The Beagle, Records of the Museums and Art Galleries of the Northern Territory, 2009 25: 17–42
Taxonomic revision of the order Halichondrida (Porifera: Demospongiae) from
northern Australia. Family Axinellidae
BELINDA ALVAREZ1 and JOHN N.A. HOOPER2
1
Museum and Art Gallery 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
johnh@qm.qld.gov.au
ABSTRACT
Nine species in ive genera of the family Axinellidae, including three new species, Axinella loribellae sp. nov. A. sinoxea
sp. nov. and Phakellia tropicalis sp. nov., are recorded for the tropical northern Australian waters of Western Australia, the
Northern Territory and the Queensland coast as part of a revision of the order Halichondrida (Porifera: Demospongiae)
in this region. One species, Dragmacidon durissimum (Dendy, 1905), generally found in the Indian Ocean, represents
a new record for Australia. Taxonomic descriptions and discussion of those species are presented here. The position of
Reniochalina within the Axinellidae is also discussed based on new evidence found in this and other studies.
Keywords: Sponge, Porifera, Halichondrida, Axinellidae, northern Australia, new species, taxonomy.
Taxonomic knowledge of northern Australian sponges
is limited to a few studies. The irst sponges collected from
this area were described by Ridley (1884) and included
24 species, of which only 17 are currently recognised as
valid species. Bergquist and Tizard (1967) later described
19 species from the rich intertidal area of Darwin Harbour.
Since 1967, there have been 50 additional records to the
fauna of northern Australia and only one revision of a
particular group, i.e. the family Halichondriidae (Hooper
et al. 1997 and references within ). Recent descriptions of
some species have also been included in major taxonomic
revisions of the demosponge families Raspailiidae (Hooper
1991) and Microcionidae (Hooper 1996).
The order Halichondrida is presently represented in
northern Australia by 41 nominal species (Hooper and
Wiedenmayer 1994; Hooper et al. 1997), and a large
number of specimens recently collected and deposited
in the Museum and Art Gallery Northern Territory and
the Queensland Museum (see abbreviations below). Data
gathered from these collections clearly indicate that new
species and records are represented in the area, and that
species previously recorded also need to be revised using
more sophisticated taxonomical tools.
The Halichondrida is a group with an uncertain
classiication and deinition. As with many other sponge
groups, it is deined by traditional morphological characters,
such as growth form, surface characteristics and skeletal
features. But in the Halichondrida, however, these characters
are extremely simple, polymorphic and few, and as a
consequence the discrimination of taxa within this group
is ambiguous. Halichondrid sponges have diverse growth
forms (e.g. encrusting, massive, ramose, tubular, labellate).
INTRODUCTION
The northern marine region of Australia, or the
Northern Province as deined by the Interim Marine and
Coastal Regionalisation of Australia (IMCRA, version
3.3, www.environment.gov.au/coasts/mpa/imcra/index.
html) includes tropical waters off the Northern Territory
(from the Admiralty Gulf in the west) and the Queensland
coasts (western coast of Cape York to Torres Strait in the
east). The continental shelf of this area is generally shallow
(less than 70 m) and extensive, reaching approximately
400 km in width in the Timor Sea and adjoining the coast
of New Guinea in the Arafura Sea and Torres Strait (Bunt
1987; Ferns 1999). The area is part of the central Indo-West
Paciic, which is well known for its high species-richness,
high levels of endemism and is considered to be centre of
origin of many tropical marine species (Veron 1995).
Sponges are one of the most diverse and prevalent
groups of marine invertebrates of northern Australia, but
also one of the most poorly known in terms of proportions
of known and new species, and levels of endemism.
According to Hooper et al. (1997) the northern sponge fauna
includes approximately 800 species, 60% of which remain
undescribed. Further studies based on ‘presence-absence’
analyses of the diversity of tropical Australian sponges
(Hooper et al. 2002) identiied at least two ‘hot spots’ of
biodiversity for the northern area, one in the region of
Darwin and Cobourg Peninsula and the other in the Wessel
Islands region. Only 30% of the sponge species included in
that biodiversity study could be assigned to a known taxon
indicating that a great percentage of the fauna of that region
is not well known.
17
B. Alvarez and J. N. A. Hooper
The skeletons are plumoreticulate, dendritic or confused,
constructed with three types of spicules (strongyles, styles
and oxeas), or transitional forms, in any combination and
not functionally localised. The order includes ive families
(Axinellidae, Dictyonellidae, Heteroxyidae, Halichondriidae
and Bubaridae), and 45 genera, most of which remain poorly
deined despite recent efforts to clarify and redeined the
taxonomy of these families (Alvarez and Hooper 2002;
Alvarez and Van Soest 2002; Hooper 2002a; Van Soest et al.
2002; Van Soest and Hooper 2002). Evidence from molecular
studies (Alvarez et al. 2000) indicates also that some of the
genera are not monophyletic. Moreover, species allocated
to some genera (e.g. Axinella, Acanthella, Phakellia)
have fuzzy boundaries and overlapping characters and
include numerous forms (or varieties/morphs). Similarly,
some allegedly widely distributed species may represent
complexes of cryptic species hiding under morphotypes
that span a continuum, and which cannot be resolved easily
using morphometric data alone.
The taxonomic confusion around the Halichondrida, has
generated long-lasting debates at higher levels of sponge
classiication. Further studies using larger groups of species,
revisions at the regional level and different kinds of genetic
and chemical approaches have been recommended to reine
the current concept of this taxon (Van Soest and Hooper
2002).
The aim of this study is to revise the fauna of the
Halichondrida from northern Australia and the status of all
the nominal halichondrid species in this region. The present
paper represents the irst part of this revision and includes
the family Axinellidae. Revision of the remaining families
represented in the area (i.e. Dictyonellidae, Halichondriidae
and Heteroxyidae) will follow in separate papers.
0˚
10˚
20˚ S
120˚
140˚ E
Fig. 1. Study area included in this taxonomic revision.
Barrier Reef were also examined if the species distribution
was included in the studied area.
Complete locality and collection data of material
included in this revision is indicated under the species
description as usual, whereas non type voucher material
deposited at the Queensland Museum and the Museum and
Art Gallery Northern Territory, is listed in Appendix 1.
Specimens were prepared for light microscopy using the
usual methods (e.g. Hooper 1996; Van Soest and Hooper
2005). Spicule measurements are in micrometres, based
on 25 spicules (otherwise indicated in brackets), of each
category and denoted as range (and mean ± 1 S.E.) of
spicule length x spicule width. Measurements were made
using a digital video camera attached to a light microscope
in combination with the software V++ Precision Digital
Imaging System v 4.0 (© Digital Optics Ltd). Scanning
Electron Microscope photographs were taken in a JEOL
JSM 5610LV. The higher systematic arrangement follows
classification in the current version of World Porifera
Database (Van Soest et al. 2008).
