The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2011 27: 55–84
Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) of
northern Australia. Family Halichondriidae
BELINDA ALVAREZ1 and JOHN N. A. HOOPER2
1
Museum and Art Gallery of the Northern Territory. GPO Box 4646, Darwin NT 0801, AUSTRALIA
belinda.glasby@nt.gov.au
2
Queensland Museum. PO Box 3300, South Brisbane QLD 4101, AUSTRALIA
john.hooper@qm.qld.gov.au
ABSTRACT
Fifteen species in six genera of the family Halichondriidae, including two new species, Halichondria (Halichondria)
carotenoidea sp. nov. and Halichondria (Halichondria) microbiana sp. nov., are recorded for northern Australia as
part of a revision of the order Halichondrida (Porifera: Demospongiae) in this region. Descriptions and discussion
of those species are presented here. Eight new combinations within the family Halichondriidae are here established,
i.e. Amorphinopsis fenestrata (Ridley, 1884, as Leucophloeus), Amorphinopsis maculosa (Pulitzer-Finali, 1996, as
Topsentia), Axinyssa bergquistae (Hooper et al., 1997, as Halichondria), Axinyssa mertoni (Hentschel, 1912, as
Ciocalypta), Axinyssa gracilis (Hentschel, 1912, as Ciocalypta rutila gracilis), Axinyssa terpnis (De Laubenfels, 1954,
as Phycopsis), Ciocalypta vansoesti (Hooper et al., 1997, as Halichondria) and Topsentia ridleyi (Hooper et al., 1997
as Halichondria) and one species is relocated into the family Dictyonellidae, i.e, Stylissa vernonensis (Hooper et al.,
1997, as Hymeniacidon). A lectotype is designated for Ciocalypta stalagmites Hentschel, 1912.
Keywords: sponge, Porifera, Halichondrida, Halichondriidae, northern Australia, new species, taxonomy.
INTRODUCTION
The family Halichondriidae was revised by Erpenbeck
& Van Soest (2002). Its deinition is based entirely on
a few skeletal characters, i.e. presence of an ectosome
with specialised skeleton, a disorganised choanosomal
skeleton, dimensions of oxeas and styles, absence of
microscleres. Other morphological characters (e.g.
general shape of megascleres, presence or size categories
among the megascleres, spicule density and its relation to
consistency, orientation of spicules at the ectosomal level)
are used in combination with the diagnostic characters to
separate species and genera within this family. All these
characteristics are very simplistic, and often displayed as a
gradient of variation which makes separation of species very
subjective as relected by the large number of synonyms
(see Van Soest et al. 2008).
The family includes 14 genera and at least 296 valid
species (Van Soest et al. 2008); however, the status and
generic allocation of many of the species listed on this
database on the Internet needs to be revised and validated
against the current concept of the genera.
The Australian Faunal Directory (Hooper 2005)
currently lists a total of 42 species of Halichondriidae.
Twenty species were reported and described for the Beagle
Gulf (northern Australia) by Hooper et al. (1997). No
other revision of species of Halichondriidae in Australia or
adjacent areas is presently available.
The present paper represents the third part of a revision
of the order Halichondrida in the northern Australian region
and includes the family Halichondriidae. Alvarez & Hooper
(2009, 2010) provided details and presented an introduction
to the revision of the order and the families Axinellidae and
Dictyonellidae.
MATERIALS AND METHODS
55
This revision includes material of the family
Halichondriidae recorded for the tropical northern
Australian waters of the Northern Territory and Queensland
coast (from Admiralty Gulf in the west to Torres Strait in
the east, approx. between the 125º E and 142º E).
Complete locality and collection data for non-type
voucher material deposited at the Queensland Museum and
the Museum and Art Gallery of the Northern Territory are
available in Appendix 1.
The distribution of species is given according to
the marine provinces deined by Spalding et al. (2007).
Spicule measurements are in micrometres and are based
on 25 spicules (unless indicated in square brackets) and
denoted as range (and mean ± 1 S.E.) of spicule length and
width. All other methods as discussed in Alvarez & Hooper
(2009, 2010).
B. Alvarez and J. N. A. Hooper
A
B
C
D
E
G
F
H
I
Fig. 1. A, Amorphinopsis excavans, specimen in situ at South Shell I., Darwin Harbour, N.T; B, Amorphinopsis fenestrata, specimen in situ
at Weed Reef, Darwin Harbour, N.T. C, Amorphinopsis maculosa: Specimen (QM G313577) dredged at Groote Eylandt, Gulf of Carpentaria,
Queensland; D, Axinyssa bergquistae, specimen (NTM Z. 5976) in situ at Stevens Rock, Darwin Harbour, N.T.; E, Axinyssa mertoni,
specimen in situ at South Shell I., Darwin Harbour, N.T.; F, Ciocalypta heterostyla, specimen in situ at Channel Rock, Darwin Harbour, N.T.;
G,H, C. stalagmites, specimens with different colouration in situ at Channel Rock and Stevens Rock, respectively; I, C. vansoesti, specimen
in situ at Channel Rock, Darwin, N.T. Photographs: A,B – M. Browne; D,G–I – H. Nuguyen, H,E,F – B. Alvarez.
56
Halichondriidae from northern Australia
ABBREVIATIONS
Abbreviations used in the manuscript are: AIMS,
Australian Institute of Marine Sciences, Townsville;
BMNH, Natural History Museum, London (formerly British
Museum (Natural History)); CRRF, Coral Reef Research
Foundation, Palau; GBR, Great Barrier Reef, MAGNT,
Museum and Art Gallery Northern Territory; MNHN,
Musée National d’Historie Naturelle, Paris, France; MSNG,
Museo Civico di Storia Naturale ‘Giacomo Doria’, Genoa,
Italy; MAGNT/NTM, Museum and Art Gallery Northern
Territory (formerly Northern Territory Museum), Darwin,
Northern Territory, Australia; NHMB – Naturhistorisches
Museum, Basel, Switzerland; SMF, Senckenberg Research
Institute and Natural History Museum, Frankfurt; QLD,
Queensland, Australia; QM, Queensland Museum,
Brisbane; WA, Western Australia; ZMA, Zoologisch
Museum, University of Amsterdam, ZMB, Museum für
Naturkunde and der Universität Humboldt zu Berlin, Berlin,
Germany.
Numbers preixed with Q666C, 0CDN, 0M9H are the
cross-reference sample number collected for the United
States National Cancer Institute, under the ‘Collection of
shallow-water organisms’ programme, by the Australian
Institute of Marine Sciences, CRRF and MAGNT
(subcontracted through CRRF) respectively.
TAXONOMY
Family Halichondriidae Gray, 1867
Genus Amorphinopsis Carter, 1887
Gender feminine. Type species, by monotypy,
Amorphinopsis excavans Carter, 1887. Recent, Indian
Ocean.
Amorphinopsis excavans Carter, 1887
(Figs 1A, 2)
Amorphinopsis excavans Carter, 1887: 77; Hooper &
Wiedenmayer 1994: 205; Hooper et al. 1997: 25; Erpenbeck
& Van Soest 2002: 791; Lim, de Voogd & Tan 2008: 115.
Amorphinopsis sacciformis.– Hooper et al. 1997: 27.
Material examined. Darwin Harbour: NTM Z.2215,
Z.4093, Z.4125 (0CDN -8016-W), Z.5213 (0M9H2184-Q);
Z.5222 (0M9H2251-O), Z.5736. Vernon Is: G303658.
Description
Shape (Fig. 1A). Thinly to thickly encrusting (up to
50 mm thick), massive to lobate, or developing short
projections and small lumps, generally growing in patches
and following substrate, semi-buried in substrate.
Colour. Olive green, yellow inside.
Oscula. Round to ovate, inconspicuous, 10 mm diameter.
Surface. Hispid, bumpy.
Skeleton. Ectosomal skeleton (Fig. 2A) thin, detachable
tangential layer, composed by a disorganised criss-cross
reticulation of paucispicular-multispicular tracts of oxeas,
57
up to 100 µm thick, with small styles tangentially to
paratangentially oriented, sometimes in disorganised tufts.
Choanosomal skeleton (Fig. 2B) halichondroid, with large
oxeas oriented in all directions, sometimes grouped in
directionless multispicular tracts; slightly cavernous at
subectosomal area with short multispicular tracts supporting
the ectosomal skeleton.
Spicules (Fig 2C). Oxeas, hastate, in a large range of
sizes, 213.4–945.3 µm (598±221) x 5.9–25.1 µm (16.3±5.5).
Smaller ectosomal styles 140.8–264.9 µm (193.8±41.3) x
4.1–7.5 µm (5.5±0.9).
Remarks. The material examined here agrees with
the description of the syntype by Erpenbeck & Van Soest
(2002: 790).
The specimen described under Amorphinopsis sacciformis
by Hooper et al. (1997) is better allocated to A. excavans. It
does not agree with the syntype of Ciocalypta sacciformis
Thiele which, as mentioned below (see under remarks of
A. carpentariensis), should be interpreted as Halichondria.
The material described by Hooper et al. (1997) is a thin crust
covering a bivalve shell with skeleton similar to the rest of
the material assigned here to A. excavans.
Species considered synonyms of Amorphinopsis
excavans by Hooper & Wiedenmayer 1994, following
Burton (1959), were excluded by Hooper et al. (1997) and
Erpenbeck & Van Soest (2002).
Distribution. The species was recorded originally from
the Mergui Archipelago (Andaman province). Records from
Singapore and northern Australia extend the distribution
of the species to the Sunda and Sahul Shelf provinces. The
northern Australian and Singaporean populations seem to
be common at the intertidal region associated with piers
and wharfs. The species is found also subtidally between
9 and 16 m.
Amorphinopsis fenestrata (Ridley, 1884) comb. nov.
(Figs 1B, 3)
Leucophloeus fenestratus Ridley, 1884: 464; Dendy
1922: 124; Burton 1928: 127.
Leucophloeus fenestratus unnamed variety. – Ridley
1884: 464.
Suberites oculatus Kieschnick, 1896: 534.
Hymeniacidon fenestratus. – Lindgren 1897: 483;
Lindgren 1898: 312 [?].
Ciocalypta oculata. – Thiele 1900: 75.
Ciocalypta oculata maxima Hentschel, 1912: 428.
Axinyssa fenestratus.–Van Soest, Díaz & Pomponi 1990:
27; Hooper & Bergquist 1992: 102.
Ciocalypta confossa Hooper et al., 1997: 23.
Ciocalypta fenestrata.– Hooper et al. 1997: 17.
Ciocalypta oscitans Hooper et al., 1997: 20.
Amorphinopsis foetida. – Hooper et al. 1997: 28.
Material examined. Type maTerial – Leucophloeus
fenestratus, HoloType, BMNH 1882.2.23.255, Darwin
Harbour, N.T., 16–24 m depth, October 1881, HMS
Alert. Leucophloeus fenestratus unnamed variety, BMNH
B. Alvarez and J. N. A. Hooper
C
A
B
Fig. 2. Amorphinopsis excavans (NTM Z.5213): A, light microphotograph of tangential section of ectosomal skeleton, showing oxeas organised
in bundles and small styles, paratangentially oriented in disorganised brushes; B, light microphotograph of section perpendicular to surface,
showing choanosomal skeleton; C, diagram of spicules. Scale bars: A,C, 100 µm; B, 500 µm.
1882.2.23.195, Arafura Sea, 64–72 m depth, 18 October
1881, coll. HMS Alert. Suberites oculatus, syntype SMF
680, Ternate, Maluku Sea, coll. Kükenthal, W, 1894.
Ciocalypta oculata maxima, SMF 971, Aru Is, between
Meriri and Leer, 6–10 m depth, 31 March 1908, coll. Merton,
H. Ciocalypta oscitans, HoloType, QM G303560, Bynoe
Harbour, E Fish Reef, N.T., 12°24.1334’ S, 130°28.16’ E,
17 m depth, 6 October 1993, coll. CCNT Ocean Rescue
2000 Program, dredge. Ciocalypta confossa, NTM Z.3106,
Parry Shoals, Arafura Sea, N.T., 11°12.5167’ S, 129°42.07’
E., 20 m depth, 15 August 1987, coll. Mussig, AM and NCI
team. addiTional specimens – Melville Is: QM G313543.
Shoal Bay, Vernon Is, Cape Hotham: QM G303558,
G303677, G303541. Bynoe Harbour: NTM Z.5207
(0M9H2319-N), Z.5208 (0M9H2543-H). Darwin Harbour:
QM G303287, G310170; NTM, Z.2018, Z.4085, Z.4122,
Z.4123, Z.5215 (0M9H2137-P), Z.5206 (0M9H2192-Y).
Cobourg Peninsula, NTM Z.1391. Gulf of Carpentaria: QM
G314246, G314247, G315205.
Description
Shape (Fig. 1B). Massive to subspherical, with tapering,
hollow, rudimentary, subconical or volcano-shaped istules,
up to 18 mm long and 30 mm diameter; basal portion buried
beneath sediment and istules protruding through substrate.
58
Halichondriidae from northern Australia
A
C
B
Fig. 3. Amorphinopsis fenestrata: A, light microphotograph of tangential section of ectosomal skeleton (NTM Z.5206), showing spicule tracts
criss-crossing and forming a reticulation of polygonal meshes; B, light microphotograph of section perpendicular to surface (NTM Z.5215)
showing choanosomal skeleton; C, diagram of spicules. Scale bars: A-B, 500 µm; C, 100 µm.
oriented and cavernous at subectosomal region. In istules
choanosomal tracts more compressed in central region
(Fig. 3B), cavernous towards periphery. Single spicules,
including smaller styles scattered through choanosome.
Spicules (Fig. 3C, Table 1). Choanosomal styles and
styloids (thicker in apical third and with basal ends narrower
than the centre), slender, straight or slightly curved at centre,
fusiform, in a great size range (163–895 x 3.3–17.4 µm).
Smaller styles can be transitional to subtylostyles. Relative
proportions of styles, styloids and subtylostyles vary among
populations.
Remarks. The species was originally described by
Ridley (1884) under Leucophloeus, a junior synonym of
Ciocalypta (Van Soest et al. 1990; Erpenbeck & Van Soest
2002) and related to species of Ciocalypta by Hooper
et al. (1997) based on the characteristics of the ectosomal
skeleton, and the presence of styles. As described above, the
growth form of Amorphinopsis fenestrata is characterised
by the presence of istule-like projections of several shapes
Colour. Yellow, brown or pale mauve, alive.
Oscula. Large, up to 10 mm in diameter or, grouped on
a terminal sieve-plate, on apex of istules.
Texture and consistency. Compressible, harsh, easily
torn.
Surface. Irregular, rugose, translucid, hispid, marked in
some specimens with longitudinal channels.
