single spicules also dispersed between
tracts. Spicule tracts composed of both large
and small oxeas, becoming more perpendicular
closer to surface and forming distinctive cavernous subectosomal region. Moderate quantities of collagen present in mesohyl associated
with spicule tracts.
Spicules. Oxeas in two size classes. Larger
choanosomal oxeas straight or slightly curved
at centre, fusiform, with slightly telescoped
points (length 470-(558.3)-615 pm, width 8-
(11.5)-15 pm). Smaller ectosomal oxeas straight
or very slightly curved at centre, fusiform, with
slightly telescoped points (length 200-(265.7)425 pm, width 2-(3.8)-6 pm).
Etymology. For Dr Rob Van Soest, Zoological Museum, University of Amsterdam, for extensive contributions to the systematics of
sponges, particularly the contemporary classification of Halichondriidae.
Remarks. Halichondria vansoesti and H.
tyleri are sibling species sharing a burrowing,
fistulose growth form and comparable spicule
sizes (mean lengths and widths 560 x 11 pm
and 265 x 4 pm, versus 540 x 14 pm and 205 x
4 pm, for choanosomal and ectosomal oxeas,
respectively). They are clearly not conspecific,
differing substantially in their field characteristics (the fistules of H. vansoesti being consistently white alive, whereas those of H. tyleri are
mauve-brown), ectosomal skeletal structure
(marginally thicker in H. vansoesti), choanosomal skeletal structure (whereby in H. vansoesti the most cavernous part of the fistulose
skeleton is found near the periphery, and in this
region spicule tracts become reticulate, whereas
in H. tyleri the deeper, central choanosomal
region is prominently cavernous whereas the
peripheral, subectosomal skeleton is dense and
radial in construction), and spicule geometry
(with telescoped points on spicules as opposed
to sharply pointed spicules).
Halichondria phakellioides
Dendy and Frederick
(Figs 25-26)
Halichondria phakellioides Dendy and
Frederick, 1924: 498-499, pl. 26, fig. 10. - Burton
1934: 600, pl. 2, fig. 1.
Type material. HOLOTYPE - BMNH
1925.11.1.185: Sandy Isle, Houtman-Abrolhos,
WA, coll. W. Dakin.
Other Material. NTM Z900: East Point
Aquatic Reserve, off Dudley Point, Darwin, NT,
12°25.0'S, 130°48.04'E, 10 m depth, 31 August
1982, coll. J.N.A. Hooper, SCUBA. NTM
Z2063: same locality, 12°25.0'S, 130°48.4'E, 610 m depth, 10 May 1984, coll. J.N.A. Hooper
and P.N. Alderslade, SCUBA. NTM Z2255:
same locality, 12°24.5'S, 130°48.0'E, 10 m
depth, 12 April 1985, coll. J.N.A. Hooper and C.
Johnson, SCUBA. NTM Z2610, 2623, 2643,
2652, 2674: same locality, 12°24.5'S, 130°48.0'E,
9-12 m depth, 03 April 1986, coll. J.N.A. Hooper
and A.M. Mussig, SCUBA. NTM Z935: same
39
J.N.A. Hooper et al.
Description. Erect, buttressed flattened ridges.
Base fixed to substrate by basal plate, attached
to fragments of coral or shell, usually burrowing into soft substrate; ridges ('fistules') thick,
flat (up to 220 mm long, 50 mm wide, 30 mm
thick) growing in several planes, anastomosing
and becoming buttressed or digitate in some
specimens. Oscules small (330-500 pm diameter) but conspicuous and scattered on lateral
surfaces of ridges, each with slightly raised membraneous lip.
Colour. Yellow-brown to pale orange alive
(Munsell 2.5Y 8/10 - 5YR 7/10), beige to grey
in ethanol.
Texture. Harsh, firm but compressible, can
be torn.
Surface. Surface smooth, even between large
di g itate projections (ridges); usually membraneous and clearly detachable (although in some
material surface roughened and minutely
conulose due to poor preservation). Surface membrane translucent, with subectosomal lacunae
and drainage channels clearly visible below.
Ectosomal skeleton. Ectosomal skeleton consisting of distinct tangential crust of both single
spicules and bundles of mostly medium-sized
spicules, forming prominent 'skin'. Paratangential tracts of subectosomal spicules meeting
tangential surface layer at regular intervals, giving ectosome patchy or striated appearance in
cross-section; ascending spicule tracts may protrude slightly through tangential layer. In transverse section, ectosome nearly regularly reticulate, with cross-hatched spicule tracts five to
eight spicule widths (20-50 pm in diameter)
forming oval meshes 80-15 pm in diameter,
distinctly detachable and clearly offset from cavernous subectosomal skeleton. Small amounts
of collagen present in ectosomal layer bonding
tangential spicule tracts.
Choanosomal skeleton. Skeletal structure
halichondroid reticulate in deeper choanosomal
skeleton and cavernous reticulate in subectosomal peripheral skeleton. Deeper choanosomal
region more disorganised than peripheral skeleton, largely composed of single spicules forming directionless halichondroid reticulation
(criss-cross), comprising both large and small
oxeas, whereas only large and medium-sized
spicules comprise tracts or bundles of approximately 20-50 spicules in width (150-250 pm in
diameter). Multispicular tracts forming prominent ascending lines arising to surface, with
tracts aligned perpendicular to surface, support-
locality, 12°24.05'S, 130°48.0'E, 12 m depth, 13
September 1982, coil. J.N.A. Hooper, SCUBA.
QM G303320: East Point `bommies', East Point
Aquatic Life Reserve, 12°24.5'S, 130°48.8'E,
10m depth, 23 September 1993, coll. J.N.A.
Hooper and L.J. Hobbs, SCUBA. QM G303261:
S of South Shell I., East Arm, 12°29.8'S,
130°53.5'E, intertidal, 19 September 1993, coll.
J.N.A. Hooper and L.J. Hobbs, by hand. NTM
Z241: Indian I., 12°35.0'S, 130°33.0'E, 3 m
depth, 18 November 1981, coll. Byers, P., FV
Skelton, snorkel. QM G303448: Fish Reef, W
side near wreck of HMAS Brisbane, Bynoe Harbour, NT 12°26.1'S, 130°26.5'E, 1 lm depth, 2
September 1993, coll. J.N.A. Hooper and L.J.
Hobbs. NTM Z3929: Bay N side of Cumberland
Strait, Wessel Is, NT 11°27.5'S, 131°28.8'E, 20
m depth, 14 November 1990, coll. J.N.A.
Hooper, SCUBA. NTM Z621: Unnamed shoal, N
of Melville I., NT, 11°38.23'S, 129°51.0'E, 24 m
depth, 17.v.1982, coll. R. Lockyer and B. Thom,
SCUBA. NTM Z3077: Parry Shoals, 11°11.41'S,
129°43.01'E, 18 m depth, 13 August 1987, coll.