Terminology used here follows Boury-Esnault and
Rützler (1997) and Alvarez and Hooper (2002).
MATERIALS AND METHODS
This revision includes material of the family Axinellidae
recorded for the tropical northern Australian waters of the
Western Australia, 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 meridians
(Fig. 1). The area does not represent a true biogeographical
area and it was delimited based on the marine bioregions
deined by IMCRA. This area also corresponds with two of
the ecoregions (i.e. Arafura Sea, Arnhem Coast to Gulf of
Carpenteria) of the Sahul Shelf marine province as deined
by Spalding et al. (2007).
All specimens recorded for the selected area and
registered under Axinellidae at the Queensland Museum
and the Museum and Art Gallery Northern Territory sponge
collections were examined and identiications were veriied.
Specimens and relevant type material from adjacent areas
including western Australia, the Ashmore, Cartier and
Hibernia reefs on the Sahul Shelf, Lesser Sunda Islands,
Aru Islands, the south coast of Papua New Guinea and Great
130˚
ABBREVIATIONS
18
Abbreviations used in the paper are: AIMS, Australian
Institute of Marine Sciences; BMNH, Natural History
Museum, London (formerly British Museum Natural
History); CRRF, Coral Reef Research Foundation, Palau;
GBR, Great Barrier Reef, NTM, Museum and Art Gallery
Northern Territory, Darwin, Australia (formerly Northern
Territory Museum); MONZ, Museum of New Zealand;
NTM, Museum and Art Gallery Northern Territory
(formerly Northern Territory Museum), Darwin; SMF,
Senckenberg Research Institute and Natural History
Museum, Frankfurt; QLD, Queensland, Australia; QM,
Queensland Museum, Brisbane; WA, Western Australia,
Axinellidae from northern Australia
Australia; ZMA, Zoologisch Museum, University of
Amsterdam, Amsterdam.
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’ program, by the Australian
Institute of Marine Sciences, CRRF and NTM (subcontracted
through CRRF), respectively.
Dragmacidon durissimum (Dendy, 1905)
Phakellia tropicalis sp. nov.
Reniochalina stalagmitis Lendenfeld, 1888
Genus Axinella Schmidt, 1862
Gender feminine. Type species, by subsequent
designation of De Laubenfels (1936), Axinella polypoides
Schmidt, 1862. Recent, Adriatic Sea.
Axinella aruensis (Hentschel, 1912)
(Figs 2 A–F, 3, 4, Table 1)
Phakellia aruensis Hentschel, 1912: 420; Hooper et al.
1992 [in part]; Pulitzer-Finali 1993: 283.
Axinella aruensis. – Hooper and Wiedenmayer 1994:72;
Alvarez et al. 2000 [form II, see below]; Alvarez, Krishan
and Gibb 2007[form II]; Holmes and Blanch 2007.
Material examined. The material examined for this
species is separated according to the morphotypes described
below. Holotype – SMF 953, E side, Aru I., Indonesia,
31 August 1908, coll. Merton, H.. AdditionAl specimens
– Lacepede Is, NW Shelf, WA: NTM Z.2284, Z.2304,
Z.2331, Z.2345. Joseph Bonaparte Gulf: QM G301197,
Cartier I.: QM G301092. Melville I. NT: NTM Z.615,
TAXONOMY
Order Halichondrida Gray, 1867
Family Axinellidae Carter, 1875
Nine species of Axinellidae, listed below, were recorded
within the studied area; three of these being new species.
Axinella aruensis (Hentschel, 1912)
Axinella loribellae sp. nov
Axinella sinoxea sp. nov
Cymbastela stipitata (Bergquist and Tizard, 1967)
Cymbastela vespertina Hooper and Bergquist, 1992
Dragmacidon australe (Bergquist, 1970)
Table 1. Comparison of spicule dimensions among specimens and varieties of Axinella aruensis. Measurements in micrometres.
Specimen
Locality
Axinella aruensis
SMF 953
Aru Is, Indonesia
Z.2304
Lacepede Is, WA
G301092
Cartier Is, WA
Z.619
Melville I., NT
Z.3141
Parry Shoals, NT
Z.5053
Darwin Harbour, NT
Z.4465
Wessel Is, NT
Oxeas
Styles
257.1–423.9 (360.6±38.1)
x 13.8–21.4 (16.5±1.9)
245.5–337.6 (285.4±21)
x 8.6–17.2 (13±2.6)
281.2–450.2 (360.7±39.9)
x 10.3–19.4 (15.6±2.2)
236.1–406 (302.4±39.4)
x 9.3–17.5 (13.9±2.3)
267.1–372.9 (307.1±23.3)
x 9.4–17.2 (13.6±2.3)
297.6–498.6 (392.1±47)
x 7.3–22.5 (16.5±3.7)
194.4–396.4 (299.5±48.6) [24]
x 5.6–17.4 (12±3.3)
249.1–382.2 (313.6±40.7) [12]
x 14.1–21.7 (17.4±2.2) [12]
213.2–271.4 (244.8±20.9) [9]
x 11.9–15.7 (13.5±1.6) [9]
242.8–419 (301.1±36)
x 12.6–20.3 (16.8±1.9)
186–362.8 (267.2±43.3) [22]
x 9.8–17.1 (14.1±1.9) [22]
248.6–294.6 (270.4±18.8) [4]
x 11.1–16.7 (13.9±2.3) [4]
263.6–417.2 (342.7±37.6)
x 12–23.5 (17.1±2.9)
204.6–331.7 (269.7±44.1) [15]
x 8.6–18 (13.8±2.8) [15]
305–451.7 (376.7±36.3)
x 13.1–25.9 (18.3±3.9)
187.1–318.2 (237.2±26.6)
x 8.5–17.2 (13.2±2.4)
266.7–354.5 (312.1±23.3)
x 13.1–18.8 (16.3±1.4)
274.8–392.8 (333±32.8)
x 8.3–16.4 (13.3±2.3)
283.1–406.6 (334.7±40.7) [6]
x 14.7–19.6 (16.5±1.8) [6]
167.2–222.5 (200±15.3)
x 9.3–15.5 (13.2±1.7)
200.1–353.2 (260.7±34.1)
x 10.3–22.8 (15.4±2.9)
248.6–363.3 (297±31) [10]
x 11.5–18 (15.1±2.3) [10]
209.1–278.6 (246.3±15.8)
x 12.7–19.9 (15.1±1.8)
173.8–247.4 (214.7±18.9)
x 7.5–16.7 (10.3±1.9)
163.6–231.4 (191.1±17.4)
x 6.5–12.5 (9.8±1.6)
166.4–262.3 (218.2±22.4)
x 7.8–17.9 (13.9±2.4)
Axinella aruensis form I
Z.5816
Bynoe Harbour, NT
Z.3068
Parry Shoals, NT
Z.5819
East Point, Darwin, NT
Z.3946
Wessel Is, NT
Axinella aruensis form II
Z.4490
Stevens Rock, Darwin
Z.5054
Wessel Is
19
B. Alvarez and J. N. A. Hooper
A
C
B
D
F
E
G
H
Fig. 2. Axinella aruensis: A, B, specimens at Raragala I., Wessel Is; C, specimen at South Shell I., Darwin Harbour; D, form I, Z.5816, Dawson
Rock, Bynoe Harbour; E, form II, specimen at Raragala I, Wessel Is; F, Axinella sinoxea sp.nov., NTM Z.2719. Axinella loribellae sp. nov.;
G, Holotype, NTM Z.4427; H, NTM Z.5662. Photos: A–B, P. Colin; C–D, G, B. Alvarez; E, D. DeMaria; F, J. Hooper; H, A. Ayling.