Skeleton. Ectosomal skeleton (Fig. 3A) detachable,
supported by subectosomal multispicular tracts. Formed
by multispicular tracts of larger choanosomal styles, up to
3 spicules abreast, lying tangential to surface, directionless
or criss-crossing, forming a nearly regular reticulation
of polygonal meshes; and, irregular bundles of smaller
ectosomal styles arranged mostly paratangential to surface
as plumose brushes or tufts. Choanosomal skeleton
disorganised, halichondroid criss-cross of both unispicular
and multispicular tracts, containing 5–20 spicules abreast,
with larger choanosomal styles mainly conined to central
region; becoming more wide-meshed, paratangentially
59
B. Alvarez and J. N. A. Hooper
Table 1. Comparison of spicule dimensions among specimens of Amorphinopsis fenestrata.
Specimen
Locality
Styles
BMNH 1882.2.23.255 (Holotype)
Darwin Harbour
193.6–611.8µm (390.4±136.4)
x 5.3–12.7µm (8.6±2)
BMNH 1882.2.23.195
Arafura Sea
252.9–895.3µm (617.7±217.3)
(Ridley’s unnamed variety specimen)
x 5.9–15.4µm (9.6±2.6)
Ternate, Indonesia
210.2–761.6µm (431.1±160)
SMF 680 (Syntype of Suberites oculatus)
x 4.7–17.4µm (8.1±2.8)
Aru Is, Indonesia
306.4–892.8µm (563.3±176)
SMF 971 (Syntype of Ciocalypta oculata maxima)
x 3.3–11.6µm (6.6±1.8)
Parry Shoals
217.8–649µm (448.9±149.7)
NTM Z.3106 (Holotype of Ciocalypta confossa)
x 6.4–12.6µm (9.2±1.9)
Bynoe Harbour
175.1–690.4µm (502.7±155.8)
QM G303560 (Holotype of Ciocalypta oscitans)
x 5.2–13.3µm (8.7±2.2)
NTM Z.5215
Darwin Harbour
178.5–891.1µm (427.1±199.8)
x 4–12.3µm (7.7±2.3)
NTM Z.5207
Bynoe Harbour
163.5–833.1µm (432.4±214.6)
x 3.7–13.7µm (7.3±2.3)
QM G314247
Gulf of Carpentaria
218.8–599.3µm (412.2±120.3)
x 4.3–11.1µm (7.2±2.2)
(e.g. pointy, hollow, pyramidal or volcano-shaped), some
with a system of exhalant channels. They project from
a semi-buried massive base and protrude through the
substrate/sand. The skeletal architecture in these istulose
projections, although similar, is considered here not to be
homologous to the one observed in Ciocalypta species,
which is characterised by a central spicular axis and extraaxial secondary tracts supporting the ectosomal skeleton
(see below under Ciocalypta). Instead, the skeleton of the
istulose projections of this species is formed by multiple
axes, oriented longitudinally and radiating towards the
surface. Based on these arguments, we suggest that the
species is better allocated to the genus Amorphinopsis. It
needs to be noted however, that the skeleton of the present
species is composed totally by styles with a great size range
and lacks of ‘true’ oxeas, a diagnostic characteristic of this
genus. In the majority of specimens examined, including the
holotype, a great proportion of the styles are ‘subacerate’
(styloids) similar to those observed in Aaptos (Hadromerida:
Suberitidae) where the thickest part of the spicule occurs in
the apical third and the basal end is substantially narrow. The
relative proportion of styloids varies among the populations
examined. Subtylostyle modiications, especially in the
smaller styles are also common. We are unable to conirm
whether this type of styles might be considered a derived
form of oxea (as stated in the current deinition of the genus)
and therefore we propose to expand slightly the deinition of
Amorphinopsis to accept species with spicules differentiated
into oxeas and/or styles in a large size range, with smaller
ones concentrated at the surface.
The examination of additional specimens allowed us to
understand the concept of this species better and to conclude
that the material from Beagle Gulf described by Hooper
et al. (1997) under Ciocalypta oscitans, C. confossa and
Amorphinopsis foetida, and the Indonesian species Suberites
oculatus Kieschnick and Ciocalypta oculata maxima
Hentschel are all conspeciic with A. fenestrata.
Amorphinopsis subacerata (Ridley & Dendy, 1886) from
the Philippine Islands is very similar to A. fenestrata and
as stated by those authors they share many characteristics
including the type of styles but differ in external form and in
having larger and thicker styles. Study of local populations
and examination of the type material is required to establish
whether the two species are conspeciic.
Distribution. Amorphinopsis fenestrata is common
throughout the Northern Territory coast (Sahul Province),
from Parry Shoals (NW of Darwin Harbour) to the Gulf of
Carpentaria. The records of Lindgren (1897, 1898) from
Vietnam and China, of Dendy (1922) from the Indian Ocean
and of Burton (1928) from the Malay Archipelago need to
be veriied.
Amorphinopsis maculosa (Pulitzer-Finali, 1996)
comb. nov.
(Figs 1C, 4)
Topsentia maculosa Pulitzer-Finali, 1996: 114.
Axinyssa aplysinoides. – Hooper et al. 1997: 4. Not
Axinyssa aplysinoides Dendy, 1922: 39.
Material examined. HoloType – MSNG 48701, Laing
Is., 4º 09’ S, 144º 52’ E, Papua New Guinea, 6 m depth, 23
August 1986. addiTional specimens – Gulf of Carpentaria:
QM G300854, G301034, G313577, G314255, G314267,
G315207, G320819, G320904. Shoal Bay: QM G303561.
Description
Shape. Thickly encrusting, following substrate with
convoluted ridges and short projections.
Colour. Light grey or yellow alive; yellow inside beige
in ethanol.
Oscula. Inconspicuous, of different diameter, lushed
and irregularly distributed or aggregated in top of the short
projections (Fig. 1C).
Surface. Smooth, lumpy, with dermal skin of reticulated
appearance.
Consistency and texture. Firm but crumbly.
60
Halichondriidae from northern Australia
A
C
B
Fig. 4. Amorphinopsis maculosa (QM G313577): A, light microphotograph of tangential section of ectosomal skeleton showing brushes of
small styles with ends projecting through surface; B, light microphotograph of section perpendicular to surface showing choanosomal skeleton;
C, diagram of spicules. Scale bars: A, 100 µm; B, 500 µm; C, 100 µm.
Skeleton. Ectosomal skeleton (Fig. 4A) consisting of
tangential to paratangential crust, approx. 200–300 µm
thick, supported by choanosomal tracts of spicules and with
disorganised brushes of small styles with ends projecting
through surface, forming a discontinuous palisade spaced
roughly at regular distances. Choanosomal skeleton (Fig.
4B) halichondroid, forming oval to round lacunae, 300–1000
µm in diameter, becoming compact towards surface, with
very little collagen and abundant spicule content. Spicule
tracts long and multispicular, running either towards surface
or parallel to surface at the subectosomal area.
Spicules (Fig. 4C, Table 2). Oxeas and much less
frequent styles in a large size variation, 207–994 x 5–38
µm; small ectosomal styles (and transitional to oxeas),
139–274 x 3–8 µm.
Remarks. The holotype of Topsentia maculosa
Pulitzer-Finali was examined and is comparable in external
morphology and skeletal characteristics to material collected
in the Gulf of Carpentaria and Shoal Bay. The species is
redescribed using this material and assigned to the genus
Amorphinopsis.
Table 2. Comparison of spicule dimensions among specimens of Amorphinopsis maculosa.
Specimen
MSNG 48701
(Holotype of Topsentia maculosa)
QM G313577
Locality
Papua New Guinea
Gulf of Carpentaria
QM G314255
Gulf of Carpentaria
Oxeas
240.2–994.6µm (633.7±255.1)
x 7.4–37.6µm (21.7±9.9)
369.1–863.5µm (675.8±124.6)
x 7.9–20.3µm (15.2±3.2)
207.9–819.5µm (560.1±166.3)
x 4.5–22.1µm (12.6±4)
61
Styles
138.5–259.2µm (211±35.5) [16]
x 3–9.7µm (6.6±1.8) [16]
143.7–273.8µm (187.7±35.2)
x 3.2–7µm (4.8±1)
147.5–240.4µm (174.2±20.9)
x 3–7.8µm (5.6±1)
B. Alvarez and J. N. A. Hooper
Amorphinopsis maculosa is very similar in habit and
skeletal organisation to A. fenestrata (Ridley, 1884) but
that species has only styles as megascleres and lacks the
oxeas of variable sizes present in this species (see above).
Fistulous projections, as observed in A. fenestrata, are not
present in the examined material of A. maculosa.
The specimen assigned to Axinyssa aplysinoides
(Dendy) by Hooper et. al. (1997) was re-examined and has
characteristics (i.e. habit, ectosomal skeleton, oxeas and
smaller styloids) that agree with Amorphinopsis maculosa.
Amorphinopsis maculosa was also compared to other
species of Amorphinopsis recorded for northern Australia and
adjacent biogeographical regions – Amorphinopsis excavans
(Dendy, 1889) from the Indian Ocean, A. foetida (Dendy,
1889) from the Gulf of Manaar, A. maza (De Laubenfels,
1954), A. oculata (Kieschnick,1896), and A. sacciformis
from Indonesia. Amorphinopsis excavans has similar habit,
skeletal structure and spicule composition, but differs in
the organisation of the ectosomal skeleton; in A. excavans
it consists of a tangential layer of thick intercrossing tracts
of large oxeas, and loose oxeas of all sizes with small styles
illing up the spaces (Erpenbeck & Van Soest 2002). The
material of A. excavans from northern Australia described
above also has a clearly tangential and detachable ectosomal
skeleton of large oxeas grouped in bundles and small styles
tangentially to paratangentially oriented, sometimes in
disorganised tufts. Other characteristics that differentiate
A. maculosa from A. excavans are the predominance of
small styles in the ectosome (instead oxeas and styles as
reported in most populations of A. excavans) and the lacunar
appearance of the choanosomal skeleton.
Amorphinopsis foetida (type specimen BMNH
1889.1.21.55, examined) is also a massive and slightly
lobose species but with a skeletal architecture and spicule
geometry different to A. maculosa. The ectosomal skeleton
of A. foetida is formed by a halichondroid dermal crust,
tangential-paratangetially oriented, 500–900 µm thick,
with a mixture of oxeas and small styles projecting through
the surface, and with vague tracts and small rounded
meshes approx. 100–300 µm. The choanosomal skeleton
is halichondroid, cavernous, with large rounded lacunae,
500–700 µm approx., and ill-formed multispicular tracts
and ibres irregularly oriented through the skeleton; less
compact than the ectosome and bounded by little collagen.
The spicules are mixture of oxeas of different thickness and
sizes, characteristically curved or sinuous (165.4–720.6 µm
x 3.3–16.6 µm). The smaller styles are curved, and some are
transitional to oxeas (154–281.7µm x 2.7–7.9µm).
Amorphinopsis maza was re-examined by Erpenbeck
& Van Soest (2002) and differs from A. maculosa in habit,
skeletal organisation and spicule composition.
Suberites oculatus Kieschnick, 1896, accepted as
Amorphinopsis oculata (Van Soest et al. 2008) is a synonym
of Amorphinopsis fenestrata (see above).
Ciocalypta sacciformis Thiele, 1900 (syntypes, SMF
685, 1818, Ternate Maluku Sea, 1894, coll. Kükenthal
W., examined) was transferred from its original genus to
Ciocalapata by De Laubenfels (1936) and related it to
species-like Ciocalypta including oxeas and styles. The
species was interpreted as Amorphinopsis by Hooper et al.
(1997). The revision of the type material indicates that
the specimens described by Thiele agree better with the
current concept of Halichondria but not with Ciocalypta,
Ciocalapata or Amorphinopsis as suggested by previous
authors. The syntypes examined are small fragments with
a pouch-like shape as described by Thiele. The ectosomal
skeleton is a thick tangential to paratangential layer up to
500–800 µm, with single spicules and short tracts crisscrossing in a disorganised manner. The choanosomal
skeleton is disorganised with bundles of spicules irregularly
spaced and running towards the surface where they merge
with the ectosomal skeleton. The skeleton is formed entirely
by oxeas in a great size range 195–706 µm. The skeleton of
SMF 1818 includes styles in low frequency 463–525 µm
(491.2±17.4) x 6–16 µm. As admitted by Thiele, the styles
might be modiications of oxeas.
Distribution. Papua New Guinea (Eastern Coral
Triangle Province), Gulf of Carpentaria and outer region of
Shoal Bay (Sahul Shelf province) between 6–28 m depth.
It is also found in Torres Strait (Northeastern Australian
Province) (Alvarez & Hooper unpublished data).
Remarks on Amorphinopsis. Amorphinopsis includes
approx. 13 valid species (Van Soest et al. 2008) distributed
mainly throughout the Indian Ocean but with species also
recorded from the Atlantic and Mediterranean oceans.
The genus is represented in northern Australia (Sahul
Shelf Province) by three species – Amorphinopsis excavans,
A. fenestrata comb. nov. and A. maculosa comb. nov.
Amorphinopsis foetida was recorded by Hooper et al.
(1997) from the Beagle Gulf but it is concluded here that
the specimen is better allocated to A. fenestrata.
The species originally described as Leucophloeus
fenestratus Ridley, 1884 is common in the study area. Its
habitat and skeletal characteristics have been interpreted as
typical from the genus Ciocalypta but as discussed above,
these similarities are considered not homologous and we
propose to include the species in Amorphinopsis. The
choanosomal skeleton of A. fenestrata is formed entirely
by styloids and lacks ‘true’ oxeas as seen in other species
of Amorphinopsis and other genera of Halichondriidae. We
propose to expand the deinition of Amorphinopsis slightly
to include species with spicules differentiated into oxeas
and/or styles.
Genus Axinyssa Lendenfeld, 1897
Gender feminine. Type species, by original designation,
Axinyssa topsenti Lendenfeld, 1897: 116. Recent, western
Indian Ocean.
62
Axinyssa bergquistae (Hooper et al., 1997) com. nov.
(Figs 1D, 5)
Halichondria bergquistae Hooper et al., 1997: 45.
Halichondriidae from northern Australia
membrane; becoming distinctively conulose at erect
columns or digits. Conules up to 2 mm long organised
in longitudinal rows along erect columns and digits, with
brushes of larger choanosomal spicules protruding through
surface.
Texture and consistency. Hispid due to projection of
spicules through surface, irm and incompressible.
Skeleton (Fig. 5A,B). Ectosome without specialised
skeleton, with lightly coloured collagenous skin.
Choanosomal skeleton halichondroid, with high spicule
density in deeper regions; becoming more organised at
subectosomal region, with multispicular spicule tracts,
50– 200 µm running longitudinally and ascending towards
surface, becoming more radial and plumose near periphery;
ending in disorganised brushes that project through
ectosome.