A.M. Mussig and NCI AIMS, SCUBA. NTM
Z3139: same locality, 11°12.28'S, 129°42.71'E,
16m depth, 14 August 1987, coll. A.M. Mussig
and NCI AIMS, SCUBA. NTM Z3075: same
locality, 11°11.72'S, 129°43.26'E, 16 m depth,
12 August 1987, coll. Mussig, AM and NCI
AIMS, SCUBA. NTM Z3349: 8 km from shore,
NW corner of Herald Reef, Coolga Point, WA,
21°29.5'S, 115°17.1'E, 5 m depth, 25 August
1988, coll. D. Low Choy and NCI AIMS,
SCUBA. NTM Z3395: 50 m from shore, 3 km
west of Hermite Island, Monte Bello Islands,
20°27.1'S, 1 15°34.2'E, 6 m depth, 29 August
1988, coll. D. Low Choy and NCI AIMS,
SCUBA. NTM Z3361: 5 km from shore, Flinders
Shoals, N of Mary Anne Island, Mary Anne Passage, WA, 25°14.2'S, 115°28.2'E, 8 m depth, 25
August 1988, coll. D. Low Choy and NCI AIMS,
SCUBA.
Distribution. Swan River region and
Houtman-Abrolhos, southern WA; Herald Reef,
Mary Anne Island and Monte Bello Islands,
northern WA; East Point, Dudley Point and
East Arm, Darwin Harbour; Indian Island and
Fish Reef; Bynoe Harbour; Wessel Islands, Gove
Peninsula; Parry Shoals and Melville Island,
Arafura Sea, NT; ? unspecified locality, Great
Barrier Reef, Qld.
Habitat. Coral rubble and soft substrates,
usually between patches of reef; intertidal to 30
m depth.
40
Beagle Gulf halichondriid sponges
Remarks. This species is probably widely
distributed in tropical and subtropical waters of
Australasia, although its recorded distribution
so far is restricted to the southwest and northwest coasts, from the Perth region, WA, to the
Wessel Islands, NT. The single Pacific record is
based on a specimen collected by Saville Kent
(Burton 1934), with no specific locality data
given. This record remains unsubstantiated, although Burton (1934) states that the Queensland specimen agrees in all details to the
holotype.
One of the prominent features of H.
phakellioides is the clearly detachable tangential ectosomal skeleton, which peels off the sur-
ing ectosomal skeleton, and traversing large
subdermal cavities; cavities (lacunae) varying
greatly in size between specimens ranging from
200-2000 pm diameter. Spongin fibres not evident but sparse collagen present in mesohyl
closely associated with spicule tracts.
Spicules. Oxeas in three size classes, all fusiform, slightly curved at centre or straight; larger
ones usually faintly telescoped, sometimes
sharply pointed or rarely styloid; smaller ones
include both prominently telescoped and sharply
pointed forms (I: length 314-(444.6)-578 pm,
width 10-(13.1)-20 pm; II: length 195-(230.6)275 pm, width 4-(6.9)-12 pm; III: length 58(196.8)-266 pm, width 1-(2.2)-3 pm).
ABC
Fig. 25. Halichondria phakellioides Dendy and Frederick. A, larger oxeas; B, medium sized oxeas; C, smaller oxeas;
D, peripheral skeletal structure; E, preserved specimen NTM Z900; F, live subtidal specimen NTM Z2610.
41
J.N.A. Hooper et al.
Fig. 26. Halichondria phakellioides Dendy and Frederick. Specimen NTM Z2255. A, perpendicular section through skeleton of
erect ridge, showing distinctive cavernous and reticulate nature of subectosomal skeleton and spicule columns supporting thin
tangential ectosomal skeleton; B, transverse section of ectosomal skeleton showing outer surface of sponge with a tangential layer
of mostly multispicular tracts composed of both larger and smaller oxeas, forming cross-hatched reticulation; C, light micrograph
of section through deeper choanosomal skeleton showing halichondroid reticulation of mostly single spicules; D, telescoped
terminations on larger oxeas (left) and smaller oxeas (right).
face relatively easily, exposing a cavernous reticulate skeleton beneath (often infested with
shrimps and polychaete worms). Initially this
species was subdivided into two morphs, one
with and the other without a detachable
ectosomal skeleton, but it was eventually recognised that this feature was largely dependent on
the state of preservation of material (in badly
preserved material the surface was minutely
conulose, lacking the tangential ectosomal peel).
Another important feature of this species, making it easily recognisable in the field, is the erect
buttressed growth form and yellowish live
colouration (although in deeper or murkier waters pigments may be absent entirely), and harsh
texture of the ectosomal skeleton. In its erect,
buttressed growth form this species is very similar to Leucophloeus massalis Carter from southern Australia (in particular to an unpublished
variety alata of Carter's (1883) in the BMNH,
with 'hypotypes' BMNH 1886.12.15.454 and
473 from Fremantle, WA). Leucophloeus
massalis, however, has much thicker lamellae
than H. phakellioides, and exclusively stylote
megascleres which places it in Ciocalypta
(Hooper and Wiedenmayer 1994). Other species referred to Halichondriidae at one time or
another with similar erect buttressed growth
forms include: L. compressus Carter (1883) from
Westernport Bay, Victoria. (syntype BMNH
1886.12.15.9) and Axinella solida Carter, (1885)
from Bass Strait, Vic. (lectotype BMNH
1887.7.11.24); L. compressus belongs to
Pseudoclathria (family Crellidae) (Dendy 1897),
and A. solida belongs to Rhaphoxya (family
Axinellidae) (Hooper and Levi 1993b).
Three spicule sizes are differentiated in the
description of H. phakellioides, although realistically only two sizes are clearly distinguished
(the smallest category probably being juvenile
42
Beagle Gulf halichondriid sponges
Fig. 27. Halichondria stalagmites (Hentschel). Specimen NTM Z131. A, perpendicular section through skeleton of fistule,
showing multispicular halichondroid reticulate skeleton, large lacunae near in the subectosomal region, and thin tangential
ectosomal peel on convoluted part of surface (top of photo); B, transverse section of ectosomal skeleton showing outer surface of
sponge with a thin tangential layer of both larger and smaller oxeas without apparent difference in their distribution; C, light
micrograph of section through peripheral skeleton showing extensive canal system in subectosomal region; D, slightly telescoped
terminations on larger oxeas (left) and smaller oxeas (right).
representatives of the medium-sized oxeas).
There is no clear size-class localisation of spicules (such as seen in Amorphinopsis and
Ciocalypta), although smaller spicules are mostly
found scattered between the multispicular tracts.
NTM Z1097: East Point, Darwin, NT, 12°25.0'S,
130°48.4'E, 6-7 m depth, 22 December 1982,
coll. J.N.A. Hooper, SCUBA. NTM Z2651: same
locality, 12°24.5'S, 130°48.0'E, 9-12 m depth, 3
April 1986, coll. J.N.A. Hooper, A.M. Mussig,
A. Howard and C. Hood, SCUBA. NTM Z1991:
W side Weed Reef, Darwin, NT, 12°29.2'S,
130°47.1'E, 8 m depth, 11 May 1984, coll.
J.N.A. Hooper, P.J. Homer and P.N. Alderslade,
SCUBA. NTM Z3147: Parry Shoals, Arafura
Sea, NT, 11°12.53'S, 129°42.08'E, 16 m depth,
15 August 1987, coll. A.M. Mussig and NCI,
SCUBA.
Distribution. Arafura Sea, Indonesia; Darwin Harbour, Cobourg Peninsula and Parry
Shoals, Arafura and Timor Seas, NT.
Habitat. Sand, mud, coral rubble, usually
with base burrowing slightly into substrate; 616 m depth.
Halichondria stalagmites (Hentschel)
(Figs 27-28)
Ciocalypta stalagmites Hentschel, 1912: 126,
pl. 21, fig. 60.