20
Axinellidae from northern Australia
Z.619, Z.630, Z.632. Parry Shoals, Arafura sea, NT: QM
G310136 (Q66C0514-X), NTM Z.3062 (Q66C87-0514-X),
Z.3141.Bynoe Harbour, NT: NTM Z.5071 (0M9H2464-U).
Darwin Harbour, NT: NTM Z.5053 (0M9H2168-X),
Z.5057 (0M9H2665-O), Z.5058 (0M9H2675-Y), Z.5072
(0M9H2579-U), Z.5830. Cobourg Peninsula, NT: NTM
Z.1363, Z.1388, Z.2511, Z.2526, Z.2529. English Company
Is., NT: NTM Z.3956. Wessel Is, Gove Peninsula, NT: QM
G3.609 (Q66C4762-R), G300768 (=Q66C4737P, QM
G311873 and NTM Z.3945), Z.3922 (Q66C4687-L), Z.3935
(=Q66C4785-R, QM G300752), Z.3936 (Q66C4831-R),
Z.4465 (0M9H2770-C), Z.5055 (0M9H2650-W). Papua
New Guinea: QM G312913, G312935.
Axinella aruensis, form I, Bynoe Harbour, NT: NTM
Z.5816, Z.5817, Z.5818. Darwin Harbour, NT: NTM
Z.2156, Z.5819-Z.5823. Wessel Is, Gove Peninsula, NT:
NTM Z.3925, Z.3946.
Axinella aruensis, form II. Darwin Harbour, NT:
QM G303332, Z.1961, Z.2249, Z.2402, Z.2632, Z.4425
(0M9H2044-O), Z.4490, Z.4491, Z.5824-Z.5829, Z.5831,
Z.5232. Parry Shoals, Arafura sea, NT: NTM Z.3137,
Z.3068. Wessel Is, Gove Peninsula, NT: QM G300759
(Q66C-4831-R), NTM Z.5054 (0M9H2648-U). Yampy
sound, WA, NTM Z.665.
Description. Three different morphotypes of this species
with one corresponding to the holotype are recognisable
among the material examined and they will be described
below separately under the heading of ‘forms’.
Axinella aruensis, typical form. Shape (Fig. 2A–C).
Thickly labellated, on broad and short, or long and narrow,
peduncle, uni or bi-planar, sometimes folded, with round
margins projecting in most cases into short and broad
extensions with square, round or pointed tips, or in long
rounded to flat branches which tend to fuse laterally.
Specimens up to 400 mm high.
Colour. Orange, pale yellow or yellowish brown alive.
Dark brown in alcohol.
Oscula. Regularly distributed in one or both sides of
fan, stellate, lush or with elevated rims, less than 5 mm
diameter.
Surface. Evenly microconulose-conulose, nodulose,
rough, marked with primary longitudinal choanosomal
ibres.
Skeleton (Fig. 3A). Plumose, vaguely reticulated to
halichondroid, very compact, with plumose columns
up to 600 µm thick, diverging toward surface, ending
in fan-shapped spicule brushes and projecting through
ectosome. Axial skeleton differentiated only towards base
of attachment or peduncle, halichondroid.
Spicules (Fig. 3B). Oxeas with blunt, pointed or
telescoped tips; slightly bent and sometimes slightly sinuous,
195–498 x 5–22 µm. Styles, less frequent or rare, similar
in size to oxeas (see Table 1) with blunt ends, enlarged or
slightly narrow bases, straight or slightly bent. Transitional
forms (e.g. styloids, strongyles) are common.
A
B
C
D
E
F
Fig. 3. Axinella aruensis: light microphotograph of skeleton and
diagram of spicules: A, B, SMF 953, holotype; C, D, Z.5819 (form
I); E, F, NTM Z.5054 (form II). Scale bars: A, 200 µm; B, D, F, 50
µm; C, E, 500 µm.
Axinella aruensis, form I. Shape (Fig. 2D). Erect,
fan-shaped or narrow long and lat digits with few simple
ramiications, generally with square margins. Specimens
up to 20 cm high and 10 cm wide.
Colour. Light orange, brown, beige or yellow. Same
colour in alcohol.
Oscula. Regularly distributed, less than 5 mm diameter,
with distinctive raised margins.
Surface. Minutely hispid, marked with choanosomal
skeletal tracts in a regular reticulation or with radial
grooves.
Skeleton (Fig. 3C). Plumose and slightly compressed
at axial region, with thick plumo-echinated multispicular
columns, up to 1 mm thick, and radiating outwards towards
surface, anastomosing or connected irregularly by short
and thick paucispicular or multispicular tracts, or by single
spicules oriented in any direction. Main tracts end at surface
in fan-shaped brushes with spicules projecting shortly
through ectosome; light spongin embedding tracts.
Spicules (Fig. 3D, Table 1). Oxeas, 187–451 x 8–25 µm,
with pointed or blunt ends, straight or bent; thinner forms
are common. Styles less frequent or rare, slightly smaller,
21
B. Alvarez and J. N. A. Hooper
including intermediate forms similar to styloids, anisoxeas or
strongyles. Fused spicules are characteristically common.
Axinella aruensis, form II. Shape (Fig. 2E). Thick
fans or lamellae with round margins, folding in more than
one perpendicular plane, or joining at angles from 45-90
degrees; or short single or digitate projections, stipitate, on
short narrow peduncles or on broad base. Generally small
with individuals reaching up to 13 cm high.
Colour. Bright or light orange alive. Light beige in
alcohol.
Oscula. Stellate with minute drainage canals, sometimes
located at margin of fans or evenly distributed in both sides
of fan, 3-5 mm in diameter.
Surface. Pierced uniformly with minute ostia.
Microconulose; minutely hispid, firm but some has
mucous consistency after collection. Marked by regular
choanosomal reticulation.