Spicules (Table 3, Fig. 5C). Mixture of oxeas of variable
thickness and length (354–948 x 4–27.8 µm), slightly bent
Material examined. HoloType – QM G303351, East
Point Bommies, Darwin Harbour, Northern Territory,
Australia, 12°24.083’ S, 130°48.14’ E, 10 m depth,
23 September 1993, coll. Hooper, JNA & Hobbs, LJ.
addiTional specimens – Cartier Island: QM G301059.
Bynoe Harbour: Z.5224 (0M9H2375-X); Z. 5901. Darwin
Harbour: Z.5976. Gulf of Carpentaria: QM 313572.
Description
Shape (Fig 1D). Massive-lobate, bulbous-digitate; with
erect columns or irregular coalescent plates. Individuals
approx. 70–100 mm high, 45–200 mm thick.
Colour. Purple-mauve, grey-brown; some individuals
with lighter tinges.
Oscula. Variable in size (3–7mm diameter), conspicuous,
discrete, with raised white and opaque membranous lips
(Fig. 1D), irregularly distributed.
Surface. Smooth to microconulose at base with
shallow and meandering channels covered by a translucent
A
C
B
Fig. 5. Axinyssa bergquistae: A, light microphotograph of perpendicular section (NTM Z. 5976) showing choanosomal spicule tracts projecting
through surface; B, light microphotograph of perpendicular section through surface (NTM Z.5224), showing choanosomal skeleton with
longitudinal plumose tracts; C, diagram of spicules. Scale bars: A-B, 500 µm; C, 100 µm.
63
B. Alvarez and J. N. A. Hooper
Axinyssa mertoni (Hentschel, 1912) com. nov.
(Figs 1E, 6)
Ciocalypta mertoni Hentschel, 1912: 424; Burton 1934:
564.
Halichondria tyleri. – Hooper & Wiedenmayer 1994:
209.
Halichondria mertoni. – Hooper et al. 1997: 52.
Pseudaxinyssa pitys De Laubenfels, 1954:178; Bergquist
1965 : 175 [?].
Axinyssa pitys. – Hooper & Bergquist 1992: 102.
Material examined. Type maTerial – Ciocalypta
mertoni, holotype, SMF 1608, Aru Is, North of Penambulai;
station 10, 8 m depth, 2 April 1908, coll. Merton exp. 1908.
Pseudaxinyssa pitys, holotype, USNM 23103, Caroline
Islands, Palau Is, Koror I., Iwayama Bay, 2 m depth, 1
September 1949, coll. De Laubenfels, M.W. addiTional
specimens –Darwin Harbour: NTM Z.5221 (0M9H2189-V).
Description
Shape (Fig. 1E, 6A). Massive with conspicuous istules
up to 50 mm long, 1–2 mm thick, projecting from semiburied basal portion up to 100 mm diameter.
Colour. Grey alive.
Oscula. On top of istules, lushed, less than 5 mm
diameter.
Surface. Regularly conulose with transluscent membrane
stretching over conules; marked with choanosomal axes.
Skeleton (Fig. 6B). Ectosomal skeleton absent.
Choanosomal skeleton plumose to halichondroid with
multispicular axes of spicules running longitudinally, nearly
parallel and close to each other, anastomosing and diverging
towards surface and becoming dendritic, bounded with
collagen and ending in disorganised plumose brushes that
project through surface. Spongin ibres ill-deined and short,
direction-less, embedding spicule tracts.
Spicules (Fig. 6C, Table 4). Oxeas, hastate, slightly bent,
straight or crooked, 476–1470.2 x 11.2–29.7 µm. Style and
strongylote modiications also present.
Remarks. Ciocalypta mertoni is conspecific with
Pseudaxinyssa pitys and is better allocated to the genus
and sometimes slightly sinuous. Styloid modiications
common.
Remarks. The species was initially described under
Halichondria. Examination of additional material
considered conspeciic with the specimen described by
Hooper et al. (1997) allowed us to conclude that the
species is better allocated in Axinyssa. The plumose conules
observed in some areas of the surface of A. bergquistae
resembles Axinyssa mertoni (described below), but that
species has a more lax skeleton with less spicular density
and more collagen. It differs also in general shape, colour
and the oscula morphology.
Axinyssa bergquistae is comparable to A. valida (Thiele,
1899: 12) [holotype NHMB 13, examined] in external
morphology, skeletal organisation and size of spicules
[(290.3–859.3 µm (599±201.8) x 9.7–34.8 µm (20.6±8.3)]
and could possibly be conspeciic. But both the skeleton and
the size of spicules of most Axinyssa species are very similar
and the separation of species is dificult and subjective
(see below under remarks on the genus). The examination
of more material from Indonesia (Alvarez & De Voogd
in progress) will help to determine whether the northern
Australian populations belong to the same species.
Distribution. Darwin Harbour and Bynoe Harbour
(Sahul Shelf Province). Probably present also in the
Northeast Australian Shelf Province (Alvarez & Hooper
unpublished data).
Table 3. Comparison of spicule dimensions among specimens of
Axinyssa bergquistae.
Specimen
QM G303351
(Holotype of
Halichondria
bergquistae)
NTM Z.5224
Locality
Oxeas
Darwin Harbour 407.3–793.8µm (619.7±82.8)
x 10.3–27.8µm (15.5±4.3)
QM G313572
Gulf of
Carpentaria
A
Bynoe Harbour
366.3–948.1µm (749.2±135.7)
x 4.5–33.2µm (21.3±7)
353.6–747.3µm (618.3±77.5)
x 4–20.9µm (14.7±4.2)
C
B
Fig. 6. Axinyssa mertoni: A, preserved specimen (NTM Z.5221); B, light microphotograph of perpendicular section through surface showing
choanosomal skeleton with longitudinal tracts projecting through surface (Holotype, SMF 1608); C, diagram of spicules. Scale bars: A,B,
500 µm; C, 100 µm.
64
Halichondriidae from northern Australia
Axinyssa. The type material of both species were examined
here, and they are identical in all their characteristics.
Ciocalypta mertoni was considered a junior synonym of
Halichondria tyleri by Hooper & Wiedenmayer (1994)
(following Burton 1959), but this synonym was later
rejected by Hooper et al. (1997) who considered it a valid
species of Halichondria, but admitted that the lack of an
ectosomal skeleton was atypical of that genus. The species
agrees well with the concept of Axinyssa, as it lacks an
ectosomal skeleton and has a halichondroid to vaguely
plumose choanosomal skeleton bounded by relatively high
amounts of collagen and formed by oxeas of variable size
and common styles and strongyles. The growth form of
this species however, is not reported for other species of
Axinyssa, thus a slight expansion in the diagnosis of the
genus is necessary to accommodate species with istulose
projections.
A single specimen found in Darwin Harbour is assigned
to this species. It differs from the type material examined
only on spicule dimensions. The oxeas of the Darwin
specimen are in average longer and thicker, but this might
correspond to intraspeciic variation within the species.
Distribution. Axinyssa mertoni is rare within the study
area with only one specimen recorded here. The distribution
of the species is extended to the Sahul Shelf and the Tropical
Northwestern Paciic provinces. The record of Bergquist
(1965) from Palau needs to be veriied as there are several
sympatric species of Axinyssa occurring in the area (Lori
Bell Colin pers. comm.).
Remarks on Axynissa. Axinyssa is represented in the
area of the Indo-Paciic by several species: A. aculeata
Wilson, 1925 (Philippines); A. aplysinoides (Dendy, 1922)
(Seychelles); A. oinops (De Laubenfels, 1954) (central
West Paciic); A. topsenti Lendenfeld, 1897 (Tanzania);
A. variabilis Lindgren, 1897 (Malaysia); and A. valida
(Thiele, 1899) (Indonesia). The new combinations
established in this revision extend the list by two additional
species – A. mertoni (Sahul Shelf and Tropical Northwest
Paciic provinces) and A. bergquistae (Sahul Shelf and
probably Northeast Australian Shelf). Two additional
species from the central Indo-Pacific region are also
referred to Axinyssa after examination of type material
– Ciocalypta rutila gracilis Hentschel, 1912 (SMF 1566,
Aru Is, examined; see under remarks of Ciocalypta), and
Phycopsis terpnis De Laubenfels, 1954 (Caroline Is, Central
Paciic, USNM 23061, examined).
We were not able to revise all the type material of the
Indo-Paciic species of Axinyssa thoroughly, therefore it
remains inconclusive whether A. mertoni and A. bergquistae
might be conspeciic with other species recorded from
the region. As mentioned above, the skeletal organisation
and the size of spicules among Axinyssa species is very
similar and separation of species is subjective. The external
morphological characteristics seem to be more discrete, but
study of individual populations is necessary to determine
the actual range of variability present within these species.
Table 4. Comparison of spicule dimensions among specimens of
Axinyssa mertoni.
Specimen
SMF 1608
Locality
Aru Island,
Indonesia
Caroline Is,
Central West
Paciic
USNM
23103
(holotype of
A. pitys)
NTM Z.5221 Darwin Harbour,
NT
Oxeas
515.6–779.6µm (691.4±61)
x 11.2–21.7µm (18±2.5)
514.3–871µm (788.2±74.9)
x 8.6–18.5µm (15±2.5)
476–1470.2µm (979.5±285.9)
x 14.7–29.7µm (21.6±4.8)
Study of different populations of Axinyssa through Indonesia
(Alvarez & De Voogd unpublished data) is currently in
progress and will help to re-deine the limits of Axinyssa
species.
The diagnosis of Axinyssa is here expanded to include
species like Axinyssa mertoni with istulose projections. We
note however, that the skeletal organisation of the istules
observed in A. mertoni is considered not to be homologous
with the organisation observed in species of Ciocalypta. The
istulose projections of Ciocalypta spp. are transparent with
the skeleton formed by a central axis of spicules and extraaxial tracts diverging towards the surface. The istulose
projections of A. mertoni are opaque, tough-cartilagineous,
arising from a massive base, and without a central axis of
spicules which is diagnostic for Ciocalypta.
Axinyssa mertoni shares with the Indian Ocean
Ciocalypta digitata (Dendy, 1905) the lack of an ectosomal
skeleton, and the presence of istulose projections; however,
the istules of C. digitata are transparent with a spicular axis
from which thick bundles of spicules diverge towards the
surface ending in conules (Erpenbeck & Van Soest, 2002).
In our view the expansion of the deinition of Axinyssa to
include species with istular projections such as A. mertoni
does not affect the current position of C. digitata or the
deinition of the Ciocalypta.
The current position of the genus Axinyssa within the
family Halichondriidae is debatable as molecular data
(Erpenbeck et al. 2005) indicate that some species currently
allocated to this genus are related to other dictyonellid genera
such as Acanthella and Dictyonella. From a morphological
point of view, the lack of an ectosomal skeleton, the
presence of abundant collagen in the skeleton and the
common occurrence of styles, strongyles and transitional
forms, also points to possible relationships with members
of Dictyonellidae. These relationships should be further
explored to conirm the placement of this genus within the
family Halichondriidae. However, taxonomic veriication
of the species of Axinyssa used in the molecular analyses
should also be taken in consideration, particularly given
the paucity of morphometric characters in this group and
our still rudimentary understanding of character homology.
65
Genus Ciocalypta Bowerbank, 1862.
Gender feminine. Type species, by monotypy, Ciocalypta
penicillus Bowerbank, 1862. Recent, eastern Atlantic Ocean.
B. Alvarez and J. N. A. Hooper
Ciocalypta heterostyla Hentschel, 1912
(Figs 1F, 7)
Ciocalypta heterostyla Hentschel, 1912: 424; Hooper
et al. 1997: 36.
Material examined. HoloType – SMF 1569, Aru Is,
N Penanbuli, 8 m depth, 2 April 1908, coll. H. Merton.
addiTional specimens – Darwin Harbour, NTM Z.5902.
Description
Shape (Fig. 1F). Fistulose, with semi-buried and massive
base. Fistules pointed, projecting perpendicularly from base,
20–30 mm long, less than 10 mm diameter wide, slightly
translucent especially at tips.
Colour. Light yellow.
Oscula. Apical on istules.
Surface. Smooth, microconulose.
Ectosomal skeleton (Fig. 7A). Thin layer formed
by tangentially oriented pauci- to multispicular tracts
of spicules, crossing over and forming a reticulation of
triangular meshes; supported by choanosomal tracts.
Choanosomal skeleton. Differentiated at fistules,
with central column formed by thick multispicular tracts
oriented longitudinally and expanding into thick brush
at tip of istule (Fig. 7B). Secondary tracts, 20–100 µm
thick, slightly plumose, diverging nearly perpendicularly
from central column toward surface, regularly spaced and
connected irregularly by unispicular-paucispicular tracts of
spicules; becoming thicker and ending in fan-like brushes
at the subectosomal area, generally with smaller spicules
oriented with their ends towards surface. Skeleton at base
C
A
B
Fig. 7. Ciocalypta heterostyla (Holotype SMF 1569): A, light microphotograph of tangential section of ectosomal skeleton, showing tracts
of spicules, forming a reticulation of triangular meshes; B, light microphotograph showing choanosomal skeleton at a istule. C, diagram of
spicules. Scale bars: A-B, 500 µm; C, 100 µm.
66
Halichondriidae from northern Australia
Ciocalypta stalagmites Hentschel, 1912
(Figs 1G–H, 8)
Ciocalypta stalagmites Hentschel, 1912: 426.
Halichondria tyleri.– Hooper et al. 1997.
Material examined. lecToType – SMF 1567 (here
designated, a representative specimen of the species,
chosen from six examined specimens deposited at SMF
and registered as syntypes with a single accession number),
Aru Is, Mimien I., Indonesia, 15 m depth, 9 April 1908,
coll. Meron, H. paralecToTypes – SMF 1537, Aru Is,
Sungi Manumbai (Kapala Sungi), Indonesia, station 17,
Merton Expedition Aru and Kei Is 1907-1908, 20 m depth,
5 May 1908, coll. Merton, H. SMF1574, Aru I., SW Lola
I, Indonesia, station 9, Merton Expedition Aru and Kei
Is 1907-1908, 8–10 m depth, 1 April 1908, coll. Merton,
H. SMF 1595, Aru Is, Bambu I., Indonesia, station 11,
Merton Expedition Aru and Kei Is 1907-1908, 10 m depth,
3 April 1908, coll. Merton, H. SMF 1623, Aru Is, SW
Mariri and Leer Is, Indonesia, Merton Expedition Aru and
Kei Is 1907-1908, 6–10 m depth, 31 March 1908, coll.
Merton, H. SMF 1627, Aru Is, N Penambulai I., Indonesia,
Merton Expedition Aru and Kei Is 1907-1908 , 8 m depth,
2 April 1908, coll. Merton, H. addiTional specimens –
Parry Shoals, NT, NTM Z.3133. Charles Point, NT, NTM
Z.5230 (0M9H2574-P). Bynoe Harbour: NTM Z.5226
(0M9H2511-V). Darwin Harbour: NTM Z.941, Z.5210
(0M9H2571-M), Z.5229 (0M9H2299-Q), Z.5977. Cobourg
Peninsula: NTM Z.592, Z.1358, Z.1395, Z.3286. Wessel Is:
Z.3920, Z.5218 (0M9H2666-P).