Type material. SYNTYPE - SMF 1595:
Meriri and Leer Is., Aru Is, Arafura Sea, Indonesia, 31 March 1908, coll. H. Merton, 10 m
depth, dredge.
Other Material. NTM Z131: Sandy Island
No. 2, Cobourg Peninsula, NT, 11°05.5'S,
132°17.0'E, 10 m depth, 21 October 1981, coll.
J.N.A. Hooper and P.N. Alderslade, SCUBA.
43
J.N.A. Hooper et al.
B
Fig. 28. Halichondria stalagmites (Hentschel). A, larger oxeas; B, smaller oxeas; C, peripheral skeletal structure; D, preserved
specimen NTM Z3I47; F, live subtidal specimen NTM Z1097.
Description. Growth form generally cushion-shaped base with fistulose apex, but specifically ranging from elongate, flattened palmatebranching (up to 160 mm high), with slightly
flattened, spreading, lobate-digitate branches (up
to 30 mm wide, 10 mm thick), slightly convoluted digits on lateral sides of sponge (presumed
older specimens), to subspherical massive habits (up to 70 mm diameter), with tapering, blind
fistule-like digits on upper surface, base attached
to hard object (coral, shells) burrowing slightly
into substrate (presumed younger specimens).
Oscules discrete, conspicuous, approximately 3
mm diameter, found at base of larger fistules, or
on lateral surface in flattened, branching specimens.
Colour. Orange-brown in life (Munsell 5YR
7/10-6/10), pale brown in ethanol.
Texture. Soft, compressible, spongy, easily
torn. Fistules easily collapsable upon collection.
Surface. Fistules opaque, collagenous, slightly
microconulose; surface even, unornamented be-
44
Beagle Gulf hatichondriid sponges
nesian population). In all other respects, including the possession of telescoped points on spicules, northwest Australian and Indonesian
populations are comparable.
tween fistules; surface of burrowing mass generally membraneous and translucent through
which cavernous choanosome visible.
Ectosomal skeleton. Thin tangential layer of
unispicular tracts or multispicular tracts (two to
three spicules abreast) composed of both smaller
and larger choanosomal oxeas, without apparent localisation, bound together with granular
collagen; spicules becoming plumose bundles
in places, with oxeas occasionally protruding
through ectosome, but mostly lying tangential
to surface.
Choanosomal skeleton. All histological
preparations contained cyanobacteria. Basal
mass containing disorganised halichondroid
unispicular reticulation of both larger and
smaller oxeas, supported by occasional
multispicular (`reticulate') tracts of larger oxeas;
towards subectosomal region larger spicules producing more prominent multispicular bundles
containing three to five spicules per tract; these
becoming oriented perpendicular to surface near
peripheral skeleton, occasionally protruding
through it. Choanosomal region cavernous
throughout, particularly in fistules, with lacunae between 100-300 pm in diameter in deeper
choanosomal region of basal mass but larger
(up to 500 pm) in subectosomal region of fistules.
Collagen relatively abundant in mesohyl, in both
basal mass and fistules, but no spongin fibres
visible.
Spicules. Oxeas, fusiform, slightly telescoped
or less frequently sharply pointed, in two size
classes although not restricted to any particular
location within skeleton (length I 335-(478.3)675 pm, width 7-(11.4)-15 pm; length II 88(171.2)-325 pm, width 2-(3.7)-5 pm).
Remarks. This species is apparently widely
distributed throughout the Aru and Kai Islands
of Indonesia, and is also relatively common
throughout the Darwin region, although this is
area appears to be its southernmost extent in
distribution. The species is easily recognisable
in the field by its orange-brown colouration and
cushion-shaped fistulose growth form.
The northwest Australian population differs
slightly from type material and the description
of Hentschel (1912) in the extent to which the
ectosomal skeleton is developed (represented
only by a thin tangential peel in material described above, but more prominent in Hentschel's
specimens), and the upper range of length of
choansomal oxeas (675 pm long in present material whereas up to 1040 pm long in the Indo-
Halichondria bergquistae sp. nov.
(Figs 29-30)
Type material. HOLOTYPE - QM G303351:
East Point `bommies', Darwin, NT, 12°24.5'S,
130°48.8'E, 10 m depth, 23 September 1993,
coll. J.N.A. Hooper, L.G. Hobbs and B. Alvarez,
SCUBA.
Distribution. East Point Aquatic Reserve,
Darwin Harbour.
Habitat. Fringing coral reef, subtidal.
Description. Massive-lobate, bulbous-digitate
(resembling a Psetidaxinella, family Axinellidae), 70 mm long, 55 mm wide, up to 45 mm
thick. Oscules small (up to 3 mm diameter),
conspicuous, discrete, with slightly raised membranous lip, mainly on apical surface of sponge.
Colour. Dark purple-mauve alive (Munsell
5RP 3/6); light brown in ethanol.
Texture. Firm, compressible.
Surface. Prominently hispid, conulose, shaggy
in appearance, with an opaque, membranous,
skin-like surface covering stretched between adjacent conules.
Ectosomal skeleton. Membranous, hispid
with brushes of larger spicules protruding
through surface, paratangential to it, supporting a sparse thin, tangential layer of smaller
oxeas on external surface.
Choanosomal skeleton. Deeper parts of skeleton consisting of halichondroid reticulate architecture of single spicules or paucispicular
tracts composed of larger oxeas. In subectosomal region spicule tracts more obvious, running predominantly longitudinally, ascending,
becoming more radial, plumo-reticulate near
periphery, with tracts composed mainly of larger
oxeas and thinner oxeas also scattered throughout mesohyl. Spicule tracts uni- or paucispicular,
loosely bound together with sparse collagen,
without any obvious spongin fibres. Collagen in
mesohyl light, nearly unpigmented. Choanocyte
chambers ovoid, 40-50 pm diameter.
Spicules. Oxeas in two size classes: larger
(predominantly choansomal) oxeas slightly
curved at centre, tapering to fusiform sharp
points (length 610-(797.2)-855 pm, width 15(21.6)-32 pm); smaller (predominantly ectosomal) oxeas straight or slightly curved, fusiform
45
J.N.A. Hooperet al .
O
O
Fig. 29. Halichondria bergguistae sp. nov. A, larger (predominantly) choanosomal oxea; B, smaller
(predominantly) ectosomal oxeas; C, peripheral skeletal structure; D, preserved holotype QM G303351;
E, live subtidal holotype.
46
Beagle Gulf halichondriid sponges
Fig. 30. Halichondria bergquistae sp. nov. Holotype QM G303351. A, perpendicular section through skeleton showing
halichondroid reticulation of directionless pauci- and unispicular tracts, becoming more plumo-reticulate in subectosomal region,
protruding through and supporting the thin tangential ectosomal crust; B, transverse section of ectosomal skeleton showing outer
surface of sponge with protruding larger oxeas and sparse tangential smaller oxeas (with an ostium, upper right); C, light
micrograph of section through peripheral skeleton showing plumo-reticulate subectosomal skeletal tracts; D, sharply pointed
spicule terminations on both larger oxeas (left) and smaller oxeas (right).
sharply pointed (length 382-(509.5)-675 pm,
width 5-(7.4)-12 pm).
Etymology. For Professor Patricia Bergquist,
University of Auckland, for her unparalleled
contributions to contemporary debate on sponge
biology and systematics, and specific contributions to the taxonomic knowledge of Australasian sponges, including material from Darwin
Harbour (Bergquist and Tizard 1967).