Skeleton (Fig. 3E). Thick and dense plumo-echinated
multispicular tracts up to 600 µm wide, forming regular,
nearly radial reticulation which is marked on surface. Main
columns end at surface in fan-shaped brushes, with spicules
projecting shortly through ectosome. Axial skeleton not
differentiated.
Spicules (Fig. 3F, Table 1). Oxeas and styles in nearly
equal proportions, 174-279 x 7-20µm. Styles are dominant
and slightly smaller than oxeas.
Remarks. Although some features allowed distinction
of two additional morphotypes within this species, the limits
among them are not clear and some individuals could be
considered intermediate forms. Axinella aruensis sensu
stricto is distinguished from its two other forms by shape
and colour, generally observed to be thickly labellate and
orange when alive, by the change of colour in alcohol (it
turns brown) and by the dominance of oxeas in relation to
styles. Form I differs slightly in shape from A. aruensis;
the colour in life is always beige or pale yellow and does
not change in alcohol; oxeas are also dominant and styles
are relatively more common. Form II is always beige in
alcohol and styles are dominant relative to oxeas. Some
other features of shape and surface consistency and texture
are also distinctive within this form.
Some data included in Hooper et al. (1992) indicate
there are some differences in the biochemistry between
populations (i.e. North West Shelf versus Darwin Harbour)
of this species, however the published results do not seem
to be related to the forms distinguished here.
Alvarez et al. (2007) detected up to 29% of intra-genomic
polymorphism within the Internal Transcribe Spacer (ITS)
of the rDNA in individuals of Axinella aruensis from Darwin
Harbour. These levels of intra-genomic variation are so far
the highest reported for Porifera and correspond in most
cases to hybrid species reported for other groups, including
corals of the genus Acropora. Thus, it is possible that the
forms here distinguished to document the variability present
within species are the result of a hybridisation processes
with sympatric species or populations. Future population
genetic studies will help to determine whether the variability
observed across these forms are signiicant to justify their
recognition as different species or as hybrids.
Axinella aruensis is very similar in shape, skeletal
architecture and spicule composition and dimensions to
some Axinella species recorded from the Indian Ocean
(e.g. A. donnani (Bowerbank, 1873); A. manus Dendy, 1905
and A. symmetrica (Dendy, 1905, as Phakellia)). Skeletal
reticulation of the Indian Ocean species, in particular
A. donnani, is much more regular, with thicker primary
lines. Interesting also is the change in colouration, from
orange to brown after few hours of collection, reported for
A. donnani (Bowerbank, 1873), a characteristic also seen in
A. aruensis. A detailed revision of the Indian Ocean species
complemented with population genetic studies is essential
to deine their limits and phylogenetic relationships with
the northern Australian populations of A. aruensis and its
forms.
Distribution. Axinella aruensis and its forms appear to
occur sympatrically and are common throughout northern
Australia (Fig. 4). The type locality is the Aru Is, Indonesia,
but the species is also known from other Indonesian
localities (Alvarez and de Voogd, unpublished data) and
from Papua New Guinea. It is found in subtidal areas from
5 to 76 m. Pulitzer-Finali’s (1993) record for East Africa,
is dubious and requires conirmation.
Axinella loribellae sp. nov
(Figs 2G–H; 5)
Material examined. H olotype – NTM Z.4427
(0M9H2041-L), Stevens Rock, Weed Reef, Darwin
Harbour, 12°29.2001´S, 130°47.1´E, NT, 5–19 m depth, 8
May 2002, coll. B. Alvarez and party. pArAtypes – ntm
Z.5834 Stevens Rock, Weed Reef, Darwin Harbour,
Northern Territory, Australia, 12°29.1667´S, 130°47.19´E,
17 m depth, 8 May 2006, coll. B. Alvarez.
Additional specimens. Melville I., NT, NTM Z.631.
Darwin Harbour, NT: QM G303388, NTM Z.822,
Z.868, Z.5662. Wessel Is, NT: NTM Z.3938, Z.5059
(0M9H2771-F).
120˚ E
128˚
136˚ E
8˚S
16˚ S
Fig. 4. Distribution of Axinella aruensis (open circles) and its forms
(I, black circles; II, grey circles) based on conirmed records from
QM and NTM.
22
Axinellidae from northern Australia
A
tracts oriented perpendicular to surface and laterally close,
connected by single spicules or uni-paucispicular tracts up
to 2 spicules long, cemented with thin and clear spongin, to
each other and ending in brushes of spicules that protrude
shortly through ectosome.
Spicules (Fig. 5C; Table 2). Styles slightly bent;
strongyles straight, slightly sinuous, bent in middle; oxeas
fusiform, straight. Transitional shapes between monoactins
and diactins are common. All types in a wide range of sizes
and thickness. Dominant types vary among specimens (see
below).
Remarks. A great variability in the shape and size of
spicules was observed among specimens of this species.
Styles and strongyles of 190–270 by 7–13 µm, are the
dominant types, however they were absent in some of the
examined specimens (i.e. NTM Z.631 and NTM Z.3938).
Apart from these differences in spicule composition and
dimensions, no other characters seem to vary among
specimens examined here, and consequently they are
considered at this stage to belong to a single species.
Further genetic studies may help to conirm whether or
not variability in spicule dimensions and composition is
indicative of sibling species differentiation.
The new species differs from Axinella aruensis and its
related species from the Indian Ocean (see above) mainly
in shape (thinly instead of thickly labellate); in skeletal
architecture (clearly differentiated into axial and extraaxial skeleton with thinner extra-axial spicular tracts and
more regular reticulation when compared to the thicker
multispicular columns present in A. aruensis) and in spicule
composition (with common transitional shapes between
oxeas and styles not observed in A. aruensis).
Axinella ceylonensis (Dendy, 1905, as Phakellia) from
the Gulf of Manaar is similar to A. loribellae in shape
(although shortly stipitate) and in thickness of the lamellae.
Both species share the variability observed in the size and
shape of oxeas and styles. The two species differ in skeletal
and surface characteristics. The skeleton of A. ceylonensis
is not differentiated in axial and extra-axial region but
described as plumose, with columns radiating outwards into
small surface conules. No other species in the study area or in
the Indian Ocean is similar to this new species. Indeed, most
nominal species of Axinella reported for the Indian Ocean,
do not agree with the current diagnosis of the genus and
need to be re-examined. Some of these are massive forms
with styles and trichodragmata in the skeleton and might
belong in the axinellid genus Dragmacidon (e.g. Axinella
C
B
Fig. 5. Axinella loribellae sp. nov.: A, Paratype, NTM Z.5834;
B, light microphotograph of skeleton; C, diagram of spicules. Scale
bars: A, 2 cm; B, 100 µm; C, 50 µm.
Description. Shape (Figs 2G–H, 5A). Fan-shaped; thin
lamellae, 1–5 mm thick, single or bifurcated, sometimes
convoluted with rounded margins and indentations on short
stalks or broad base. Specimens are 300 mm high and up
to 400 mm wide.