Description
Shape (Figs 1G–H, 8A). Flat cushion-shaped or
massive base, buried or semi-buried, strongly attached to
substrate, up to 35 mm thick, 4–100 mm diameter, with
istules projecting perpendicularly above surface. Fistules
halichondroid, with directionless multispicular tracts and
single spicules in confused reticulation.
Spicules (Fig. 7C, Table 5). Mixture of styles in a large
range of sizes, straight to slightly curved, some sinuous,
some with rounded ends, 184–809 x 4–19 µm.
Remarks. Ciocalypta heterostyla was originally
described from the Aru Is (Indonesia) and it has not been redescribed or recorded until this present study. The material
from Darwin Harbour agrees closely with the type. Only
the spicule dimensions are on average slightly larger in the
specimen from Darwin. The species is not very conspicuous
due to its cryptic habit. It is apparently rare in the study area
with only one specimen collected so far.
Hooper &Wiedenmayer (1994) followed Burton (1959)
and considered this species a synonym of Ciocalypta tyleri
Bowerbank. The two species are certainly similar in external
morphology and skeletal organisation, however the skeleton
of C. heterostyla is formed exclusively by styles instead of
oxeas, and the skeleton within the istules is much more
organised with a regular reticulation. Because of these
characters we considered that C. heterostyla is not only a
valid species but also it can be easily differentiated from its
South African relative.
Distribution. Known only from the type locality (Aru
Is, Indonesia) and from Darwin Harbour (Sahul Shelf
province). It is found between 8–12 m depth.
Table 5. Comparison of spicule dimensions between specimens of
Ciocalypta heterostyla.
Specimen Locality
SMF
Aru Is, Indonesia
1569
Darwin Harbour
NTM
Z.5902
Styles
199.9–584.1µm (387.3±137)
x 3.5–14.8µm (8.3±3.2)
184–809.2µm (466.5±229.1)
x 3.9–18.6µm (9.4±5.5)
A
B
D
C
Fig. 8. Ciocalypta stalagmites (Lectotype, SMF 1567): A, Preserved lectotype; B, light microphotograph showing choanosomal skeleton at
istule; C, light microphotograph showing choanosomal skeleton at base; D, diagram of spicules. Scale bars: A, 2 cm; B-C, 500 µm; D, 100 µm.
67
B. Alvarez and J. N. A. Hooper
diagnostic to separate C. stalagmites from other Ciocalypta
species.
Ciocalypta stalagmites is very similar to C. tyleri
Bowerbank, 1873 from South Africa, and it was interpreted
as such by Hooper et al. (1997) but allocated to the genus
Halichondria (following Van Soest et al. 1990; Van Soest
1991). Both species are now referred to Ciocalypta under
the revised concept of Halichondriidae (Erpenbeck &
Van Soest 2002). The two species are similar in external
morphology and skeletal organisation, however, the skeleton
of C. stalagmites seems to be less organised. Despite
similarities between these two species, it is unlikely that
the Indonesian and northern Australian populations are
conspeciic with their South African relatives, thus we
propose to reserve C. stalagmites for those populations
inhabiting the Sahul Province and adjacent areas. Future
independent evidence might demonstrate whether
C. tyleri and C. stalagmites belong to a complex of cryptic
species that cannot be easily separated using traditional
morphological characters, or are instead conspeciic and
genuinely a widely distributed species.
Ciocalypta stalagmites is also very similar to C. vansoesti
(Hooper et al., 1997). comb. nov. The two species have
similar growth form and skeletal characteristics, but they
can be differentiated from each other by some distinctive
characteristics (see below).
Distribution. Ciocalypta stalagmites is very common
throughout the northern Australian localities of the Sahul
Shelf province and its distribution extends to adjacent
provinces including the Northeast Australian Shelf,
Papua New Guinea (Alvarez & Hooper unpublished), and
Indonesia (Alvarez & De Voogd unpublished). It is found
from the intertidal zone to 40 m.
sharply pointed, globular or lattened, mammiform, tubular,
rounded, 5–125 mm long, laterally fused sometimes or
branching at tips.
Colour. Base generally pale-beige. Fistules, transparent,
mauve-brown-purple, greenish-yellow. Internally, beige.
Oscula. Generally at apex of fistules or tubes but
also observed irregularly distributed along istules, with
membranous rims.
Consistency and Texture. Compressible-spongy, easily
torn.
Surface. At istules, smooth to slightly conulose, marked
with longitudinal rows of minute conules. Choanosomal
tracts of spicules, visible through the ectosome of translucent
specimens. At base smooth, opaque, rough, spiculous.
Skeleton. Ectosomal skeleton, tangential layer of variable
thickness (5–300 µm) easily peeled in some specimens,
formed by a dense mass of smaller oxeas and supported by
choanosomal skeleton. Choanosomal skeleton (Fig. 8C)
at base, halichondroid, densely spiculous, with mixture of
large and smaller oxeas and no distinct tracts of spicules.
In istules (Fig. 8B), becoming distinctive and organised,
with central column of large spicules oriented longitudinally
and smaller spicules criss-crossing. Multispicular tracts
generally formed by medium-size spicules radiating from
central column towards surface, connected by shorter tracts
and single spicules, forming a regular reticulation. Large
subectosomal spaces (up to 5 mm diameter) in some areas.
Spicules (Fig. 8D, Table 6). Two size classes of oxeas:
I, smaller, thinner, fusiform, straight or slightly lexuous
(147–321 x 4–10 µm); II, large, thick and slightly curved,
fusiform, occasionally crooked-sinuous (378–886 x11–40
µm), and styles in equivalent size categories are common.
Remarks. The growth form of Ciocalypta stalagmites is
remarkably variable. Fortunately this variation is also well
represented in the type material. The external colouration
is also variable and is possibly related to the presence
of different cyanobacterial associations. The skeletal
organisation, composition and size of spicules, however, are
consistent and very similar among specimens with different
habits and colouration. A very consistent characteristic
through all the populations we examined is that the oxeas are
differentiated in size categories not only by their length but
also by their thickness. Proper statistic and morphometric
analyses could be employed to demonstrate whether this is
Ciocalypta vansoesti (Hooper et al., 1997) comb. nov.
(Figs 1I, 9)
Halichondria vansoesti Hooper et al., 1997: 37.
Material examined. As listed by Hooper et al. (1997).
addiTional specimens – Darwin Harbour: Z.5904, Z.5217
(0M9H2568-J), Z.5903, Z.5978. Bynoe Harbour: Z.5212
(0M9H2498-I).Gulf of Carpentaria: QM G303524.
Remarks. This species was well described by Hooper
et al. (1997), but allocated to the genus Halichondria
(following Van Soest et al. 1990; Van Soest 1991). It is
Table 6. Comparison of spicule dimensions between specimens of Ciocalypta stalagmites.
Specimen
SMF 1567 (Lectotype)
Locality
Aru Is, Indonesia
NTM Z.5212
Bynoe Harbour
NTM Z.5210
Darwin Harbour
NTM Z.5218
Wessel Is
Oxea type I
167.8–240.2µm (200.8±17.4)
x 4.4–10µm (7.9±1.3)
165.3–320.6µm (238.2±34)
x 5.5–10.3µm (7.6±1.2)
188.3–258.7µm (214±19.1)
x 5.6–9.2µm (7.4±0.9)
147.2–202.7µm (169.1±12.8)
x 5.3–9.4µm (6.7±0.9)
68
Oxea type II
377.7–837µm (626.2±102.8)
x 12.3–37.1µm (18.7±4.7)
411.4–659.5µm (524.9±56.8)
x 11.1–26.3µm (18.9±4.1)
440.1–662.8µm (601.8±53.9)
x 11.3–38.5µm (21.9±7.7)
437.7–886.5µm (588.1±135.1)
x 8.7–39.5µm (21.7±7.9)
Halichondriidae from northern Australia
A
B
Fig. 9. Ciocalypta vansoesti: A, preserved paratype specimen (QM G303450); B, light microphotograph showing choanosomal skeleton at
istule. Scale bars: A, 20 mm; B, 500 µm.
now referred to Ciocalypta under the revised concept of
Halichondriidae (Erpenbeck & Van Soest 2002).
As mentioned by Hooper et al. (1997) and above,
Ciocalypta vansoesti is closely related to C. stalagmites,
both with similar growth form, skeletal organisation and
type of spicules. Both species have istules projecting from
a buried-semiburied mass, but in the case of C. vansoesti
the istules are transluscent-white with the surface regularly
conulose and subectosomal tracts and central column visible
beneath (Figs 1I, 9A). However, the skeleton in the istules,
when compared to C. stalagmites, is denser and slightly
disorganised; the central column is not as condensed and the
extra axial reticulation is vague (Fig. 9B, Hooper et al. 1997,
igs. 22a, 23a). The dimensions of the oxeas are similar and
overlap with those of C. stalagmites, but in C. vansoesti
there is not a clear difference in size categories, with the
two classes of spicules overlapping both in length and in
width (Table 7).
Ciocalypta vansoesti is also similar in some of the
ield characteristics to Axinyssa mertoni, indicating once
again that istule-like growth forms are common among
halichondrids and so are not useful to differentiate species
unless they are used in combination with other skeletal
characters.
Distribution. Ciocalypta vansoesti is common in
Darwin Harbour and Bynoe Harbour. It is also recorded
from Cobourg Peninsula and the Gulf of Carpentaria. It is
found intertidally and subtidally to 40 m depth.
Remarks on Ciocalypta. Burton (1959) considered
many species of halichondriid genera with a fistulose
habit conspeciic with Ciocalypta penicillus (type species
of Ciocalypta). These synonyms were not properly
substantiated and some of them have been rejected by
Hooper et al. (1997) and the present revision, i.e. Ciocalypta
heterostyla, C. stalagmites, C. tyleri, C. oculata maxima
(referred here to Amorphinopsis fenestrata), and C. mertoni
(referred here to Axinyssa mertoni).
Additional species of Ciocalypta recorded from the Sahul
Shelf Province and adjacent areas besides the three revised
here (i.e. C. heterostyla, C. stalagmites and C. vansoesti) are
C. rutila gracilis Hentschel, 1912 (see below), C. digitata
(Dendy, 1905, as Collocalypta), C. melichlora Sollas, 1902,
C. rutila Sollas, 1902, and C. simplex Thiele, 1900: 76.
One of the syntypes of Ciocalypta rutila gracilis
Hentschel, 1912 (SMF 1566, examined) belongs in Axinyssa.
Both the external morphology (based on Hentschel’s
description) and the arrangement of skeleton agree with the
concept of that genus. The skeleton of the material examined
is formed by two classes of oxeas: straight 480.8–706.4 µm
(601.6±76.7)x 9–24 µm (17±3.1) and vermicular, crooked
sinuous, relatively thinner, and often bent up to a 90 degree
angle, 193–600.7 µm (406.7±110.6) x 5.3–16 µm (10.6±3)
[18]; less often styles are also present. Specimens recorded
for Northeast Australian Shelf and Indonesia correspond
with the type examined (Alvarez & Hooper unpublished
data; Alvarez & De Voogd unpublished data) and will
be redescribed under the name of Axinyssa gracilis in
forthcoming publications.
Ciocalypta digitata (Dendy, 1905) resembles Ciocalypta
stalagmites and C. vansoesti in habit but differs in skeletal
Table 7. Comparison of spicule dimensions between specimens of Ciocalypta vansoesti.
Specimen
NTM Z.2648
QM G303450
Locality
Darwin Harbour,
East Point
Bynoe Harbour
QM G303524
Gulf of Carpentaria
Oxea type I
185.3–459.7µm (320±60.9)
x 6.4–9.5µm (8±0.9)
194.1–361.7µm (275.3±45.6)
x 2.1–9.8µm (6.8±1.8)
237.5–429.2µm (314.9±56.5)
x 6.4–11.9µm (8.4±1.3)
69
Oxea type II
475.2–616.4µm (546.2±38.3)
x 14–21.6µm (17.9±2.3)
382–662.1µm (536.1±78.2)
x 6.8–30.8µm (16.8±6)
432.8–677.3µm (541.2±58.7)
x 8.2–31.9µm (17.5±4.8)
B. Alvarez and J. N. A. Hooper
subectosomal region, with a vague reticulation of ascending
tracts connected by short bundles and single spicules.
Near surface, spicule tracts become more deined and
even slightly plumose, supporting ectosomal skeleton and
occasionally protruding through surface (Fig. 11B).
Skeleton of some specimens is obscured by granular
pigments and cyanobacteria when examined under light
microscope.
Spicules (Fig. 11C, Table 8). Mixture of oxeas not
separable into size categories 112–389 x 2–12 µm, hastate,
pointed, straight or occasionally slightly bent in middle.
Remarks. Halichondria carotenoidea is diagnosed
by a unique combination of features (i.e. growth form,
organisation of the skeleton, and the size and composition of
spicules) not found in other species of Halichondria revised
here or reported for adjacent areas (see Table 9).
The skeleton and spicule composition of all the
material examined is almost identical to Halichondria
(H.) microbiana sp. nov. (see below). Both species are
sympatric and include a high density of cyanobacteria in
the choanosome. The two species clearly differ in shape,
which is branching to arborescent in H.(H.) carotenoidea
and massive to cushion shape in H. (H.) microbiana. Both
species can be further distinguished by the size of the
oxeas, which are separated into class categories in H.(H.)
microbiana but not in H.(H.) carotenoidea.
The material described by Hooper et al. (1997) as
Halichondria stalagmites (Hentschel) is a composite of H.
(H.) carotenoidea (specimens NTM Z.2651 and Z.3147)
and H (H.) microbiana (specimens nTm Z. 131, Z.1097, Z.
1991), and it does not agree with the concept of the species
described by Hentschel (1912) as Ciocalypta stalagmites
(see above).
Distribution. Halichondria carotenoidea is common in
Darwin and Bynoe Harbours, and it was also observed at
Wessel Is (Raragala I.).
Etymology. Named after carotenoid, a pigment which
might give the red-orange colour to the species. It is intended
as a noun in apposition.
organisation and composition and size of spicules. This
species lacks an ectosomal skeleton, but the fistular
processes have spicular axis and extra-axial tracts as in
other species of Ciocalypta.
Genus Halichondria Fleming, 1828
Gender feminine. Type species, by subsequent
designation (of Bowerbank 1862), Spongia panicea Pallas,
1766. Recent, East England.
Subgenus Halichondria (Halichondria) Fleming, 1828
Halichondria (Halichondria) carotenoidea sp. nov.
(Figs 10A,B, 11)
Halichondria stalagmites. – Hooper et al. 1997:
43 [in part, not Z.131, Z.1097, Z.1991 = Halichondria
(Halichondria) microbiana sp. nov.]