Remarks. This species was initially thought
to belong to H. melichlora (Sollas) (from the
Straits of Malacca (Sollas 1902) and Am Island, Indonesia (Hentschel 1912)) based on descriptions of growth form, spicule geometry,
possession of a slightly radial peripheral skeleton, and range of spicule sizes. But for several
reasons this is not now accepted. Unfortunately,
the holotype of H. melichlora has not yet been
located, and both Sollas' (1902) and Hentschel's
(1912) descriptions are fairly brief. However,
from both these authors we know that H. melichlora has a massive growth form with flattened surface processes, a smooth unsculptured
surface, a longitudinal central skeleton, a plumose subectosomal skeleton and a cavernous
peripheral skeleton, and published spicule dimensions ranging from 350-960 x 30-42 pm for
larger oxeas and 168-208 x — pm for smaller
oxeas. By comparison, the growth form of H.
bergquistae is lobate, bulbous-digitate, its surface is prominently conulose, shaggy (very reminiscent of Pseudaxinella), the choanosomal
skeleton is halichondroid reticulate, becoming
more radial towards the periphery, but without
cavernous construction, and spicule dimensions
are 610-855 x 15-32 pm for larger oxeas and
382-675 x 5-12 pm for smaller oxeas. On the
basis of these differences it is warranted to recognise H. bergquistae and H. melichlora as
distinct taxa.
47
J.N.A. Hooper et al.
O
O
Fig. 31. Halichondria darwinensis sp. nov. A, larger choanosomal oxea; B, smaller ectosomal oxea; C, peripheral
skeletal structure; ll, preserved paratype QM G303252; E, live intertidal paratype.
48
Beagle Gulf halichondriid sponges
Halichondria darwinensis sp. nov.
(Figs 31-32)
Type material. HOLOTYPE - NTMZ3205:
Dudley Point Reef, East Point, Darwin, NT,
12°24.5'5, 130°48.0'E, intertidal, 25 September 1987, coll. N. Smit, by hand. PARATYPE QM G303252: S of South Shell Island, East
Arm, Darwin Harbour, NT, 12°29.8'S,
130°53.5'E, intertidal, 19 September 1993, coll.
J.N.A. Hooper, L.J. Hobbs and B. Alvarez, by
hand.
Distribution. South Shell Island and East
Point Aquatic Reserve, Darwin Harbour, NT.
Habitat. Fringing coral reef, binding coral
nibble substrate, growing on algae and dead
coral; intertidal.
Description. Encrusting base, up to 15 mm
thick, with stoloniferous, arborescent and bushy,
fibrous, insubstantial digits on upper surface,
with digits standing erect or meandering across
substrate; digits up to 85 mm long, 6 mm diameter. Oscules small, discrete and conspicuous,
up to 2 mm diameter, scattered on upper surface
between upright surface processes.
Colour. Live colouration blue-grey on branch
tips (Munsell 2.5B 8/2) and pale grey on base
(5G 8/2); beige in ethanol.
Texture. Very soft, friable, easily torn.
Surface. Translucent, even, with fibrous and
porous surface and underlying spicule tracts
visible through ectosomal skeleton.
Ectosomal skeleton. Poorly developed, not
detachable, thin tangential layer two to three
spicules wide, composed mainly of smaller oxeas,
overlaying larger choanosomal oxeas in
paratangential subectosomal skeleton.
Choanosomal skeleton. Halichondroid reticulate skeleton of mainly longitudinal, multispicular tracts, with bundles of 5-10 oxeas ar-
Fig. 32. Halichondria darwinensis sp. nov. Paratype QM G303252. A, perpendicular section through skeleton showing
halichondroid unispicular reticulation and directionless multispicular tracts of the choanosomal skeleton; B, transverse section of
ectosomal skeleton showing outer surface of sponge with a thin tangential unispicular layer of predominantly smaller spicules;
C, light micrograph of section through choanosomal skeleton showing disorganised halichondroid multispicular tracts of larger
spicules and thin, vestigial ectosomal skeleton (at right); D, sharply pointed spicule terminations on larger choanosomal oxeas
(left) and abruptly telescoped (mucronate) points of smaller ectosomal oxeas (right).
49
J.N.A . Hooper et al.
moreso than their length, the shorter found
mainly in the ectosomal skeleton, the larger
found mainly in choanosomal tracts), spicule
geometry (with sharply pointed choanosomal
oxeas and prominently telescoped, mucronate
points on ectosomal oxeas), skeletal structure
(mainly longitudinal multispicular tracts of
choanosomal oxeas, with single spicules dispersed between tracts producing a disorganised
halichondroid-reticulate skeleton). In addition
to these features seen in preserved material, the
species also has distinctive field characteristics
including a meandering stoloniferous growth
form suited to growing on coral rubble, binding
the substrate, a soft, easily torn consistency and
a blue-grey live colouration. Relationships between H. darwinensis and other Halichondria
are presently uncertain given that most of the
field characters of this species have not been
ranged paratangentially in subdermal region
supporting surface skeleton, and unispicular
tracts dispersed between, both composed mainly of larger oxeas. Spongin fibres not evident.
Collagen light to moderate in mesohyl, associated predominantly with spicule tracts.
Spicules. Oxeas in two sizes: thicker
choanosomal oxeas straight or slightly curved at
centre, with sharply pointed, tapering fusiform
points (length 5254622.0)-808 pm, width 6-(8.4)12 pm); thinner ectosomal oxeas straight or rarely
slightly curved, tapering fusiform, with prominently telescoped points (mucronate) (length 320(392.2)-485 pm, width 1.5-(2.2)-3 pm).
Etymology. Named for the type locality, Darwin Harbour.
Remarks. Halichondria darwinensis is distinctive in its spicule size (having two categories of oxeas differentiated by their thickness
C
C
I 5-
Fig. 33. Halichondria ridleyi sp. nov. A, larger choanosomal oxea; B, smaller ectosomal oxea; C, peripheral skeletal structure;
D, live intertidal holotype QM G303309; E, preserved holotype.
50
Beagle Gulf halichondriid sponges
well documented in other publications on Indowest Pacific halichondriids.
mm diameter at base. Oscules small, less than 2
mm diameter, scattered between surface processes.
Colour. Slightly mottled khaki dark yellowbrown exterior (Munsell 2.5Y 7/8-6/6) and beige
interior alive; uniform beige-white in ethanol.
Texture. Firm, friable.
Surface. Opaque, hispid, even, unornamented.
Ectosomal skeleton. Thin, hispid, collagenous, non-detachable layer, with larger choanosomal spicules protruding through ectosome,
erect or paratangential to surface, overlayed by
smaller spicules lying tangential to surface, forming sparse tracts or occurring individually.
Choanosomal skeleton. Disorganised halichondroid, directionless tracts of single spicules
throughout skeleton, composed mainly of larger
oxeas, with tracts becoming only vaguely plumo-
Halichondria ridleyi sp. nov.
(Figs 33-34)
Type material. HOLOTYPE - QM G303309:
Dudley Point Reef, East Point, Darwin, NT,
12°25.3'S, 130°49.0'E, intertidal, 20 September 1993, coll. J.N.A. Hooper and L.J. Hobbs,
by hand.
Distribution. East Point Aquatic Reserve,
Darwin Harbour.