Colour. Burnt orange alive, brown in alcohol.
Oscula. Small, 2–5 mm diameter, with stellate drainage
canals, evenly distributed.
Consistency. Flexible, easy to tear, rubbery.
Surface. Smooth, velvety, marked irregularly with
ribs.
Skeleton (Fig. 5B). Plumose, differentiated in extraaxial and axial region. Axial skeleton compressed with
wavy longitudinal paucispicular-multispicular tracts,
lightly embedded in collagenous spongin, interwoven,
and radiating towards the extra-axial region. Extra-axial
region a close-set reticulation of plumose paucispicular
Table 2. Comparison of spicule dimensions among specimens of Axinella loribellae sp. nov. Measurements in micrometres.
Specimen
Z.4427 (Holotype)
Locality
Darwin Harbour
Z.5059
Wessel Is
Z.631
Melville Is
Styles
196.3–352.9 (274.6±47.7)
x 8.33–18.3 (13.5±2.7)
159.7–365.4 (250.6±50.4)
x 5.8–14.43 (10.0±2.1)
187.6–307.9 (228.9±29.0)
x 4.3–11.3 (7.3±1.8)
23
Strongyles
103.6–396.3 (190.1±74.0)
x 6.01–13.09 (10.3±1.9)
93.7–531.8 (243.1±128.6)
x 5.8–13.3 (8.8±2.3)
–
–
Oxeas
148.5–440.2 (226.8±63.6)
x 4.8–12.9 (8.7±2.1)
100.8–302.2 (208.5±50.1)
x 3.7–11.4 (6.5±2.1)
142.8–351.6 (217.8±44.1)
x 3.5–13.2 (7.1±2.7)
B. Alvarez and J. N. A. Hooper
surface; slightly compressed in axial region. Spicule tracts
bound only slightly with clear collagenous spongin.
Spicules (Fig. 6D; Table 3). Styles robust, bent, or less
often, straight, enlarged in the middle section, 159–245 x
7–17 µm; thinner category, 97–201 x 2–6 µm also present.
Long thin raphids abundant. Smaller oxeas and thick and
short strongyles, very rare.
Remarks. This species conforms in most of its
characteristics with the current concept of Axinella. The
absence or low frequencies of oxeas observed in the
examined specimens are also seen in other Caribbean
species of the genus (e.g. Axinella waltonsmithi (de
Laubenfels, 1953) and A. pomponiae Alvarez, Van Soest
and Rützler, 1998), which might be considered a common
feature among Axinella species.
The new species resembles Axinella aruensis in gross
morphology and as such can be easily mistaken for it
in the ield; both are fan-shaped on a common stalk and
both have a similar surface pierced with minute ostia and
microconulose. But A. sinoxea is clearly different from
A. aruensis in skeletal architecture and spicule composition,
having a regular plumoreticulated skeleton of ascending
tracts and long thin raphids in the skeleton. As is the case
with A. loribellae, no other species recorded in the study
area or in the Indian Ocean was found to be related to
A. sinoxea.
Distribution. Common in the vicinity of East Point
Sponge Gardens, Darwin Harbour, but also found in deeper
waters (down to 40 m) of Western Australia.
Etymology. Latin, sine- without; sinoxea referring to
the lack of proper oxeas characteristic of the species. It is
intended as a noun in apposition.
Remarks on Axinella. Axinella is a widespread genus
of sponges with approximately 100 accepted species (Van
Soest et al. 2008), many of which, however, need to be
veriied against the current deinition of the genus (Alvarez
and Hooper 2002). Ongoing revisions of species of Axinella
and related genera by one of the authors (BA) are undertaken
on a regional basis with the purpose to verify the identity of
the reported species and the monophyly of the genus, which
is currently proven as polyphyletic based on molecular
studies (Alvarez et al. 2000; Erpenbeck et al. 2005).
Three species of Axinella (A. aruensis, A. loribellae,
A. sinoxea) are reported in this work. No other species of the
genus, as far as we know, have been reported within the area
of northern Australia that is the subject of this present study.
Axinella echidnaea reported by Ridley 1884 is accepted as
Reniochalina stalagmitis (see below).
bidderi Burton, 1959 and A. massalis Burton, 1959). Some
others are Stylissa-like, or other dictyonellid genera, with
the surface marked by ridges or conules and with a dense
and irregular skeleton of multispicular tracts of styles
(e.g. Axinella bubarinoides Dendy, 1922; A. dragmaxioides
Burton, 1959 [?]; A. labelloreticulata (Burton, 1959); A.
labyrinthica Dendy, 1889; A. minor Thomas, 1981; A.
proliferans Ridley, 1884; A. tenuidigitata Dendy, 1905;
A. ventilabrum Burton, 1959), or closer to halichondrid
genera (e.g. Axinella halichondrioides Dendy, 1905, which
is encrusting and has only oxeas) or to the raspailiid genus
Ceratopsion (i.e. Axinella lamellata Dendy, 1905, with a
dermal and tangential layer of small oxeas).
Distribution. Axinella loribellae seems to be restricted
to northern Australia between Darwin Harbour and the
Wessel Is. It is found between 11–32 m depth.
Etymology. Named after Lori Bell, Coral Reef Research
Foundation, Palau, for her considerable contribution to the
knowledge of Indo-Paciic sponge diversity and distribution.
It is intended as a noun in apposition.
Axinella sinoxea sp. nov.
(Figs 2F, 6A–D)
Material examined. Holotype – NTM Z.940, East
Point, Darwin Harbour, NT, 12°24.05´S,130°48.01´E, 12 m
depth, 13 September 1982, coll. Hooper, J.N.A. pArAtypes
– Z.5833, East Point, Darwin Harbour, NT, 12º 24.484´S,
130º 48.471´E, 11 m depth, 7 June 2007, coll. B. Alvarez.
Additional specimens. NW Shelf, WA: NTM Z.2310,
Z.2322. Darwin Harbour, NT: NTM Z.2246, Z.2719.
Description. Shape (Figs 2F, 6A). Single or multiple
fans, 4–6 mm thick, 8–14 cm long and up to 30 cm wide,
on common stalk, 3–5 cm long and 7–10 mm in diameter;
erect, uniplanar with digitate to irregular margins or
bifurcate tips.
Colour. Orange, pale yellow or beige with light pink
tinge alive; brown-grey in alcohol.
Oscula. Regularly distributed in one or both sides of fan,
round to elongated or irregularly shaped, some stellate, with
slightly elevated rims, less than 1 mm diameter.
Consistency. Soft, floppy, flexible, slightly
compressible.
Surface. Smooth but slightly rough to touch; pierced
regularly by minute pores, microhispid due to projections
of brushes of choanosomal spicules. Encrusted irregularly
with detritus in some specimens.