Phakellia sp. 614. – Hooper et al. 1992.
Axinella sp. 244. – Hooper et al. 1992.
Material examined. HoloType – nTm Z.5909 , off
Dudley Point, Fannie Bay, Darwin Harbour, N.T., 12°24.96’
S, 130°48.83’ E, 4–7 m depth, 4 June 2002, coll. Alvarez,
B and party. paraTypes.– NTM Z.4451 (0M9H2036-G),
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula,
Darwin Harbour, N.T., 12°29.103’ S, 130°47.111’ E, 8–14 m
depth, 7 May 2002, coll. Alvarez, B and party. NTM Z.5225
(0M9H2462-S), Spencer Point, Indian I., Bynoe Harbour,
N.T., 12°35.351’ S, 130°31.454’ E, 6–8 m depth, 11 June
2003, coll. Alvarez, B and party. addiTional specimens – N
Adele I., Collier Bay, NW Shelf: Z.712. Parry Shoals, NT:
NTM Z.3147. Darwin Harbour NT: NTM Z.194, Z.919,
Z.934, Z.986, Z.1979, Z.2086, Z.2245, Z.2651, Z.2697.
Description
Shape (Figs 10 A,B). Fan shaped, digito-palmate,
multiplanar, arborescent, erect, stalked, bifurcated at base,
with complex branching; branches or digits flattened
generally with pointed tips but round tips also common;
specimens reaching up to 350 mm high, 300 mm wide.
Colour. Orange. Blue-grey, or beige in alcohol
Consistency and texture. Soft, easily torn, rubbery.
Oscula. Regularly distributed along lateral side of
branches (Fig 10B) less than 5 mm in diameter, generally
in rows, with membranous rims slightly elevated. Some
specimens with subectosomal thin canals ending in oscula.
Surface. Smooth, wrinkled, evenly covered with microconules when exposed to air. Marked with subectosomal
drainage canals in some specimens.
Skeleton (Fig 11 A, B). Ectosomal skeleton formed
by 100–200 µm thick continuous layer, supported by
choanosomal skeleton and packed mainly with smaller
category of oxeas, in short, ill-deined and criss-crossing
bundles, oriented tangentially- paratangentially, protruding
occasionally through surface in disorganised manner
(Fig. 11A). Choanosomal skeleton, halichondroid to
plumose, formed by ill-deined bundles of larger spicules,
criss-crossing in all directions in the inner region of
choanosome. Skeleton becomes more organised at
Table 8. Comparison of spicule dimensions between specimens of
Halichondria (Halichondria) carotenoidea.
Specimen
NTM
Z.5909
NTM
Z.5225
70
Locality
Darwin Harbour,
Dudley Point
Bynoe Harbour
Oxeas
113.2–388.9µm (263.2±87.1)
x 2.3–12.3µm (7.3±3)
111.7–379.5µm (245.3±80.2)
x 2.3–10.7µm (6.2±2.4)
Halichondria (Halichondria) darwinensis Hooper et al.,
1997
(Fig. 10C)
Halichondria darwinensis Hooper et al., 1997: 49.
Material examined. Specimens as listed in Hooper et
al. (1997). addiTional specimens – Darwin Harbour, NTM
Z.5211.
Remarks. Halichondria darwinensis was described
by Hooper et al. (1997). Additional material, an in situ
Halichondriidae from northern Australia
A
B
C
D
E
F
G
Fig. 10. Halichondria (Halichondria) carotenenoidea sp. nov.: A, specimen in situ at Dudley Point, Darwin Harbour, N.T.; B¸ specimen in
situ at Spencer Point, Bynoe Harbour, N.T., showing oscula distributed along lateral side of branches. C, Halichondria (H.) darwinensis,
specimen in situ at Weed Reef, Darwin Harbour, N.T. Halichondria (H.) phakellioides: D, specimen in situ exposed at the reef lat of East
Arm, Darwin Harbour during the low tide of 18 October 2001; E, specimen in situ at Raragala I. Wessel Is, 20 m depth. F, Halichondria (H.)
microbiana sp. nov, specimen in situ at Lee Point, Darwin Harbour, N.T. G, Topsentia dura, specimen in situ at Nightcliff bommies, Darwin
Harbour, N.T. Photographs: A,D,G – B. Alvarez; B – M. Browne; C,F – H. Nguyen; E – P. Colin.
71
B. Alvarez and J. N. A. Hooper
Table 9. Halichondria (Halichondria) species from the Central Indo-Paciic (after Van Soest et al. 2008 ), with remarks on current taxonomic
allocation, type material and description.
Species
Remarks
Halichondria (Halichondria) bergquistae
Hooper, Cook, Hobbs & Kennedy, 1997
Referred here to Axinyssa (see Remarks of Axinyssa this revision).
Halichondria (Halichondria) darwinensis
Hooper, Cook, Hobbs & Kennedy, 1997
Valid (see under genus Halichondria, this revision).
Unrecognisable from the description; no type material available.
Halichondria (Halichondria) armata
Lindgren, 1897
Halichondria (Halichondria) cartilaginea
(Esper, 1794)
Valid, common species through Indo-Paciic. Always associated to symbiotic green algae
(Lim et al. 2008).
Halichondria (Halichondria) fragilis
Kieschnick, 1896
Unrecognisable from the description; no type material available.
Halichondria (Halichondria) pelliculata
Ridley & Dendy, 1886
Valid, recognisable from the description. Type material not examined. Distinctive shape
and surface characteristics, not comparable to the new species recorded here.
Halichondria (Halichondria) stalagmites
(Hentschel, 1912)
Referred here to Ciocalypta (see remarks of Cioclaypta this revision)
Halichondria (Halichondria) incrustans
Kieschnick, 1896
Unrecognisable from the description; no type material available.
Halichondria (Halichondria) ridleyi
Hooper, Cook, Hobbs & Kennedy, 1997
Referred in this revision to Topsentia (see Remarks of Topsentia this revision)
Halichondria (Halichondria) syringea
Pulitzer-Finali, 1996
Halichondria (Halichondria) vansoesti
Hooper, Cook, Hobbs & Kennedy, 1997
Valid. Type material examined (MSNG 48700). Coalescent istules on a buried base [?].
Detachable ectosome differentiated into a thick layer, with spicules oriented paratangentially, disorganised. Choanosome halichondroid with some ill deined, multispicular,
ascending to surface and supporting ectosome. Oxeas several sizes, 256.4-706.7µm
(539.3±137) x 5.2-19µm (13±3.7); few styles.
Refered here to Ciocalypta (see under genus Ciocalypta, this revision)
photograph (Fig 10C), and spicule measurements (Table 10)
are provided here to complement the original description.
Halichondria darwinensis is very inconspicuous and
represented so far by only three individuals (including
the holotype and the paratype). Its encrusting habit with
small and insubstantial digits makes it inconspicuous and
hard to ind. The re-examination of the available material
and additional measurements of spicules, indicate that the
differences in the thickness of the spicules reported by
Hooper et al. (1997) as a distinctive character for the species,
no longer appear to be signiicant (see Table 10). Both the
length and the thickness of the oxeas are variable, but not
divisible into size classes.
The species seems to have afinities with the genus
Axinyssa. The skeleton is poorly developed, with very little
spongin and relatively less spicule density when compared
to other Halichondria species; the ectosomal skeleton is
quite undifferentiated and the spicules are similar in shape
and dimensions to other Axinyssa species. Therefore, the
assignment of this species to the genus Halichondria is
inconclusive.
Distribution. Halichondria darwinensis is presently
known only from Darwin Harbour. It occurs intertidally
and subtidally to 10 m.
Table 10. Comparison of spicule dimensions among specimens of
Halichondria (Halichondria) darwinensis.
Specimen
NTM Z.3205
(Holotype)
Locality
Darwin
Harbour, East
Point
QM G303252 Darwin
Harbour, East Arm
(Paratype)
NTM Z.5211 Darwin
Harbour, Weed
Reef
Oxeas
272.3–659.6µm
(467.2±112)
x 3.7–13.2µm (8.5±2.5)
307–627.5µm (538.8±90.6)
x 4.3–16µm (10.2±3.2)
386.2–663.7µm (524±68)
x 7–14.9µm (11.8±1.9)
72
Halichondria (Halichondria) phakellioides Dendy &
Frederick, 1924
(Figs 10 D, E, 12)
Halichondria phakellioides Dendy & Frederick, 1924:
498; Burton 1934: 600; Hooper et al. 1997.
Material examined. Specimens as listed in Hooper et al.
(1997). addiTional specimens – Bynoe Harbour, NT: Z.241,
Z.5223 (0M9H2321-P). Darwin Harbour, NT: Z.2026,
Z.4100, (0CDN-8022-F), Z.5219 (0M9H2057-C), Z.5905,
Z.5906, Z.5915, Z.5923, Z.5925, Z.5928, Z.5948, Z.5949,
Z.5950, Z.5965, Z.5970. Parry Shoals, N.T., QM G310137.
Boucat Bay, E of Maningrida, Arnhem Land, N.T.: Z.5623.
Wessel Is: Z.5228 (0M9H2655-C).
Remarks. Halichondria phakellioides was described
by Hooper et al. (1997). Additional material, illustrations
(Figs 10D, E, 12), and spicule measurements (Table 11)
are provided here to complement the original description.
Halichondriidae from northern Australia
A
C
B
Fig. 11. Halichondria (Halichondria) carotenenoidea sp. nov. (NTM Z.5909): A, light microphotograph of tangential section of ectosomal
skeleton showing ill-deined and criss-crossing bundles of spicules; B, light microphotograph of perpendicular section through surface showing
organisation of choanosomal skeleton and part of ectosomal skeleton layer (left upper corner of section); C, diagram of spicules Scale bars:
A, 100 µm; B, 500 µm; C, 50 µm.
depth, 7 August 2003, coll. Nguyen, H. paraTypes.– nTm
Z.1097, Dudley Point Reef, East Point, Darwin, N.T.,
12°25.0001’ S, 130°48.01’ E, 6–7 m depth, 22 December
1982, coll. Hooper, JNA. NTM Z.5908, Moira Reef, Bynoe
Harbour, N.T., 12°30.799’ S, 130°30.527’ E, 5–8 m depth,
25 June 2003, coll. Nguyen, H. addiTional specimens.–
Darwin Harbour NT: NTM, Z.1991. Cobourg Peninsula,
N.T.: NTM Z.131.
Description
Shape (Fig.10F). Cushion-shaped with globular,
subspherical or massive base, sometimes semi-buried
Distribution. Halichondria phakellioides is widely
distributed through the Northwest Australian Shelf and
Sahul Shelf provinces. It is found from the intertidal to
20 m depth.
Halichondria (Halichondria) microbiana sp. nov.
(Figs 10F, 13)
Halichondria stalagmites.–Hooper et al. 1997: 43 [in
part; not Z.2651 and Z.3147 = H. (H.) carotenoidea].
Material examined. HoloType. –NTM Z.5907, Lee
Point, Darwin, N.T., 12°20.538’ S, 130°52.184’ E, 9–12 m
Table 11. Comparison of spicule dimensions among specimens of Halichondria (Halichondria) phakellioides.
Specimen
NTM Z.5223
Locality
Bynoe Harbour
NTM Z.5219
Darwin Harbour
NTM Z.5228
Wessel Is
Oxea type I
170.1–308.4µm (232±30.7)
x 4.1–10.3µm (8±1.5)
154.2–293.2µm (228.6±34.5)
x 5–10.3µm (7.3±1.4)
162.1–313.9µm (242.4±48.6)
x 3.7–7.5µm (5.6±0.9)
73
Oxea type II
363.2–524.6µm (440.6±40.8)
x 11.5–24.4µm (17.9±3.7)
356.7–499.6µm (425.5±38.7)
x 10.5–20.2µm (16.3±3)
326.5–519.3µm (440.5±45.5)
x 10.6–21.1µm (15.6±3)
B. Alvarez and J. N. A. Hooper
A
B
Fig. 12. Halichondria (Halichondria) phakellioides (NTM Z.5228): A, light microphotograph of tangential section showing organisation
of ectosomal skeleton; B, light microphotograph of perpendicular section through surface, showing organisation of choanosomal skeleton.
Scale bars: A-B, 500 µm.
Paciic (Table 7) and those reported in this revision. It is
diagnosed by a unique combination of features (i.e growth
form, organisation of the skeleton, size and composition of
spicules and presence of cyanobacteria in the ectosome and
the choanosome) not found in those species.
All the specimens examined here have a high density
of ilamentous cyanobacteria in the choanosome, generally
concentrated at the subectosomal region, a character shared
with H. (H.) carotenoidea. As mentioned above, this new
species is also very similar in skeletal characteristics to
H.(H.) carotenoidea sp. nov., but it differs in habit and size
of spicules, which are larger and separated in size categories
in H. (H.) microbiana.
Hooper et al. (1997) interpreted this species as
Halichondria stalagmites. However, a thorough
re-examination of the type material of that species indicates
that it is not conspeciic with H. (H.) microbiana (see
description of C. stalagmites above).
Distribution. Halichondria microbiana is relatively
common at Darwin and Bynoe Harbours, and it was also
observed at Wessel Is (Raragala I.) and recorded from
the Northeastern Australian Shelf (Alvarez and Hooper,
unpublished data). It occurs between 6–12 m depth.
Etymology. Named after the symbiotic microorganisms
hosted by the species. The speciic name is intended as a
noun in apposition.
Remarks on Halichondria (Halichondria). The
subgenus Halichondria is large with approx. 95 valid
species (Van Soest et al. 2008). Table 7 lists species
distributed within the Central Indo-Paciic Realm (following
Spalding et al. 2007) which includes the study area and
adjacent areas. Some of those species, in particular the
in substrate, and small tapering erect projections (from
10 mm long and less than 5 mm diameter) at apex. Erect
projections vary in shape, i.e. istulose-like, convoluted,
bifurcating and lobate; generally small 30–70 mm diameter,
10–20 mm thick.
Colour. Orange. Blue-grey, in alcohol.
Consistency and texture. Soft, easily torn, rubbery.
Oscula. Apical and conspicuous on small digits, with
membranous rims.
Surface. Semitransparent at apex of digits in some
specimens.
Skeleton (Fig. 13A,B). Ectosomal skeleton,
halichondroid, formed by a thin (less than 50 µm),
wavy, loose membranous layer, with oxeas tangentially
oriented and supported by choanosomal tracts (Fig. 13A).
Choanosomal skeleton, halichondroid, formed by ill-deined
bundles of larger spicules, criss-crossing and without any
particular orientation connected by short bundles and
single spicules. In subectosomal region, skeleton becoming
cavernous with large lacunae, 100–600 µm diameter
(Fig. 13B) and vague reticulation of pauci-multispicular, illdeined tracts, diverging towards surface or condensed and
running nearly parallel to it, supporting ectosomal skeleton.