Habitat. Dead coral and laterite rock reef;
intertidal.
Description. Partly burrowing, excavating
base, boring into dead coral, with massive erect,
irregular surface processes, buttressed and
fistulose, with fistules up to 54 mm long, 35
Fig. 34. Halichondria ridleyi sp. nov. Holotype QM G303309. A, Perpendicular section through skeleton showing halichondroid
unispicular reticulation of directionless tracts throughout the choanosomal skeleton. B, Transverse section of ectosomal skeleton
showing outer surface of sponge with protruding larger oxeas and tangential smaller oxeas. C, Light micrograph of section
through peripheral skeleton showing directionless halichondroid uni- or paucispicular reticulation in choanosome, and vaguely
plumo-reticulate subectosomal skeleton. D, Sharply pointed spicule tenninations on both larger choanosomal oxeas (left) and
smaller ectosomal oxeas (right).
51
J.N.A. Hooper et al.
Genus Hymeniacidon Bowerbank
reticulate in subectosomal region, forming erect
or paratangential tracts ascending to (and protruding through) surface. Few multispicular
tracts observed, meshes produced by skeletal
reticulation small, ovoid, up to 200 pm diameter. Mesohyl collagen abundant, lightly
pigmented and dispersed evenly throughout
choanosome. Choanocyte chambers small, ovoid,
up to 50 pm diameter.
Spicules. Oxeas in two size classes: larger
choanosomal oxeas moderately or slightly curved
at centre, with tapering fusiform, sharply pointed
ends, occasionally stylote (length 305-(448.5)565 pm, width 9-(16.3)-21 pm); smaller
ectosomal oxeas straight or slightly curved, tapering fusiform, sharply pointed or blunt terminations (length 112-(164.2)-188 pm, width 4(5.9)-8 pm).
Etymology. Named for Stuart 0. Ridley, a
pioneer of sponge taxonomy and author of the
first comprehensive scientific treatise on Australian sponges, including material from Darwin Harbour (Ridley 1884).
Remarks. This species is similar to H. mertoni
(Hentschel) from Aru Island (Hentschel 1912)
in its burrowing, excavating base with robust,
irregularly fistulose digits on the upper surface
(unlike other fistulose species such as H. tyleri
which have pencil-like digits). It differs from H.
mertoni and other Indo-west Pacific Halichondria in having a khaki-brown live colouration,
an even, unsculptured surface, a firm texture, a
non-detachable tangential ectosomal skeleton, a
choanosomal skeleton that is nearly completely
disorganised halichondroid-reticulate throughout, including the peripheral skeleton (whereas
many species of Halichondria have at least some
skeletal organisation near the surface), and two
size classes of oxeas (305-565 x 9-21 pm and
112-188 x 4-8 pm), the smaller distributed in
the ectosomal skeleton. In H. mertoni the
colouration is white, the fistules are prominently
conulose and the outer surface is shaggy, the
choanosomal skeleton has differentiated halichondroid central choanosomal skeleton and
radial subectosomal skeleton, and there is only
a single size class of (choanosomal) oxea (640785 x 18-22 pm). Hentschel (1912) reported
that there was no ectosomal skeleton in H.
mertoni (in which case it would not belong to
Halichondria at all), but one of the syntypes
(SMF1608) does have a tangential ectosome
although spicule sizes are undifferentiated.
Hymeniacidon Bowerbank, 1863: 1112.
Amorphilla Thiele, 1898: 44.
Stylohalina Kirk, 1909: 539.
Thieleia Burton, 1932a: 329.
Uritaia Burton, 1932b: 198.
Rhaphoxiella Burton, 1934: 554.
Rhaphidostyla Burton, 1935: 651.
Nailondra de Laubenfels, 1954: 182.
Type species. Hymeniacidon caruncula Bowerbank, 1864, by original designation (junior
synonym of Spongia perlevis Montagu, 1818).
Diagnosis. Massive, encrusting or fistulose
growth forms. Choanosomal skeleton poorly defined directionless spicule tracts with vaguely
reticulate multispicular bundles and halichondroid criss-cross of single spicules between major tracts. Cavities (lacunae) often occurring in
choanosomal and subetosomal regions, with
vaguely ascending or oblique spicule tracts supporting ectosomal crust. Ectosomal skeleton thin
tangential crust of single spicules or spicule
bundles, usually detachable from surface.
Megascleres exclusively styles and stylotes, usually of a single category.
Remarks. According to recent revisions (Van
Soest et al. 1990; Diaz et al. 1991), Hynzeniacidon is very similar to (and a sister taxon of)
Halichondria. The two genera differ only substantially in having styles instead of oxeas, respectively (i.e. with the assumption that oxeas
have been secondarily lost in Hymeniacidon).
Hymeniacidon also has a characteristically more
fleshy choanosomal structure, and its tangential
ectosomal crust may be more poorly developed
than in Halichondria, but there are few other
significant differences between these genera.
There is some doubt about the proposed inclusion of Stylotella into synonymy with Hymeniacidon (Van Soest et al. 1990: 51). This doubt
was implied by Hooper and Wiedenmayer (1994)
in their recognition of Stylotella as a distinct
genus from Hymeniacidon. This interpretation
was based on characters taken from a single
syntype of the type species, Stylotella digitata
Lendenfeld, 1888 (AM G9152) from Port
Jackson, NSW, which has a predominantly
plumose subectosomal skeleton (best developed
towards the periphery of the skeleton), a more
disorganised halichondroid reticulate skeleton
in the central portion of the choanosome, and
lacks a tangential skeleton completely (having
52
Beagle Gulf halichondriid sponges
instead plumose bundles of styles protruding
through the surface), unlike typical Hymeniacidon. These characters essentially agree with the
redescription of S. digitata by Hallmann (1914:
349), although it is unclear which material his
description was based. However, a redescription
of a syntype of Hymeniacidon agminata Ridley,
1884 (BMNH 1881.10.21.347) also from Port
Jackson, NSW, by Van Soest et al. (1990) (allegedly a senior synonym of S. digitata;
Hallmann 1914), conflicts with this diagnosis,
particularly in the fact that it does have "horizontally disposed spicules" in the ectosomal
skeleton (Van Soest's et al. 1990: 55) and thus
belongs to Hymeniacidon. (Van Soest et al.'s
(1990) use of the term `holotype' for this specimen is implicitly a lectotype designation). It is
possible that H. agminata is not a synonym of S.
digitata, contrary to the opinion of Hallmann
(1914), but this must be ascertained by careful
re-examination of the 25 or so type specimens
of S. digitata and its alleged synonyms (see
Hooper and Wiedenmayer 1994: 212).
Species of Hymeniacidon are widely distributed in both tropical and temperate waters of all
oceans, including polar seas (see Diaz et al.
1991), recorded from shallow subtidal waters to
depths exceeding 3500 m. Within the IndoAustralian region nine species have been recorded so far: H. agminata Ridley from eastern
Australia (discussed above), H. centrotyla
Hentschel,H. fernandezi Thiele, H. kerguelensis
Hentschel and H. torquata Topsent from Macquarie Island, Subantarctic and Antarctic waters (Thiele 1905; Hentschel 1914; Koltun 1964,
1976), H. halichondroides (Thiele) from the
south coast of NSW and Tasmania (Carpay
1986), H. rigida Dendy from Bass Strait, Vic.