Skeleton (Figs 6B-C). Plumoreticulated, with ascending
multispicular tracts connected regularly by single spicules or
unispicular tracts, 1 or 2 spicules long, ending in brushes at
Table 3. Comparison of spicule dimensions among specimens of Axinella sinoxea sp. nov. Measurements in micrometres.
Specimen
Z.940 (Holotype)
Locality
East Point, NT
Z.2310
NW Lacepede
Is WA
Thick styles
Thin styles
Raphids
184.5–245.1 (223.7±12.92) 97.77–201.3 (179.64±22.84) [23] 192.9–249.6 (227.2±14.9)
7.35–17.4 (13.11±2.49)
x 2.48–5.6 (3.8±1) [23]
x 0.8–3.0 (2.0±0.6)
153.4–197.6 (183.4±10.2)
131.3–191.2 (152.1±14.8)
131.9–258.6 (201.5±35.1)
x 7.8–12.2 (10.2±1)
x 2.2–6.4 (4.4±1.)
x 0.37–2.69 (1.6±0.5)
24
Axinellidae from northern Australia
A
B
C
D
Fig. 6. Axinella sinoxea sp. nov. NTM Z.940: A, photograph of the holotype; B, light microphotograph of the skeleton; C, SEM, raphids in
choanosomal skeleton; D, scale bars: A, 5 cm; B, 500 µm; C, 100 µm; D, 20 µm.
In the present study area, the genus Axinella seems to be
less speciose than in other taxonomically revised regions:
seven species in the Western Central Atlantic (Alvarez
et al. 1998); six (recorded) species and nine possible new
species (Kelly et al. 2009); at least ive species in Indonesia
(Alvarez and De Voogd, unpublished data). Unfortunately
little is known about the biology of these species to explain
why the genus might be more diverse in some areas than
others. Many more putative Axinella species are known for
the GBR (pers. obs.), but these remain unresolved pending
future studies.
Distinction of Axinella species continues to be
subjective and is based on a combination of characters
as discussed by Alvarez et al (1998). Variability and
plasticity of all the morphological characters that
characterise the species are seen in all the species
described above. The morphological variability of A.
aruensis, for example, is remarkable and suggestions
from molecular data (Alvarez et al. 2007) that it may be
due to hybridisation should be further explored.
Genus Cymbastela Hooper & Bergquist, 1992
Gender feminine. Type species, by original designation,
Pseudaxinyssa stipitata Bergquist and Tizard, 1967. Recent,
Darwin Harbour, Arafura Sea.
25
Cymbastela stipitata (Bergquist and Tizard, 1967)
(Figs 7A–B)
Pseudaxinyssa stipitata Bergquist and Tizard, 1967:189;
Hooper et al. 1992: 265.
Cymbastela stipitata. – Hooper and Bergquist 1992: 106;
Hooper and Wiedenmayer 1994 : 75; Alvarez et al. 2000:
195; Alvarez and Hooper 2002: 733.
Material examined. Specimens as listed in Hooper and
Bergquist (1992). AdditionAl specimens – Bynoe Harbour,
NT: Z.5065 (0M9H2333-C). Darwin Harbour, NT: QM
G303262, NTM Z.4078 (0CDN8001-H, Fig. 7A), Z.4104
(0CDN8026-J), Z.4131, Z.4435 (0M9H2008-Y), Z.5064
(0M9H2134-M), Z.5835, Z.5836, Wessel Is, NT: Z.5066
(0M9H2658-H), Z.5067 (0M9H2785-T).
Remarks. Cymbastella stipitata, was re-described
extensively by Hooper and Bergquist (1992) and the type
material re-examined by Alvarez and Hooper (2002).
This is one of the most common sponges in the studied
area. It is particularly abundant in the intertidal zone of
Darwin Harbour, which becomes greatly exposed during
the nocturnal king tides of the dry season (May–June) and
the diurnal king tides during the wet season (SeptemberNovember, Fig. 7B). It is found, but less commonly, in
subtidal areas down to 19 m depth. New records indicated
that its distribution within northern Australia extends from
B. Alvarez and J. N. A. Hooper
A
B
C
D
E
F
H
G
Fig. 7. Cymbastela stipitata: A, NTM Z.4078 (0CDN-8001-H); B, specimens exposed at the reef lat of East Arm, Darwin Harbour during the
low tide of 20 September 2001. Dragmacidon australe: C, specimen at Channel I. Darwin Harbour; D, QM G304246, Lizard I, GBR, QLD.
Phakellia tropicalis sp. nov.; E, Holotype (NTM Z.5847); F, Paratype (NTM Z.5845). Reniochalina stalagmites: G, specimen at East Point,
Darwin; H, specimen at Cotton I., Wessel Is. Photos: A, B, E, F, B. Alvarez; C, H. Nguyen; D, J. Hooper; G, A. Ayling; H, P. Colin.
26
Axinellidae from northern Australia
125˚E
135˚
Genus Dragmacidon Hallmann, 1917
Gender neuter. Type species, by original designation,
Thrinacophora agariciformis Dendy, 1905. Recent, Gulf
of Manaar, Indian Ocean.
145˚
5˚
Dragmacidon australe (Bergquist, 1970)
(Figs 7 C–D, 9 A–B)
Pseudaxinella australis Bergquist, 1970: 20; Hooper
and Lévi 1993: 1441; Hooper and Wiedenmayer 1994: 80;
Alvarez et al. 2000: 196.
Dragmacidon australe. – Alvarez and Hooper 2002:
735; Kelly et al. 2009 (In press).
Material examined. Holotype – NMNZ Por. 26, Takatu
Channel, Northland, New Zealand, 11 m. AdditionAl
specimens – Cartier I, WA: QM G301089. Bynoe Harbour,
NT: G303444. Darwin Harbour, NT: NTM Z.5068. Coral
Sea, GBR, QLD: QM G300295, G304182, G304246,
G304253, G320664, NTM Z.2727.
Description. Shape (Fig. 7 C–D). Thickly encrusting,
following substrate, globular, bulbous or semispherical,
approx. 100 mm in diameter by 20 mm thick.
Colour. Bright red, orange alive.
Consistency. Slightly compressible or stiff. Mucous
surface.
Oscula. Irregularly distributed, less than 1 mm diameter,
with slightly elevated rims surrounded by thin drainage
channels in stellate arrangement.
Surface. Highly conulose; evenly pierced by pores
120–400 µm in diameter. Conules, approx. 1–3 mm long,
single or grouped in reticulated pattern, unevenly echinated
by spicules, 2–3 mm apart.
Skeleton (Fig. 9A). Plumoreticulate to halichondroid;
formed by thick plumose or plumo-echinated multispicular
tracts, up to 500 µm thick, ascending nearly perpendicularly
from base and becoming thicker and bushy near surface;
projecting through ectosome into surface conules. Main
tracts connected by shorter and thinner plumose tracts,
sometimes ill-deined, forming irregular reticulation of
large round meshes.