High densities of ilamentous cyanobacteria present in both
ectosome and choanosome, especially near surface. Spongin
or collagen scarce.
Spicules (Fig. 13C; Table 12). Oxeas, hastate, straight,
possibly in two size categories, larger and thicker (249–587
x 5–14 µm) and smaller and thinner (88–217 x 3–7 µm).
Style modiications slightly common within larger category.
Remarks. We compared Halichondria microbiana
with the valid species recorded for the Central West
Table 12. Comparison of spicule dimensions between specimens of Halichondria (Halichondria) microbiana.
Specimen
NTM Z.5907
Locality
Darwin Harbour, NT
NTM Z.5908
Bynoe Harbour, NT
Oxea type I
87.9–203µm (126.5±24.8)
x 3.3–7.1µm (5±0.9)
91.3–217µm (137.8±32)
x 2.9–7.2µm (5.1±1)
74
Oxea type II
249.2–587µm (482.9±75.1)
x 6.3–14.2µm (8.9±1.8)
282.5–572.1µm (446.6±93.5)
x 5.2–13.2µm (8.8±2)
Halichondriidae from northern Australia
A
C
B
Fig. 13. Halichondria (Halichondria) microbiana sp. nov. (NTM Z.5908): A, light microphotograph of tangential section of ectosomal
skeleton, showing oxeas on membranous thin layer; B, light microphotograph of perpendicular section through surface, showing organisation
of choanosomal skeleton and large lacuna at subectosomal level (left upper corner of section); C, diagram of spicules. Scale bars: A, 200
µm; B, 500 µm; C, 50 µm.
ones described by Kieschnick (1896) and Lindgren (1897),
are unrecognisable and their type material has never been
relocated. Other species reported for the area have been
allocated to other genera as result of this revision, thus
Halichondria (Halichondria) is represented within this
realm by nine species including the two new species
described above.
The subgenus Eumastia Schmidt, 1870 is not represented
in the study area at all and it is reserved for Halichondrialike species from high latitudes (Erpenbeck & Van Soest
2002).
Genus Hymeniacidon Bowerbank, 1858
Gender: feminine. Type species, by subsequent
designation of Bowerbank (1864), Hymeniacidon caruncula
Bowerbank, 1958. Recent, Tenby, Wales.
75
Hymeniacidon gracilis (Henstschel, 1912)
(Fig. 14)
Stylotella digitata gracilis Henschel, 1912: 356.
Hymeniacidon gracilis. – Hooper et al. 1997.
Material examined. As listed by Hooper et al. (1997).
Remarks. The species was well described by Hooper
et al. (1997) and elevated to full species rank in the genus
Hymeniacidon. We provide a new illustration of the type
material (Fig. 14) and additional spicule measurements
(Table 13) to complement the description given by Hooper
et al. (1997).
Only three specimens are recorded for the study area; no
new material was located in recent collections.
Distribution. As recorded by Hooper et al. (1997).
Remarks on the genus Hymeniacidon. Hymeniacidon
gracilis is the only valid species of the genus from the
study area. Other species of Hymeniacidon recorded for the
B. Alvarez and J. N. A. Hooper
A
B
Fig. 14. Hymeniacidon gracilis (Synype SMF 970). A, light microphotograph of perpendicular section through surface, showing organisation
of choanosomal skeleton; B, diagram of spicules. Scale bars: A, 500 µm; B, 50 µm.
these style modiications are common among the genus.
Population genetics and additional morphometric analyses
might reveal whether these style modiications of Stylissa
are reliable characters for the separation of species.
Sahul Shelf Province and adjacent areas are H. vernonensis
Hooper et al., 1997 and H. laccida Pulitzer-Finali, 1996.
The material described under H. vernonensis by Hooper et
al.(1997) was revised and it does not agree with the current
concept of the genus. The species is formally transferred
here to the dictyonellid genus Stylissa. As admitted by
Hooper et al. (1997), is very similar to Stylissa labelliformis
but S. vernonensis includes distinctive styles, curved at the
centre, sinuous or rhabdose and frequent anisoxeas with
one telescoped point. It should be noted however, that
these spicule modiications are common among species
of Stylissa and therefore they are not reliable for the
delimitation of species. Both species are similar in growth
form, surface characteristics and skeletal organisation. The
choanosomal skeleton of S. vernonensis however, is nearly
halichondroid with only vague tracts of spicules and with a
much higher spicule density and no spongin ibres (whereas
in S. labelliformis is vaguely plumo-reticulated and with
well developed spongin ibres).
The type specimen of Hymeniacidon laccida (MSNG
48703) from Laing Is., Papua New Guinea, was re-examined
and it does not correspond to the genus Hymeniacidon.
The species belongs also to the dictyonellid genus
Stylissa and is likely to be conspeciic with Stylissa massa
(Carter, 1887). The specimen examined however, includes
distinctive subtylostyles transitional to strongyles with
tylote modiications. As is the case with S. vernonensis,
Genus Topsentia
Gender feminine. Type species, by original designation,
Anisoxya glabra Topsent, 1898. Recent, Azores Is.
Topsentia dura (Lindgren, 1897)
(Figs 10G, 15)
Halichondria dura Lindgren, 1897 : 480.
Topsentia dura.– Hooper et al. 1997: 14.
Material examined. As listed by Hooper et al. (1997).
addiTional maTerial.– Darwin Harbour, N.T.: Z.5209,
Z.5233. Wessel Is: Z.5234.
Remarks. Hooper et al. (1997) assigned material from
the Beagle Gulf to this species under the genus Topsentia.
Additional specimens from recent collections agree also
with this material and are assigned to this species. Further
illustrations (Figs 10G, 15) and spicule measurements
(Table 14) are provided here to complement that description.
The description agrees with the current concept of Topsentia,
however it remains inconclusive whether the material from
the Beagle Gulf is conspeciic with Lindgren’s species from
Indonesia as the type was not examined. Examination of
additional specimens from Indonesia (Alvarez & De Voogd
unpublished data) and a re-description of the type might
provide additional evidence to conirm if these populations
belong in the same species.
Distribution. Indonesia [?], Darwin Harbour and Wessel
Is. Topsentia dura is occurs the intertidal zone to 25 m depth.
Remarks on Topsentia. Hooper et al. (1997)
described material under the Red Sea species Topsentia
halichondrioides (Dendy, 1905) that seems very similar
Table 13. Comparison of spicule dimensions between specimens of
Hymeniacidon gracilis.
Specimen
SMF 970
Locality
Indonesia
NTM Z.883 Darwin
Harbour
Styles
220.5–261.1µm (238.4±9.6)
x 3.3–8.9µm (6.1±1.5) [25]
243.1–279.1µm (265.6±11.1)
x 3–7.4µm (5.1±1.1)
76
Halichondriidae from northern Australia
A
B
C
D
E
F
Fig. 15. Topsentia dura (NTM Z.5209): A, light microphotograph of perpendicular section through surface, showing organisation of choanosomal
skeleton and palisade of erect oxeas at surface; B, diagram of spicules. Topsentia halichondrioides (QM G303442); C, light microphotograph
of perpendicular section through surface, showing organisation of choanosomal skeleton and oxeas oriented perpendicularly at surface level;
D, diagram of spicules. Topsentia ridleyi (QM G303309, Holotype); E, light microphotograph of perpendicular section through surface,
showing organisation of choanosomal skeleton; F, diagram of spicules. Scale bars: A,C,E, 500 µm; B,D,F, 100 µm.
to T. dura (Fig. 15B,C). These species are massive, of
hard consistency with skeletons made of a confused mass
of oxeas of similar dimensions, not clearly differentiated
into size classes (see Table 14) and differing only in
colouration and shape of oscules (i.e. volcano-shaped in
77
T. halichondrioides, and sunken and small in T. dura).
Halichondria ridleyi Hooper et al., 1997 is referred here to
the genus Topsentia (comb. nov.) and it is also very similar
to T. dura (Fig. 15E,F). It differs from T. dura in having
some surface istules and processes.
B. Alvarez and J. N. A. Hooper
Table 14. Comparison of spicule dimensions among specimens assigned to Central Paciic species of Topsentia.
Species
Topsentia ridleyi
Specimen
Locality
QM G303309 Darwin Harbour
Topsentia ridleyi
NTM Z.3262
Cobourg Peninsula
Topsentia halichondrioides
G303442
Bynoe Harbour
Topsentia halichondrioides
NTM Z.5233
East Point
Topsentia dura
NTM Z.5234
Wessel Is
Topsentia dura
NTM Z.5209
Darwin Harbour
Topsentia dura
NTM Z.3178
Darwin Harbour
Topsentia dura
NTM Z.1442
Gunn Point
Topsentia indica
SMF 997
Aru Is, Indonesia
Oxea type I
142.3–273.5µm (207.7±39.4)
x 3.3–7.6µm (5.4±1.2)
160.6–331.2µm (257.4±50.3)
x 4.2–12.3µm (8.2±2.3)
111.4–165.9µm (142.3±13.8)
x 5.9–8.9µm (7.2±0.9)
142.4–333.5µm (220.1±47.2)
x 3.7–10.3µm (7.3±1.7)
151.2–399.7µm (248.3±61.9)
x 3–11.7µm (7.6±2.3)
189.4–357.5µm (269.9±41.9)
x 4.1–9.7µm (7.1±1.4)
189.6–387.1µm (254.5±52.5)
x 5.5–12.5µm (7.8±2)
An additional species of Topsentia recorded from Aru
Is, Indonesia, is T. indica Hentschel, 1912 (syntype SMF
995 and 997, examined).
The differentiation of these species using traditional
morphological characters is extremely subjective. Molecular
and morphometric studies of local populations might
contribute to a better understanding of the concept of this
species.
Other species of Topsentia recorded for the Sahul
Province and adjacent areas that are better placed elsewhere
include Topsentia maculosa Pulitzer-Finali, 1996 from
Papua New Guinea (it belongs in Amorphinopsis, see above)
and Topsentia plurisclera Pulitzer-Finali, 1996 (holotype,
MSNG 48702, examined) is a species of Petrosia.
DISCUSSION
This revision of species of Halichondriidae from northern
Australia recognises a total of 15 species belonging to the
genera Amorphinopsis, Axinyssa, Ciocalypta, Halichondria
(Halichondria), Hymeniacidon and Topsentia. Other genera
of the family (i.e. Epipolasis, Laminospongia, Vosmaeria,
Ciocalapata and Spongosorites are not represented in the
Sahul Shelf Province. Epipolasis and Spongosorites are
however represented in the Northeast Australian Shelf
(Alvarez and Hooper, unpublished data).
Of the species reported in this revision, Axinyssa
bergquistae, Ciocalypta vansoesti and the two new species
Halichondria (Halichondria) carotenoidea and H (H.)
microbiana are so far known only from northern Australian
waters. The rest of the species have extralimital distributions
through the Central Indo-Paciic realm. Axinyssa mertoni
(Hentschel, 1912) in its new generic combination is recorded
for northern Australia and it represents a new record for the
study area.
78
Oxea type II
335.6–576.7µm (443.9±52.7)
x 6.7–16.6µm (12±2.6)
503–848.1µm (653.8±83.5)
x 13.4–31.5µm (20.9±4.6)
421.6–716.5µm (555.3±62.7)
x 14–24.4µm (19.3±2.6)
487.6–914.9µm (711.2±99.1)
x 13–30µm (18.9±4.4)
348.6–568.1µm (448.9±57.8)
x 11–23.7µm (16.8±3.2)
373.9–656.5µm (477.9±79.2)
x 10–23.9µm (15±3.2)
352.5–525.1µm (441.4±46.2)
x 9.9–18.5µm (13.7±2.4)
341.2–611.2µm (482.6±69.2)
x 8.3–22.7µm (15.6±3.4)
493.7–1078.1µm (734.4±125)
x 11.1–39.4µm (21.7±5.5)
As in other members of the order Halichondrida,
particularly in Axinellidae and Dictyonellidae, species
within and across all genera of Halichondriidae are
extremely dificult to delimit. This problem is demonstrated
by the large number of misidentiications in previous studies
due to the lack of adequate generic deinitions and also
to the poor understanding of the importance, or indeed
relevance, of some of the alleged pivotal characters that
currently differentiate both species-groups and genera
within the Halichondriidae. The additional information
obtained from the collection of new material plus the revised
generic deinitions of the family (Erpenbeck & Van Soest
2002) allowed us to clarify the concept of halichondriid
species of northern Australia and to allocate them to more
appropriate genera.
Nevertheless, differentiating species within
Halichondriidae continues to be ambiguous based solely
on the present limited suite of accepted morphological
characters, with a number of them shared among species
and even genera. For example, Ciocalypta heterostyla,
C.vansoesti, and Axinyssa mertoni are species with istulelike growth form, nearly indistinguishable in the ield
(Fig 1). However, they all are easily diagnosed based on
skeletal characteristics. This suggests that the growth form
of these species in particular, might be an adaptation to the
habit where they occur (i.e. soft and muddy sediments).
Amorphinopsis foetida and A. maculosa are also very similar
in habit and in their skeletal characteristics but they can
be distinguished by the predominance of oxeas and styles.
Separation of species based on the dominance of styles or
oxeas however might be debatable, as this could possibly be
related to intraspeciic variation as seen in some species of
Axinellidae (Alvarez et al. 1998; Alvarez & Hooper 2009).
Spicule morphometrics (i.e. size variation of spicules)
within Halichondriidae seems to be a useful tool for the
Halichondriidae from northern Australia
ACKNOWLEDGEMENTS
differentiation of species. Ciocalypta stalagmites, for
example, is distinguished from the other species by the
distinctive and nearly constant size of two categories of
oxeas. Similarly, Halichondria (H.) microbiana can be
distinguished from H. (H.) carotenoidea by the size of the
larger category of oxeas. On the other hand, all the Axinyssa
species we have studied have always a mixture of oxeas in a
large size range, thus spicule sizes is not a useful character
for the distinction of species within this genus.
These examples indicate that the characters currently
used to separate genera and species within the family are
extremely homoplastic and suggest that genera and species
within Halichondriidae might be non-monophyletic. The
study of local populations using both morphological and
genetic methods will help to clarify whether these taxa are
monophyletic.
This study represents the inal contribution to the present
taxonomic revision of the order Halichondrida of northern
Australia, restricted to the marine province identiied as the
Sahul Shelf in the classiication of Spalding et al. (2007).
The result of this and previous studies (Alvarez & Hooper
2009, 2010) indicates that the group is represented in the
study area mainly by the families Axinellidae (Alvarez &
Hooper 2009), Dictyonellidae (Alvarez & Hooper 2010)
and Halichondriidae (this revision).