(Dendy 1897; Hallmann 1914), H. gracilis
(Hentschel) and H. suberitoides (Brondsted)
from the Java and Arafura Seas (Hentschel 1912;
Brondsted 1934).
Fig. 35. Hymeniacidon vernonensis sp. nov. Holotype NTM Z4085. A, perpendicular section through skeleton showing
disorganised unispicular halichondroid reticulation and cavernous, highly collagenous skeleton; B, transverse section of ectosomal
skeleton showing the thin layer of collagen with scattered single spicules in halichondroid arrangement tangential to the surface;
C, slightly telescoped points (left) and hastate base of smaller style (right); D, prominently telescoped point (left) and evenly
rounded base of larger style (right).
53
J.N.A. Hooper et al.
NI III EV
11111
Fig. 36. Hymeniacidon vernonensis sp. nov. A, thicker style (predominantly in choanosomal skeleton); B, thinner style
(predominantly in ectosomal skeleton); C, peripheral skeletal structure; D, preserved paratype QM G303570; E, live specimen
(on deck) QM G303565.
Hymeniacidon vernonensis sp. nov.
Other Material. QM G303565: SSE of Lyne
Reef, Vernon Islands, NT, 12°07'S, 130°59'E,
30m depth, 11 October 1993, coll. CCNT stn.
139 (1959), dredge.
Distribution. Known only from the Vernon
Islands, Van Diemen Gulf, NT.
Habitat. Shell and rock substrate; subtidal,
6-30 m depth.
Description. Flat, plate-like, fan-shaped
growth form, encrusting, following contours of
substrate, with prominently shaggy and conulose
(Figs 35-36)
Type material. HOLOTYPE - NTM Z4085
(fragment QM G303617): S of Northwest
Vernon Island, NT, 12°03.9'S, 131°01.9'E, 25
m depth, 10 October 1993, coll. CCNT stn. 148
(1716). PARATYPE - QM G303570: SSE of
Lyne Reef, Vernon Islands, NT, 12°07'S,
130°59'E, 30 m depth, 11 October 1993, coll.
CCNT stn. 139 (1976), dredge.
54
Beagle Gulf halichondriid sponges
surface, up to 90 mm long, 15 mm thick. Oscules
small, up to 2 min diameter, conspicuous, discrete, scattered over upper surface, with thin
membraneous lip.
Colour. Yellow, yellow-brown, or orangebrown, alive in air) (Munsell 7.5YR 7/8 - 2.5Y
8/8))„ pale grey in ethanol.
TOxture. Soft, compressible, easily torn.
Surface. Transparent outer surface, hispid,
uneven, shaggy, conulose, with fibrous-membraneous appearance.
Ectosomal skeleton. Detachable, thin, membranous layer of granular pigmented collagen,
with sparse layer of mostly thinner styles lying
tangential to surface, usually in unispicular tracts
and in disorganised halichondroid arrangement,
and larger choanosomal styles protruding
slightly through surface.
Choanosomal skeleton. Deeper regions of
skeleton disorganised halichondroid reticulate,
with single spicules or occasionally bundles of
two or three, usually not forming tracts, consisting of mostly larger styles but also with thinner
spicules dispersed between unispicular tracts.
Subectosomal skeleton marginally more organised towards periphery, with choanosomal spicules generally ascending to surface, producing
slightly plumose paucispicular tracts corresponding to position of surface conules. Spongin fibres absent, but collagen abundant in choanosomal mesohyl, containing large pigment granules. Choanosomal region irregularly cavernous, with large lacunae up to 1 mm diameter.
Spicules. Styles is two size classes: thicker
styles (mostly found in choanosomal tracts)
curved at centre, sinuous or rhabdose, fusiform,
with evenly rounded bases,. occasionally
anisoxeote or mucronate, and prominently tetescoped points or less often sharply pointed end
(length 4954584.4)-6615 pm,. width 12416‘.6).
22 pm); thinner styles (mostly fbund in tangem-tial ectosomal skeleton) slightly curved at centre, with sharp pointed or slightly telescoped
p6,ints, and tapering hastate rounded bases.
(length 252-(422.0)-565 pm, width 546.4)-8'
pm).
Etymology. ;`lamed for the type locality,
Vernon Islands.
were initially
Remarksmisidentified as Stylotella j1L-Ll.)elliformis
Hentschel, 19 111?-sitite both taxa have a cr9 mi
-netlyshagWik,fbterowhm
essentially styloterriegascleres forming disorganised, halichondroid reticulate skeletal struc-
tures only slightly more organised near the surface forming vaguely ascending spicule tracts,
than in the centre of the skeleton. However, H.
cernonensis has a tangential ectosomal skeleton
composed of generally thinner styles than those
inside the choanosomal skeleton (whereas the
ectosome in S. flabellifOrmis is simply fleshy,
membraneous, without tangential spicules but
with protruding, vaguely erect choanosomal spicules piercing the surface). Its megascleres are
also exclusively styles, many with telescoped
points (whereas spicules of S. flabellifOrmis are
either sharply pointed or modified to strongylote
forms, never telescoped); the choanosome is
fleshy, halichondroid-reticulate, with distinct
horizontal and vertical spicule tracts throughout the skeleton (whereas in S. fiabelliformis
spicule tracts are essentially plumo-reticulate,
vaguely ascending to the surface); and growth
form is flattened fan-shaped, appearing to grow
flat on the substrate (as opposed to erect,
flabellate typical for S. fiabellifortids).
Hooper and Levi (1993b) recently redescribed
S. flabelliformis from its Indonesian holotype as
well as many other specimens from a number of
localities throughout the Indo-west Pacific. They
included it in the genus Stylissa, within the
family Axinellidae, although both its generic
and family assignments are still debatable, whereas
the present species clearly belongs to Hymeniacidon in its spicule and skeletal characters.
Hymeniacidon gracilis (Hentschel)
(Figs 37-38)
Stylotella digitata var. gracilis Hentschel,
1912: 356-357, pl. 19, fig. 27.
Type material. HOLOTYPE - SMF 970: Between Meriri. L and Leer I., Aru Is, Arafura Sea,
6- ha mi depth,. 31 March 1908.
Other Material. NTM Z945: East Point
`bommies% Darwin, NT, 12°24.05'5, 130°48.0'E,
m depth, 13 September 1982, coll. J.N.A.
Hooper, SCUBA. NTM Z987: same locality, 8
in depth, 26 October 1982. NTM, Z8'83:. Dudley
Point Reef, East Point, Darwin, NT,. 12°25.0'S,
30°48.04'E, 10 m depth, 31 Alugust.1982, coll.
J.N.A. Hooper, SCUBA.
Distribution. Aru Islands, Arafura Sea; Darwin Harbour.
Habitat. Rocky reef, sandy substrate; intertidal-subtidal depths.
1;e5cription. Thickly encrusting with erect,
conulose iamellae forming thickly branching,
55
J.N.A. Hooper et al.
I=
imm
ma
I= Ell II
Fig. 37. Hymeniacidon gracilis (Hentschel). A, choanosomal styles; B ,. peripheral skeletal structure; C, specimen NTM Z945.
irregularly anastomosing, subspherical mass, up
to 95 mm long, 55 mm diameter. Oscules small,
less than 2 mm diameter, scattered over surface
of lamellae.
Colour. Pale orange-red in life, (Munsell lOR
7/8), pink-beige in ethanol.
Texture. Soft, compressible, slimy.