Spicules (Fig. 9B; Table 4). Oxeas and styles in equal
proportions, 176–510 x 7–21 µm (Table 4).
Remarks. The species was originally assigned to
Pseudaxinella and transferred to Dragmacidon by Alvarez
15˚ S
Fig. 8. Distribution of Cymbastela stipitata, based on conirmed
records from QM and NTM.
Bynoe Harbour to the Wessel Is (Fig. 8), inferring it is a
narrow range endemic within northern Australia.
Cymbastela vespertina Hooper and Bergquist, 1992
Cymbastela vespertina Hooper and Bergquist, 1992:
110; Hooper and Wiedenmayer 1994: 75; Alvarez et al.
2000: 195; Alvarez and Hooper 2002: 733.
Pseudaxiyssa sp. nov. Hooper et al. 1992: 265.
Material examined. Specimens as listed in Hooper and
Bergquist (1992).
Remarks. Cymbastela vespertina is a sibling species
of C. stipitata. Separation of the two species based in
morphology is dificult and very subjective. Biochemical
and molecular evidence (Hooper et al. 1992; Alvarez et al.
2000) indicates the sympatric populations are heterogeneous.
Future genetic population studies might reveal whether or
not these populations can be reliably separated into different
species, but based on current external gross morphological
differences and skeletal characters the two taxa are
maintained as distinct.
Remarks on Cymbastela. Two species of Cymbastela are
present within the area of the present study (i.e. C. stipitata
(Bergquist and Tizard, 1967) and C. vespertina Hooper and
Bergquist, 1992). Other species of Cymbastela represented
in other regions including Australia are: C. cantharella
(Lévi, 1983), New Caledonia; C. concentrica (Lendenfeld,
1887), Queensland coast; C. coralliophila Hooper and
Bergquist, 1992, GBR; C. marshae Hooper & Bergquist,
1992, Houman-Abrolhos, WA; C. notiaina Hooper and
Bergquist, 1992, South Australia; and C. tricalyciformis
(Bergquist, 1970) from New Zealand. Comprehensive
descriptions of these species are given by the respective
authors of the species.
Phylogenetic relationships of the genus with other
axinellid species based on molecular characters indicate
that the northern Australian species of Cymbastela are
closely related to other members of Dictyonellidae, such
as Acanthella (Alvarez et al. 2000; Erpenbeck et al. 2005).
There is no doubt, however, that species of Cymbastela are
related to the Axinellidae based on their morphology. Thus
the phylogenetic relationships derived from these molecular
analyses remain enigmatic at this stage.
A
B
Fig. 9. Dragmacidon australe: A, light microphotograph of skeleton;
B, diagram of spicules. Scale bars: A, 500 µm; B, 50 µm.
27
B. Alvarez and J. N. A. Hooper
Table 4. Comparison of spicule dimensions among specimens of Dragmacidon australe. Measurements in micrometres.
Specimen
Locality
Styles
Oxeas
G303444
Bynoe Harbour
254.1–510.4 (349.1±78.8)
x 7.3–20.9 (13.6±3)
286.2–434.5 (361.5±39.8)
x 7.3–17.9 (13.4±2.4)
Z.5068
Darwin Harbour
176.4–397.3 (290.1±61.4)
x 7–17.2 (13±2.8)
271.9–412.5 (340.7±31.9)
x 9.5–18.9 (14.3±2.4)
and Hooper (2002) because it conformed more closely with
the type species of that genus.
Hooper and Lévi (1993) compared specimens from the
GBR with the holotype of Dragmacidon australe from
New Zealand and with material from New Caledonia
described as D. debitusae (Hooper and Lévi, 1993). Very
subtle differences were found between the two species. The
material from northern Australia reported here agrees with
D. australe in the majority of its features and is therefore
assigned to this species. This species is also very similar to
D. reticulatum (Ridley and Dendy, 1886) from the central
West Atlantic both in external morphology and spicule
composition.
Distribution. Dragmacidon australe was irst recorded
for New Zealand and additional records from the GBR were
reported in Hooper and Lévi (1993). The present revision
extends the distribution range of this species into northern
Australia. The species is not very common in this region
with only isolated records registered through the extension
of the studied area (Fig. 10) and thus is probably at the edge
of its range. It is also found along more temperate areas of
the Queensland coast (Hooper pers. obs.).
125˚E
135˚
145˚
5˚
15˚ S
Fig. 10. Distribution of Dragmacidon australe in northern Australia,
based on conirmed records from QM and NTM. Species distribution
extends along the Queensland coast (Hooper pers. obs.) and New
Zealand (type locality).
A
C
Dragmacidon durissimum (Dendy, 1905)
(Figs 11 A–C)
Thrinacophora durissima Dendy, 1905:187.
Sigmaxinella durissima. – Dendy 1922: 113.
Axinella durissima. – Burton 1959: 259.
Pseudaxinella durissima. – Alvarez et al. 2000: 196.
Dragmacidon durissima. – Hallmann 1917: 639; Alvarez
and Hooper 2002: 735.
Material examined. Ashmore Reef, WA: QM
G300181.
Description. Shape (Fig. 11A). Hemispherical, cushionshaped.
Colour. Orange alive, red on deck, beige in ethanol.
Oscula. Round, irregularly distributed at top, with
slightly elevated rims.
Surface. Very rugose, composed of minute projections
or conules, compact and close-knit; membranous skin
stretched over conules.
Skeleton (Fig. 11B). Plumoreticulate. Multispicular,
plumose or plumoechinated spicule tracts, ascending toward
surface and connected by shorter and thinner ones, or loose
spicules, forming irregular reticulation of oval to square
meshes; projecting through ectosome in surface conules
or projections.
B
Fig. 11. Dragmacidon durissimum: A, QM G300181; B, light
microphotograph of skeleton; C, diagram of spicules. Scale bars:
A, 1 cm; B, 500 µm, C, 50 µm.
28
Axinellidae from northern Australia
Spicules (Fig. 11C). Styles 203.1–312.5 µm (251.8±33.4)
by 11.7–16.4 µm (13.7±1.2). Oxeas in equal proportions,
229.8–312.7 µm (283.2±18) by 7.4–19 µm (13.4±3.1).
Trichodragmata short and thick, 15–20 by 5–10 µm.
Distribution. Indian Ocean, including Seychelles Is,
Amirante, Providence, Saya de Malha (Dendy 1905; Dendy
1922; Burton 1959), Maldive Is (Alvarez and de Voogd,
unpublished data) and Ashmore Reef, Australia.
Remarks. The material examined here agrees in all its
characteristics with Dragmacidon durissimum, an Indian
Ocean species never previously recorded in Australia. Only
one specimen from WA was found among the collections
examined in this revision. More isolated populations might
be present along the WA coast, given that the species is
widely distributed throughout the Indian Ocean.