One additional family, the Heteroxyidae (formerly
Desmoxyidae see Van Soest & Hooper 2005), is represented
in the area by two very common species: Myrmekioderma
granulatum (Esper, 1830), which is documented and
illustrated by Hooper et al. (1997); and Higginsia mixta
(Hentschel, 1912). The type of H. mixta (SMF 968) was
examined and it agrees with material deposited at the
collections of QM and NTM and recorded for the area of
study. The ectosomal skeleton of the specimens studied
have a relatively thick tangential crust formed by a dense
mass of spined microxeas and interrupted by disorganised
brushes of thin raphidiform oxeas and extra long stylesstrongyles (mostly broken in the preparations) projecting
through surface. The similarities of this type of skeletal
organisation with raspailiid genera such as Ceratopsion are
remarkable and worth further investigation. Phylogenetic
afinities based on molecular data (Erpenbeck et al. 2005)
indicated that members of Heteroxyidae (i.e. Didiscus and
Myrmekioderma) are closely related to the axinellid genera
Reniochalina and Ptilocaulis. The position of these genera
within Axinellidae (Halichondrida) and its relationships
based on molecular data with other raspailiid genera such
as Axechina has already been discussed (Alvarez 2009 and
references within). Sequencing data of additional genera
currently allocated to Heteroxyidae, and in particular of the
species represented in the Sahul Shelf province will help to
clarify these relationships and classiication.
The family Bubaridae is the only family of Halichondrida
not represented in the area of study.
This work was funded by an Australian Biological
Research Studies Research grant (No 205-10) and
by the ‘Collection and Taxonomy of Shallow Water
Marine Organisms’ program for the U.S. National
Cancer Institute (Contract N02-CM-27003 and Contract
N02CO-2009-00012) subcontracted to the MAGNT through
CRRF.
We thank the following people: Michael Browne and Huy
Nguyen, for their invaluable assistance during 2002–2004
MAGNT ield collections; Dr Pat Colin, CRRF, and Don
DeMaria, for their assistance and photographic work during
the MAGNT ield collections in the 2004 Wessel Is; Terry
Yumbuluy, Wessel Is, for allowing collections of sponges
in his sea area; Merrick Ekins, QM, for his assistance
interrogating QM database and making specimens available
for study and Dr Kathryn Hall for substantial re-checking
voucher specimens housed in the collections of the QM
to conirm or refute previous species diagnoses. Drs Rob
W.M. Van Soest (ZMA) and Nicole De Voogd are thanked
for their input and valuable discussions on Indo-Paciic
halichondrid sponges. Mr Swee-Cheng Lim is thanked for
his assistance locating type material at MSNG. Drs Richard
Willan and Chris Glasby (MAGNT) are thanked for their
continuous advice and suggestions during the preparation
of this manuscript. And lastly, we thank the two referees for
their valuable suggestions.
REFERENCES
Alvarez, B. & Hooper, J.N.A. 2009. Taxonomic revision of the
order Halichondrida (Porifera: Demospongiae) from northern
Australia. Family Axinellidae. The Beagle, Records of the
Museums and Art Galleries of the Northern Territory 25: 17–42.
Alvarez, B. & Hooper, J.N.A. 2010. Taxonomic revision of the
order Halichondrida (Porifera: Demospongiae) from northern
Australia. Family Dictyonellidae. The Beagle, Records of the
Museums and Art Galleries of the Northern Territory 26: 13–36.
Alvarez, B., Van Soest, R.W.M. & Rützler, K. 1998. A revision of
the species of Axinellidae (Porifera: Demospongiae) in the
Central-West Atlantic region. Smithsonian Contributions to
Zoology 598: 1–47.
Bergquist, P.R. 1965. The sponges of Micronesia, Part 1. The Palau
Archipelago. Paciic Science 19: 123–204.
Bowerbank, J.S. 1862. On the anatomy and physiology of the
Spongiadae. Part III: On the generic characters, the speciic
characters and on the method of examination. Philosophical
Transactions of the Royal Society of London 152: 1087–1135,
pls. 72–74.
Bowerbank, J.S. 1864. A monograph of the British Spongiadae.
Volume 1. The Ray Society, London.
Bowerbank, J.S. 1873. Contributions to a general history of the
Spongiadae. Part 4. Proceedings of the Zoological Society of
London for the year 1873: 3–25 pls. 1–4.
Burton, M. 1928. Report on some deep-sea sponges from the Indian
Museum collected by the R.I.M.S. ‘Investigator’. Part II.
Tetraxonida (concluded) and Euceratosa. Records of the Indian
Museum, Calcutta 30: 109–138, pls.1–2.
79
B. Alvarez and J. N. A. Hooper
Kieschnick, O. 1896. Silicispongiae von Ternate nach den
sammlungen von Herrn Prof. Dr. W. Kükenthal. Zoologischer
Anzeiger 19: 526–534.
Lendenfeld, R.V. 1897. Spongien von Sansibar. Abhandlungen
Herausgegeben von der Senckenbergischen Naturforschenden
Gesellschaft 21: 93–133, pls 9–10.
Lim, S-C., De Voogd, N. & Tan, K-S. 2008. A guide to sponges of
Singapore. Science Centre Singapore: Singapore.
Lindgren, N.G. 1897. Beitrag zur kenntniss der spongienfauna
des Malaiischen Archipels und der Chinesischen Meere.
Zoologischer Anzeiger 20: 480–487.
Lindgren, N.G. 1898. Beitrag zur kenntniss der spongienfauna
des Malayischen Archipels und der Chinesischen Meere.
Zoologische Jahrbücher. Abteilung für Systematik, Geographie
und Biologie der Thiere 11: 283–378, pls 17–20.
Pulitzer-Finali, G. 1996. Sponges from the Bismarck Sea. Bolletino
dei Musei e degli Istituti Biologici dell’Università di Genova
60–61: 101–138.
Ridley, S.O. 1884. Spongiida. Pp. 366–684. Report on the Zoological
Collections made in the Indo-Paciic Ocean during the Voyage
of H.M.S. ‘Alert’ 1881-2. British Museum, Natural History:
London.
Ridley, S.O. & Dendy, A. 1886. Preliminary report on the Monaxonida
collected by H.M.S ‘Challenger’. Part I. The Annals and
Magazine of Natural History (Series 5) 18: 325–351.
Sollas, I.B.J. 1902. On the sponges collected during the Skeat
expedition in the Malay Peninsula (1899-1900). Proceedings
of the Zoological Society of London 2: 210–221.
Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña,
Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A.,
Lourie, S.A., Martin, K.D., McManus, E., Molnar, J., Recchia,
C.A. & Robertson, J. 2007. Marine ecoregions of the world:
a bioregionalization of coastal and shelf areas. Bioscience 57:
573–583.
Thiele, J. 1899. Studien über Paziische Spongien. II. Zoologica 24:
1–33, pls. 1–5.
Thiele, J. 1900. Kieselschwämme von Ternate. I. Abhandlungen der
Senckenbergischen Naturforschenden Gesellschaft 25: 19–80.
Van Soest, R.W.M. 1991. Demospongiae higher taxa classiication
re-examined. Pp. 54–71. In: Reitner, J. & Keupp, H. (eds) Fossil
and Recent Sponges. Springer-Verlag: Berlin.
Van Soest, R.W.M, Boury-Esnault, N., Hooper, J.N.A., Rützler,
K, de Voogd, N.J., Alvarez de Glasby, B., Hajdu, E., Pisera,
A.B., Manconi, R., Schoenberg, C., Janussen, D., Tabachnick,
K.R., Klautau, M., Picton, B., Kelly, M., Vacelet, J. 2008.
World Porifera database. Available online at http://www.
marinespecies.org/porifera. Last consulted 27 July 2011.
Van Soest, R.W.M., Díaz, M.C. & Pomponi, S.A. 1990. Phylogenetic
classiication of the halichondrids (Porifera, Demospongiae).
Beaufortia 40: 15–62.
Van Soest, R.W.M. & Hooper, J.N.A. 2005. Resurrection of Desmoxya
(Porifera: Halichondrida), with the description of a new species
from Rockall Bank bathyal coral reefs, North Atlantic. Journal
of the Marine Biological Association of the United Kingdom
85: 1367–1371.
Wilson, H.V. 1925. Silicious and horny sponges collected by the U.S.
Fisheries Steamer ‘Albatross’ during the Philippine expedition,
1907-10. Bulletin of the United States National Museum 2:
273–532.
Burton, M. 1934. Sponges. Scientiic Reports of the Great Barrier
Reef Expedition 1928-29 4: 513–621, pls.1–2.
Burton, M. 1959. Sponges. Scientiic Reports of the John Murray
Expedition 1933-34 10: 151–281.
Carter, H.J. 1887. Report on the marine sponges, chiely from King
Island, in the Mergui Archipelago, collected for the Trustees
of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S.,
Superintendent of the Museum. Journal of the Linnean Society
of London, Zoology 21: 61–84, pls 5–7.
De Laubenfels, M.W. 1936. A discussion of the sponge fauna of the
Dry Tortugas in particular, and the West Indies in general, with
material for a revision of the families and orders of the Porifera.
Papers from the Tortugas Laboratory 30: 1–225, pls. 1–22.
De Laubenfels, M.W. 1954. The Sponges of the West-Central Paciic.
Oregon State Monographs, Studies Zoology: 1–306, pls. 1–12.
Dendy, A. 1889. Report on a second collection of sponges from the
Gulf of Manaar. Annals and Magazine of Natural History 3:
73–99, pls. 3–5.
Dendy, A. 1905. Report on the sponges collected by Professor
Herdman, at Ceylon, in 1902. Pp. 57–246, pls 1–16. In:
Herdman, W.A. (ed.) Report to the Government of Ceylon
on the pearl oyster Fisheries of the Gulf of Manaar. Royal
Society: London.
Dendy, A. 1922. Report on the Sigmatotetraxonida collected by
H.M.S. ‘Sealark’ in the Indian Ocean. Transactions of the
Linnean Society of London 18: 1–164, pls. 1–18.
Dendy, A. & Frederick, L.M. 1924. On a collection of sponges from
the Abrolhos Islands, Western Australia. Journal of the Linnean
Society of London, Zoology 35: 477–519 pls. 25–26.
Erpenbeck, D., Breeuwer, J. & Van Soest, R.W.M. 2005. Implications
from a 28S rRNA gene fragment for the phylogenetic
relationships of halichondrid sponges (Porifera: Demospongiae).
Journal of Zoological Systematics and Evolutionary Research
43: 93–99.
Erpenbeck, D. & Van Soest, R.W.M. 2002. Family Halichondriidae.
Pp. 773–786. In: Hooper, J.N.A. & Van Soest, R.R.M. (eds)
Systema Porifera. A guide to the supraspeciic classiication of
the phylum Porifera. Plenum Press: New York.
Hentschel, E. 1912. Kiesel-und hornschwämme der Aru und KeiInseln. Abhandlungen Senckenbergiana Naturforschende
Gessellschaft: Hamburg.
Hooper, J.N.A. 2005. Porifera. Australian Faunal Directory.
Australian Biological Resources Study, Canberra. Last
consulted 18 May 2011. http://www.environment.gov.au/
biodiversity/abrs/online-resources/fauna/afd/taxa/.
Hooper, J.N.A. & Bergquist, P.R. 1992. Cymbastela, a new genus
of lamellate coral reef sponges. Memoirs of the Queensland
Museum 32: 99–137.
Hooper, J.N.A., Capon, R.J., Keenan, C.P., Parry, D.L. & Smit, N. 1992.
Chemotaxonomy of marine sponges: families Microcionidae,
Raspailiidae and Axinellidae, and their relationships with other
families in the order Poecilosclerida and Axinellida (Porifera:
Demospongiae). Invertebrate Taxonomy 6: 261–301.
Hooper, J.N.A., Cook, S.D., Hobbs, L.J., Hooper, L.G. &
Kennedy, J.A. 1997. Australian Halichondriidae (Porifera:
Demospongiae): I. Species from the Beagle Gulf Marine Park.
Pp. 1–65. In: Hanley, J.R., Caswell, G., Megirian, D. & Larson,
H.K. (eds) The marine lora and fauna of Darwin Harbour,
Northern Territory, Australia. Museums and Art Galleries of
the Northern Territory: Darwin.
Hooper, J.N.A. & Wiedenmayer, F. 1994. Porifera. Pp. 1–624. In:
Wells, A. (ed.) Zoological Catalogue of Australia. Volume 12.
Pp 1–624. CSIRO: Melbourne.
Accepted 4 November 2011
80
Halichondriidae from northern Australia
APPENDIX I
G300854
G301034
G303252
G303287
G303309
G303351
G303442
G303450
G303524
G303541
G303558
G303560
G303561
G303595
G303658
G303677
G310137
G310170
G313543
G313572
G313577
G314246
G314247
G314255
G314267
G315205
G315207
G320819
G320904
Collection and locality data of material examined in the collections of QM and NTM
QM material
Gulf of Carpentaria, northern central region, QLD, 9°36.0001’ S, 136°6.01’ E, 52 m , 23 Nov 1991, coll. Cook, SD. on
CSIRO RV Southern Surveyor
SW Vrilya Point, SW, Gulf of Carpentaria, QLD, 11°29.0167’ S, 142°55.09’ E, 18 m , 1 Dec 1991, coll. Cook, SD. on
CSIRO RV Southern Surveyor
South Shell I., reef N of boatramp, East Arm, Darwin Harbour, NT, 12°29.1334’ S, 130°53.09’ E, 0 m , 19 Sep 1993,
coll. Hooper, JNA & Hobbs, L.J.
South Shell I., reef N of boatramp, East Arm, Darwin Harbour, NT, 12°29.1334’ S, 130°53.09’ E, 0 m, 19 Sep 1993,
coll. Hooper, JNA & Hobbs, LJ
Dudley Point Reef, East Point, Darwin, NT, 12°25.05’ S, 130°49.01’ E, 0 m , 20 Sep 1993, coll. Hooper, JNA & Hobbs,
L.J.
East Point Bommies, Darwin Harbour, NT, 12°24.0834’ S, 130°48.14’ E, 10 m , 23 Sep 1993, coll. Hooper, JNA &
Hobbs, LJ
Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 m , 26 Sep 1993, coll. Hooper, JNA &
Hobbs, LJ
Fish Reef, west side, Bynoe Harbour, NT, 12°26.0167’ S, 130°26.09’ E, 11 m , 26 Sep 1993, coll. Hooper, JNA &
Hobbs, LJ
Duyfken Point, W Gulf of Carpentaria, QLD, 12°41.0501’ S, 141°3.01’ E, 42 m, 11 Nov 1993, coll. Cook, SD. &
Kennedy,J. on CSIRO RV. Southern Surveyor
Vernon Is, W of South West Vernon I., NT, 12°6.15’ S, 131°4.14’ E, 13 m, 10 Oct 1993.
Cape Hotham, NW of cape, NT, 12°1.05’ S, 131°13.16’ E, 34 m, 9 Oct 1993.