Surface. Opaque, collagenous; uneven and
papiliose-conulose.
Ectosomal skeleton. Dense collagenous
ectosomal skeleton, with single spicules or tracts
of styles, 5-10 spicules abreast, aligned tangen56
Beagle Gulf halichondriid sponges
Fig. 38. Hyrneniacielon gracilis (Hentschel). Specimen NTM Z945. A, perpendicular section through skeleton showing
halichondroid reticulate choanosomal structure; B, transverse section of ectosomal skeleton showing outer surface of sponge with
paratangential-plurnose bundles of styles from the subectosomal skeleton and irregularly dispersed single or paucispicular tracts
of styles lying tangential to the surface; C, light micrograph of section through peripheral skeleton showing plumo-reticulate
development of the outer portion of subectosomal region; D, fusiform point (left) and evenly rounded base of style (right).
tial to surface; erect paucispicular plumose bundles of styles protruding through tangential layer,
arising from deeper choanosomal skeleton.
Choanosomal skeleton. Interior of choanosome consists of single spicules or
paucispicular bundles of two or three spicules
abreast, forming partially organised, mostly
halichondroid reticulate structure; spicule tracts
becoming more organised towards surface,
with distinct ascending multispicular plumose
tracts in peripheral skeleton; paucispicular bundles protruding slightly through surface but
only partially beyond collagenous layer. Collagen in choanosome less dense than ectosomal
region and mostly associated with spicule bundles. In some specimens collagen appearing
granular.
Spicules. Styles of single size class, straight
or slightly curved, fusiform, sharply pointed,
with evenly rounded bases (length 218-(251.2)285 pm, width 2-(5.1)-8 pm).
Remarks. This species was originally described by Hentschel (1912) as a new variety
(subspecies) of Stylotella digitata Lendenfeld,
but there is very little resemblance in skeletal
structure or growth form between these taxa
and the Arafura Sea population is elevated here
to full species rank. Spicule geometry, spicule
dimensions and skeletal structure are comparable between southeast Indonesian and northwest Australian populations, whereas the
holotype is substantially thinner than present
material, with a concomitant reduction in the
extent of development in the (organised portion) of the peripheral skeleton. The possession
of a tangential ectosomal skeleton and a halichondroid-reticulate choanosomal skeleton, the
latter only showing significant organisation near
57
J.N.A. Hooper et al.
tember 1981, coll. J.N.A. Hooper and P.N.
Alderslade, by hand. PARATYPE - QM
G303414: N of South Shell I., Darwin Harbour,
NT, 12°29.8'S, 130°52.9'E, 14 m depth, 25
September 1993, coll. J.N.A. Hooper and L.J.
Hobbs, SCUBA.
Other Material. NTM Z272: Dudley Point
Reef, East Point, Darwin, NT, 12°25.0'S,
130°49.01'E, 17 September 1981, coll. J.N.A.
the periphery, indicates that the species belongs
in Hymeniacidon.
Hymeniacidon hapalia sp. nov.
(Figs 39-40)
Type material. HOLOTYPE - NTM Z184:
Dudley Point Reef, East Point, Darwin, NT,
12"25.0'S, 130°49.01'E, 0.5 m depth, 13 Sep-
a
Fig. 39. Hymeniacidon hapalia sp. nov. A, larger styles; B, smaller strongylote styles; C, peripheral skeletal structure;
D, paratype QM G303414; E, live intertidal specimen QM G303280; F, live subtidal holotype NTM Z 184.
58
Beagle Gulf halichonclri id sponges
Hooper, by hand. NTM Z286, Z291: same locality, lm depth, 18 September 1981, coll. J.N.A.
Hooper, by hand. NTM Z2146: same locality, 27
September 1984, coll. J.N.A. Hooper, by hand.
NTM Z2560: Fannie Bay reef, Darwin, NT,
12°24.50'S, 130°48.0'E, intertidal, 12 December
1985, coll. A.M. Mussig and C. Hood, by hand.
NTM Z775: Channel Island, Darwin Harbour,
NT, 12°33.0'S, 130°52.03'E, intertidal, 24 May
1982, coll. P.N. Alderslade, by hand. QM
G303280: S of South Shell Island, East Arm,
Darwin Harbour, NT, 12°29.8'S, 130°53.5'E,
intertidal, 19 September 1993. coll. J.N.A.
Hooper, L.J. Hobbs and B. Alvarez, by hand.
Distribution. East Point Aquatic Reserve,
Fannie Bay Reef, Lee Point, Channel Island and
South Shell Island, Darwin Harbour.
Habitat. Rocky reef, mudflats, fringing coral
reef and coral pinnacles; intertidal-subtidal
depths.
Description. Thickly encrusting, massive-bulbous, stoloniferous or elongate ridges, up to 30
mm thick, closely following contours of
substrate. Oscules relatively large, conspicuous,
discrete or multiple (sieve-plate), up to 5 mm
diameter, usually on apex of surface ridges,
between conules, surrounded by slightly raised
membraneous lip.
Colour. Bright orange to dark orange alive
(Munsell IOR 6/10 - 5YR 7/8), pale orangebrown in ethanol.
Texture. Soft, insubstantial, membraneous,
easily torn, fragile (often disintegrating upon
collection), producing clear mucus.
Surface. Long, sharply pointed or bifurcated,
bushy, digitate and conulose surface processes
scattered over surface, interconnected by fibrous
ridges; surface opaque, prominently membranous, collagenous.
Ectosomal skeleton. Collagenous, membraneous, detachable ectosomal peel composed
of granular, heavily pigmented collagen together
with sparse, tangential or paratangential tracts
or individual spicules, composed of both larger
Fig. 40. Hymeniacidon hapalia sp. nov. Holotype NTM Z184. A, perpendicular section through skeleton showing irregular
paucispicular reticulate fibres in subectosomal skeleton supporting collagenous ectosome; B, transverse section of ectosomal
skeleton showing the aspicular collagenous outer surface; C, rounded, slightly asymmetrical ends of smaller strongylote styles;
D, slightly telescoped point (left) and evenly rounded base of larger style (right).
59
J.N.A. Hooperet al.
ertheless detachable, thus conforming to the
strict definition of the genus Hymeniacidon. It
was absent in some specimens, such as poorly
fixed frozen samples, where the ectosome had
presumably become detached from the remainder of the sponge prior to histology.
Hymeniacidon hapalia is also clearly unusual amongst other species of the genus in having smaller spicules of different morphology
than the main structural megascleres. These
strongylote spicules are clearly derived from
styles as they have asymmetrical strongylote
ends. They are not localised to any particular
region of the sponge but are dispersed throughout the ectosomal and choanosomal regions,
albiet sparsely.
Hymeniacidon hapalia should be compared
to Batzella corticata (Thiele) and Nailondra
maza de Laubenfels, both of which have styles
together with modified strongylote styles (see
brief redescriptions in Van Soest et al. 1990),
but in both these species all spicules are approximately of equivalent length. It is possible
that one of the genera synonymised under Hymeniacidon by Van Soest et al. (1990) should
be resurrected for H. hapalia, such as
Rhaphoxiella Burton (i.e. based on possession
of modified stylote spicules as well as styles), or
Uritaia Burton (with differentiated larger and
smaller styles), or Thieleia Burton (with a predominantly collagenous ectosomal peel) - differences which Van Soest et al. (1980) considers to differ from Hymeniacidon s.s. only by
degree - but such action is not warranted without a more in-depth study of the family, and for
the present H. hapalia fits with a broad definition of Hymeniacidon.