Bynoe Harbour, 2 nmls E Fish Reef, NT, 12°24.1334’ S, 130°28.16’ E, 17 m, 6 Oct 1993, coll. CCNT Ocean Rescue
2000 Program
Shoal Bay, outer region of bay, NT, 12°6.15’ S, 130°49.16’ E, 18 m, 12 Oct 1993, coll. CCNT Ocean Rescue 2000
Program
Vernon Is, W of Knight Reef, NT, 12°1.0334’ S, 131°3.16’ E, 22 m, 11 Oct 1993.
Vernon Is, N marsh Shoal, NT, 12°07.0001’ S, 130°56.1’ E, 16 m, 11 Oct 1993, coll. CCNT stn. 138. Dredge
Shoal Bay, middle of bay, NT, 12°13.0167’ S, 130°56’ E, 17 m, 12 Oct 1993, coll. CCNT Ocean Rescue 2000 Program
Parry Shoals 35nm W Bathurst I., NT, 11°7.0321’ S, 129°25.9’ E, 16 m, 12 Aug 1987, coll. mussig, AM and NCI team
Darwin Harbour, NT, 12°15.1834’ S., 130°29.11’ E., 9 m depth, 17 August 1987, coll. mussig, AM and NCI team
N Bathurst I., Timor Sea, NT, 11°13.98’ S, 130°34.21’ E, 41.2 m, 5 Oct 1997, coll. Cook, SD. on RV Southern Surveyor
SW Groote Eylandt, NT, 14°25.0801’ S, 135°58.51’ E, 20.3 m, 13 Oct 1997, coll. Cook, SD. on RV Southern Surveyor
SW Groote Eylandt, NT, 14°20.22’ S, 136°34.98’ E, 19.6 m, 14 Oct 1997, coll. Cook, SD. on RV Southern Surveyor
N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136°18.13’ E, 20 m, 27 Sep 1998, coll. Leys, SP. on RV
Southern Surveyor
N Groote Eylandt, Gulf of Carpentaria, NT, 13°32.2801’ S, 136°18.13’ E, 21.7 m, 27 Sep 1998, coll. Leys, SP. on RV
Southern Surveyor
W of Groote Eylandt, Gulf of Carpentaria, NT, 14°8’ S, 136°8’ E, 13 m, 6 Oct 1998, coll. Leys, SP. on RV Southern
Surveyor
SW of Groote Eylandt, Gulf of Carpentaria, QLD, 14°20’ S, 136°2’ E, 22.1 m, 6 Oct 1998, coll. Leys, SP. on RV
Southern Surveyor
W Groote Eylandt, Gulf of Carpentaria, NT, 14°8.5667’ S, 136°16.51’ E, 21.4 m, 13 Oct 1998, coll. Wassenberg T
SW of Groote Eylandt, Gulf of Carpentaria, NT, 14°22.5’ S, 136°9.12’ E, 22.2 m, 13 Oct 1998, coll. Wassenberg T
Gulf of Carpentaria, QLD, 15°20.037’ S, 140°19.84’ E, 28 m, 24 may 2003, coll. Bartlett C, Cook S on RV Southern
Surveyor 2380403 CSIRO “Effects of Trawling”
Gulf of Carpentaria, QLD, 15°20.037’ S, 140°19.84’ E, 28 m, 11 mar 2003, coll. Bartlett C, Cook S on RV Southern
Surveyor 2380403 CSIRO “Effects of Trawling”
81
B. Alvarez and J. N. A. Hooper
APPENDIX I (continued)
Z.84
Z.131
Z.194
Z.241
Z. 592
Z 712
Z.919
Z.934
Z.941
Z.945
Z.986
Z.987
Z.1097
Z.1358
Z.1391
Z.1395
Z.1442
Z.1979
Z.1991
Z.2018
Z.2026
Z.2086
Z.2215
Z.2245
Z.2648
Z.2651
Z.2697
Z.3106
Z.3133
Z.3147
Z.3178
Z.3195
Z.3205
Z.3262
Z.3920
Z.4085
Z.4093
Z.4100
Z.4122
Z.4123
Z.4125
Z.4451
Collection and locality data of material examined in the collections of QM and NTM
NTM material
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.5001’S, 132°2’E, 18 Oct 1981, coll. Hooper, JNA &
Alderslade, PN
Sandy I. No.2, Cobourg Peninsula, NT, 11°5.5001’S, 132°17’E, 10 m, 21 Oct 1981, coll. Hooper, JNA & Alderslade, PN
Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, 0–0.5 m, 13 Sep 1981, coll. Hooper, JNA and
party
Indian I., Bynoe Harbour, NT, 12°35’S, 130°33.01’E, 3 m, 18 Nov 1981, coll. Byers,P.,F.V. Skeleton
Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5’S., 132°10.51’E., 3 m depth, 4 May 1982, coll. Hooper,
JNA
N Adele I.,Collier Bay, NW Shelf, WA 15°58.0167’S., 122°39.07’E., 59 m depth, 21 April 1982, coll. R.V.SPRIGHTLY,
dredge.
Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, 10 m, 31 Aug 1982, coll. Hooper, JNA
East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA
East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA
East Point Reef, East Point, Darwin, NT, 12°24.05’S, 130°48.01’E, 12 m, 13 Sep 1982, coll. Hooper, JNA
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 26 Oct 1982, coll. Hooper, JNA
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 26 Oct 1982, coll. Hooper, JNA
Dudley Point Reef, East Point, Darwin, NT, 12°25.0001’S, 130°48.01’E, m, 22 Dec 1982, coll. Hooper, JNA
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3’S, 132°3.71’E, .5–6 m, 16 May 1983, coll. Hooper, JNA
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3’S, 132°3.71’E, 6 m, 17 May 1983, coll. Hooper, JNA
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°10.4’S, 132°2.8’E, 2 m, 19 May 1983, coll. Hooper, JNA
Blue Hole, Gunn Point, NT, 12°9.0001’S, 131°0’E, 25 m, 19 Aug 1983, coll. Alderslade, PN
West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party
West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party
West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E, m, 11 May 1984, coll. Hooper, JNA and party
West side of Weed Reef, Darwin, NT, 12°29.2001’S, 130°47.1’E , 11 May 1984, coll. Hooper, JNA and party
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 20 Jul 1984, coll. Hooper, JNA
Vestey’s Beach, Bullocky Point, Darwin, NT, 12°26.2’S, 130°49.89’E 21 Jan 1985, coll. Hooper, JNA
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 10 m, 12 Apr 1985, coll. Hood, C and party
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, m, 3 Apr 1986, coll. Hooper, JNA and party
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 3 Apr 1986, coll. Hooper, JNA and party
Parry Shoals, Arafura Sea, NT, 11°12.5167’S, 129°42.07’E, 20 m, 15 Aug 1987, coll. Mussig, AM and NCI team
Parry Shoals, Arafura Sea, NT, 11°11.4’S., 129°43.01’E., 18 m depth, 13 August 1987, coll. Mussig, AM and NCI team
Parry Shoals, Arafura Sea, NT, 11°12.5167’S, 129°42.07’E, 16 m, 15 Aug 1987, coll. Mussig, AM and NCI team
East Point Reef, East Point, Darwin, NT, 12°29.5’S, 130°48.01’E, 0.5 m, 10 Sep 1987, coll. Smit, N
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 9 m, 16 Sep 1987, coll. Smit, N
Dudley Point Reef, East Point, Darwin, NT, 12°24.5’S, 130°48.01’E, 25 Sep 1987, coll. Smit, N
Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5’S, 132°10.51’E, 11 Sep 1986, coll. Hooper, JNA &
Johnson,C
Cumberland Strait, NE bay, Wessel Is, Gove Peninsula, NT, 11°26.8’S, 136°30.2’E, 13 m, 14 Nov 1990, coll. Hooper,
JNA
Near Boat Ramp, East Arm Port, Darwin, NT, Australia, 12º29.8’S, 130º53.5’E, intertidal, 20 September 2001, coll. B.
Glasby & party, by hand
Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 20 September 2001, coll. B. Glasby &
party, by hand
Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 18 October 2001, coll. B. Glasby &
party, by hand
Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 18 October 2001, coll. B. Glasby &
party, by hand
Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 19 October 2001, coll. B. Glasby &
party, by hand
Near Boat Ramp, East Arm Port, Darwin, NT, 12º29.8’S, 130º53.5’E, intertidal, 19 October 2001, coll. B. Glasby &
party, by hand
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.103’S, 130°47.111’E, 8–14 m, 7
May 2002, coll. Alvarez, B and party
82
Halichondriidae from northern Australia
Z.5206
Z.5207
Z.5208
Z.5209
Z.5210
Z.5211
Z.5212
Z.5213
Z.5215
Z.5216
Z.5217
Z.5218
Z.5219
Z.5221
Z.5222
Z.5223
Z.5224
Z.5225
Z.5226
Z.5228
Z.5229
Z.5230
Z.5233
Z.5234
Z.5736
Z.5901
Z.5902
Z.5903
Z.5904
Z.5905
Z.5906
Z.5907
Z.5908
South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 10–12 m, 21 Aug 2002, coll. Alvarez, B and
party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5–10 m, 23 May 2003, coll.
Alvarez, B and party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 7–12 m, 24 Jul 2003, coll.
Alvarez, B and party
Nightcliff bommies, off Nightcliff jetty, Darwin Harbour, NT, 12°22.751’S, 130°50.116’E, 5–8 m, 8 Aug 2003, coll.
Alvarez, B and party
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22
Aug 2003, coll. Alvarez, B and party
Weed Reef, entrance to West Arm, Darwin Harbour, NT, 12°29.25’S, 130°47.54’E, 9–12 m, 6 Sep 2003, coll. Nguyen, H
off Herbert Point, Indian I., Bynoe Harbour, NT, 12°34.586’S, 130°31.419’E, 0–5 m, 24 Jun 2003, coll. Alvarez, B and
party
South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7–11. m, 19 Aug 2002, coll. Alvarez, B and
party
West Arm, 2.5 km N of Stokes Point, Darwin Harbour, NT, 12°31.300’S, 130°48.500’E, 4–5 m, 3 Aug 2002, coll.
Alvarez, B and party
Wickham Point, 2.5 km SW of East Arm Wharf, East Arm, Darwin Harbour, NT, 12°30.12’S, 130°52.39’E, 4–7 m, 15
Sep 2002, coll. Alvarez, B and party
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22
Aug 2003, coll. Alvarez, B
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004,
coll. Alvarez, B and party
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2001’S, 130°47.1’E, 5–19 m, 8 May 2002, coll. Alvarez, B and
party
South Shell I., East Arm, Darwin Harbour, NT, 12°29.869’S, 130°53.141’E, 7–14 m, 20 Aug 2002, coll. Alvarez, B and
party
East Arm Wharf, East Arm, Darwin Harbour, NT, 12°29.19’S, 130°53.35’E, 0.6 m, 1 Mar 2002, coll. Alvarez, B and
party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.238’S, 130°35.557’E, 5–10 m, 23 May 2003, coll.
Alvarez, B and party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.207’S, 130°35.459’E, 3–7 m, 25 May 2003, coll.
Alvarez, B and party
Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.351’S, 130°31.454’E, 6–8 m, 11 Jun 2003, coll. Alvarez, B and
party
Moira Reef, Bynoe Harbour, NT, 12°30.799’ S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Browne, M
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004,
coll. Alvarez, B and party
Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 12–18 m, 16 Sep
2002, coll. Alvarez, B and party
Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, 130°38.371’E, 9–12 m, 23 Aug 2003, coll. Browne,
M
off Dudley Point, Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4–7 m, 4 Jun 2002, coll. Alvarez, B and
party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.600’S, 136°17.839’E, 17–20 m, 30 Mar 2004,
coll. Alvarez, B and party
Mandorah jetty, NW Cox Peninsula, Darwin Harbour, NT, 12°26.55’S, 130°46.05’E, 9–12 m, 5 Sep 2003, coll. Alvarez,
B and party
Moira Reef, Bynoe Harbour, NT, 12°30.799’S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Alvarez, B and party
Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 12–24 m, 3 Sep
2002, coll. Alvarez, B and party
Approx. 3 km NE Charles Point, Cox Peninsula, NT, 12°22.782’S, 130°38.371’E, 9–12 m, 23 Aug 2003, coll. Nguyen,
H
West Arm, 2.5 km N of Stokes Point, Darwin Harbour, NT, 12°31.300’S, 130°48.500’E, 4–5 m, 3 Aug 2002, coll.
Alvarez, B and party
Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m,
7 Nov 2006, coll. Alvarez, B
Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, 12°33.09’S, 130°52.43’E, intertidal 0.02 m,
7 Nov 2006, coll. Alvarez, B
Lee Point, Darwin, NT, 12°20.538’S, 130°52.184’E, 9–12 m, 7 Aug 2003, coll. Nguyen, H
Moira Reef, Bynoe Harbour, NT, 12°30.799’S, 130°30.527’E, 5–8 m, 25 Jun 2003, coll. Nguyen, H
83
B. Alvarez and J. N. A. Hooper
Z.5909
Z.5915
Z.5923
Z. 5925
Z.5928
Z.5948
Z.5949
Z.5950
Z.5965
Z.5970
Z.5976
Z.5977
Z.5978
off Dudley Point, Fannie Bay, Darwin Harbour, NT, 12°24.96’S, 130°48.83’E, 4–7 m, 4 Jun 2002, coll. Alvarez, B and
party
Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 20 m depth, 22 April 2009,
coll. Sultana, S, SCUBA.
Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009,
coll. Sultana, S, SCUBA.
Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009,
coll. Sultana, S, SCUBA.
Larrakeyah sewerage outfall, Darwin Harbour, NT, Australia, 12º28.04’S., 130º49.77’E., 19.5 m depth, 22 April 2009,
coll. Sultana, S, SCUBA.
Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May
2009, coll. Alvarez, B and Sultana, S, SCUBA
Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May
2009, coll. Alvarez, B and Sultana, S, SCUBA
Stevens Rock, near Weed Reef, Darwin Harbour, NT, Australia, 12º29.17’S., 130º47.19’E., 10–16 m depth, 21 May
2009, coll. Alvarez, B and Sultana, S, SCUBA.
East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12º25.01’S., 130º48.88’E., 6 m depth, 21 May 2009, coll.
Alvarez, B and Sultana, S, SCUBA
East Point, Fannie Bay, Darwin Harbour, NT, Australia, 12º25.01’S., 130º48.88’E., 6 m depth, 21 May 2009, coll.
Alvarez, B and Sultana, S, SCUBA
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.188’S, 130°47.110’E, 8–14 m, 22
Aug 2003, coll. Alvarez, B and party
Channel Rock, 4 km NE West Pt on Cox Peninsula, Darwin Harbour, NT, 12°24.94’S, 130°47.04’E, 13–16 m, 5 Sep
2003, coll. Alvarez, B and party
Larrakeyah sewerage outfall, Darwin Harbour, NT, 12º28.04’S, 130º49.77’E, 5.8 m depth, 22 July 2010, coll. Sultana, S,
SCUBA
84