The present species is also similar to
Stongylacidon intermedia Burton, 1934, from
the Great Barrier Reef (which de Laubenfels
(1954) suggested belonged to Ulosa, Hooper
and Wiedenmayer (1994) subsequently referred
to Batzella, but is still poorly known).
Stongylacidon intermedia is so far known only
from the holotype, and has not yet been described from the living population. Nevertheless, its distinctive external appearance (massive, subspherical, with prominent, short digitate processes), skeletal structure (irregularly
reticulate, branching, anastomosing tracts of
megascleres surrounded by heavy collagen) and
spicule geometry (strongyles and styles) are similar to those in H. hapalia, although spicule sizes
are considerably different between the two spe-
styles and smaller strongylote styles; choanosomal fibres and spicules also protruding through
surface only at surface conules.
Choanosomal skeleton. Deeper regions of
choanosome predominantly collagenous, very
granular; larger styles scattered individually
throughout mesohyl and also forming paucispicular, irregular, directionless, halichondroid
tracts, with both horizontal and vertically reticulate elements, bound together with abundant
collagen. Subectosomal skeleton more obviously reticulate than central skeleton with pauciand multispicular tracts easily visible. Collagen
dominant over spicules throughout skeleton,
particularly in periphery. Smaller strongylote
styles also scattered within mesohyl, not confined to any particular region of skeleton, but
generally not forming spicule tracts. Spicule
meshes cavernous, up to 600 pm diameter in
places.
Spicules. Spicules in two categories: larger
styles straight or slightly curved at centre, fusiform, sharply pointed or with slightly telescoped
points, with evenly rounded bases, some slightly
rhabdose (length 375-(583.5)-1130 pm, width
3-(8.1)-15 pm); smaller strongylote styles
straight, usually asymmetrical, with evenly
rounded ends (length 182-(231.9)-275 pm, width
1-(2.1)-2.5 pm).
Etymology. Named for the soft, delicate, collagenous consistency, from hapalus (L.), soft to
touch.
Remarks. This species is distinctive in the
field, with dark orange colouration, prominently
conulose fleshy surface, and very soft, relatively
fragile, collagenous texture. It is predominantly
intertidal, being a prominent member of the
coral rubble community fringing the shores of
Darwin Harbour, and when it is exposed to air
during spring low tides it literally drips mucus
(like many other tropical intertidal sponges and
soft corals in this region), presumably with a
function to protect the species from dessication
during prolonged exposure to air. Although the
species appears to be fragile to touch and collect, individuals exposed to air for many hours
do not appear to undergo tissue necrosis or any
other visible effects, unlike the widespread
necrosis seen in many other subtidal species.
Hymeniacidon hapalia is unusual to the genus in having a predominantly collagenous
ectosomal peel with only sparse tangential spicules or spicule tracts supporting the ectosomal
peel. This paper-like ectosomal skeleton is nev-
60
Beagle Gulf halichondriid sponges
Berg, C. 1899. Substitution de nombres genericos.
III. Communicationes de Museo Nacional Buenos Aires 1: 77-80.
Bergquist, P.R. 1961. Demospongiae (Porifera) of
the Chatham Islands and Chatham Rise, collected by the Chatham Islands 1954 Expedition. New Zealand Department of Scientific and
Industrial Research Bulletin 139. Biological
Results of the Chatham Islands 1954 Expedition (5): 169-206.
Bergquist, P.R. 1965. The sponges of Micronesia,
Part I. The Palau Archipelago. Pacific Science
19: 123-204.
Bergquist, P.R. 1970. The Marine fauna of New
Zealand: Porifera, Demospongiae, Part 2.
(Axinellida and Halichondrida). New Zealand
Department of Scientific and Industrial Research Bulletin. New Zealand Oceanographic
Institute Memoir 197:1-85, pls. 1-20.
Bergquist, P.R. 1978. Sponges. Hutchinson: London.
Bergquist, P.R. and Tizard, C.A. 1967. Australian
intertidal sponges from the Darwin area.
Micronesica 3: 175-202.
Bowerbank, J.S. 1863. On the anatomy and physiology of the Spongiaidae. Part III. One the generic characters, the specific characters, and on
the method of examination. Philosophical
Transactions of the Royal Society 152 (1862):
1087-1135, pls 72-74.
Bowerbank, J.S. 1864. A Monograph of the British
Spongiadae. Ray Society: London. Volume 1.
Pp 1-290, pls 1-37.
Bowerbank, J.S. 1873. Contributions to a general
history of the Spongiadae. part IV. Proceedings
of the Zoological Society of London 1873: 325, pls 1-4.
Brondsted, H.V. 1923. Sponges from New Zealand.
Part I. In: Mortensen, T. (ed.) Papers Dr. Th.
Mortensen's Pacific Expedition 1914-16. XV
Videnskabelige Meddelelser fra Dansk
Naturhistorisk Forening 77: 435-483.
Brondsted, H.V. 1924. Sponges from the Auckland
and Cambell Islands. In: Mortensen, T. (ed.)
Papers from Dr. Th. Mortensen's Pacific Expedition 1914-16. XV Videnskabelige Meddelelser
fra Dansk Naturhistorisk Forening 75: 117167.
Brondsted, H.V. 1934. Resultats Scientifiques du
voyage aux Indies Orientales Neerlandaises.
Sponges. Memoires de Brusseles Institut Royal
des Sciences Naturelles de Belgique 2(15): 326.
Burton, M. 1926. Observations on some British
species of sponges belonging to the genus
Reniera. Annals and Magazine of Natural History 22: 71-83.
Burton, M. 1927. III. Report on the sponges. In:
Zoological results of the Suez Canal Expedi-
cies, 375-1130 x 3-15 pm and 182-275 x 1-2.5
pm for styles and strongyles, respectively, in H.
hapalia; versus 140 x 3 pm for both strongyles
and styles in S. intermedia. This latter species is
incertae sedis until living populations are rediscovered.
ACKNOWLEDGEMENTS
We thank Sandy Bruce and Phil Alderslade
(Northern Territory Museum of Arts and Sciences) for facilitating our access to the NT Museum sponge collections, and various technical
staff who have assisted the senior author during
ten years of research on the sponge fauna in
northwest Australia; Ron Billyard (Parks and
Wildlife Commission of the Northern Territory,
Ocean Rescue 2000 Program) for trawled specimens from the Beagle Gulf Marine Park; Neil
Smit (Northern Territory University) for assisting in various field surveys; Daryl Grey, Dave
Ramm and Anne Coleman (Northern Territory
Fisheries) for collection permits, logistic field
support and generous access to the fisheries
research vessel over the years; Peter Murphy
and colleagues (NCI AIMS) for providing samples from Parry Shoals; and Penny Barents (AM,
Sydney), Shirley Stone (BMNH, London),
Claude Levi (MNHN, Paris), Manfred Grasshoff
(SMF, Frankfurt) and Dieter KOhlmann (ZMB,
Berlin) for providing access to extensive type
collections. We especially thank Leonie G.
Hooper for illustrating spicules and skeletons of
halichondriids, funded in part by ABRS grant
to the senior author (87/5996). This paper
benefitted greatly from the comments of two
referees.
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Accepted 15 August, 1995
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