The species was included originally in Thrinacophora
due the presence of trichodragmata, later transferred to
Sigmaxinella by Dendy (1922), and subsequently to Axinella
by Burton (1959) without suficient justiication. Hallman
(1917) erected Dragmacidon for D. agariciformis (Dendy,
1905), D. durissimum, D. clathriformis (Lendenfeld, 1888)
and D. incrustans (Whitelegge, 1897). All these species are
very similar in habitat, spicule composition and all include
trichodragmata. They closely resemble to the West African
species D. lunaecharta (Ridley and Dendy, 1886). Also
similar are the Western Atlantic species D. reticulatum
(Ridley and Dendy, 1886) and D. australe (see above), but
they lack trichodragmata.
Remarks on Dragmacidon. An additional species of
Dragmacidon (described as Pseudaxinella sp. in Alvarez
et al. 2000: 196) remains undescribed as no additional
material has yet been found to fully characterise the species.
The existing material is a fragment of a thin asymmetric
lamella, found detached from original substrate that does
not agree with the characteristically thickly encrusting shape
of Dragmacidon species.
A
C
B
Fig. 12. Phakellia tropicalis sp nov.: A, light microphotograph of
skeleton in cross section; B, schematic drawing of skeleton; C,
diagram of spicules. Scale bars: A, 500 µm; B, 100 µm.
Genus Phakellia Bowerbank, 1862
Gender feminine. Type species, by original designation,
Spongia ventilabrum Linnaeus, 1767. Recent, Lervig,
Norway, North Sea.
Phakellia tropicalis sp. nov
(Figs 7 E–F, 12 A–B)
Phakellia sp. Alvarez et al., 2000: 195; Holmes and
Blanch 2007: 761; Alvarez et al. 2007: 1600.
Material examined. H olotype – NTM. Z.5847
(Fig. 7e), Stevens Rock, West Arm, Darwin Harbour,
12°29.1667´S, 130°47.19´E, NT, 9 m depth, 8 May 2006,
coll. Alvarez, B. pArAtypes – NTM Z.5845 (Fig. 7F),
Stevens Rock, West Arm, Darwin Harbour, 12°29.1667´S,
130°47.19´E, NT, 9 m depth, 8 May 2006, coll. B. Alvarez.
AdditionAl specimens – Bynoe Harbour, NT:, NTM Z.4198,
Z.4486, Z.4488. Darwin Harbour, NT: NTM Z.866, Z.877,
Z.1948, Z.4197, Z.4428, Z.5665, Z.5839–Z.5842, Z.5844,
Z.5848, QM G303365, G303383. Wessel Is: NTM Z.4463.
29
Papua New Guinea: QM G312926, G312937. Malaysia:
NTM Z.5843.
Description. Shape (Fig. 7E–F). Small convoluted thin
fans, up to 250 mm high and 300 mm wide, on short and
thin peduncle, lexible, less than 5 mm thick, arranged in
multiple planes, with luted or planar langes, ragged or
crenulated margins.
Colour. Bright orange, beige-orange, or yellow-brown
(Darwin and Bynoe Harbour specimens) alive.
Oscula. Star-shape oscula, minute.
Surface. Velvety, felty, with choanosomal spicules
projecting shortly; marked with ine network of excurrent
channels ending in oscula and reinforced close to the
peduncle by thick choanosomal axes or ‘veins’.
Skeleton (Fig. 12A, B). A core of interwoven spicules,
occupying most of specimen’s thickness, laterally
compressed, echinated by a dense palisade of single
spicules, sometimes aggregated in loose brushes, protruding
through surface.
Spicules (Fig. 12C). Strongyles wavy, 222–800 in length
by 3–8 µm thick and styles 231–703 in length by 6–16 µm
thick (Table 5).
Remarks. (see Carvalho et al. 2007 for extensive review
of Phakellia species.)
B. Alvarez and J. N. A. Hooper
The species is atypical of Phakellia, although it agrees
with the diagnosis given by Alvarez and Hooper (2002) in
most aspects. It is fan-shaped, with styles projecting through
the ectosome, and includes the typical spicule composition
of the genus. However the reticulation of thick ‘veins’ or
thick axes observed in most species of Phakellia, seems to
be either incomplete or obscured by the habit of the species
here described.
The skeleton of this species (especially when seen in
cross section, as in Fig. 12A) resembles some species of
Acanthella (Dictyonellidae), a genus often confused with
Phakellia. Phakellia tropicalis however, lacks two of the
main diagnostic features of Acanthella – the cartilaginous
to membranous surface and the cavernous structure of the
choanosomal skeleton, with sheets of aspiculous collagen
joining the primary axes in the skeleton. Instead, the surface
of P. tropicalis is velvety, felty, with choanosomal spicules
projecting shortly through the surface, a character shared
with other axinellid genera such as Axinella and Cymbastela,
and the skeleton is formed by a core of interwoven strongyles
with a regular and dense palisade of erect styles that occupies
most of the thickness of the sponge.
A phylogenetic analyses based on morphological
characters by Alvarez et al. (2000) showed this species to be
closely related to other species of Acanthella (i.e. A acuta,
A. cavernosa and A. pulcherrima). In the same study,
however, an analysis based on 28S rDNA sequences with
the same set of species, indicated that P. tropicalis was
closely related to typical axinellid genera such as Axinella
and Dragmacidon supporting its allocation to Axinellidae
instead of Dictyonellidae.
Furthermore, the skeletal architecture of P. tropicalis
corresponds with the basic plan observed in species of
Bubaris Gray, 1867, a genus of the family Bubaridae and
currently used in the strict sense to include encrusting forms
(Alvarez and Van Soest 2002). Thus, one could interpret
the new species as an ‘erect Bubaris’ and be tempted to
allocate it to that genus. It is possible that species with erect
forms originally described under Bubaris, but transferred
to Acanthella and Phakellia in order to preserve the revised
concept of Bubaridae by Alvarez and Van Soest (2002),
might be related to the new species. If that is the case, those
species could be grouped under a new genus following a
revised version of Hentschel’s (1923) concept of Bubaridae,
which accepted sponges of erect forms, where the core
of interwoven strongyles echinated by styles is placed in
the centre (or in the axis) instead of at the base, as in the
encrusting forms. However, it would be premature to erect
a new genus here based on the characteristic of one single
species and without re-examining species currently hidden
under Acanthella or Phakellia which might also share such
characteristics and could justify the creation of the new
genus. In the absence of such evidence, we assign the new
species provisionally to Phakellia and propose to expand
the deinition of Alvarez and Hooper (2002) (and modiied
by Carvalho et al. 2007) as: Axinellidae of planar habit,
with skeleton formed by multiple axes or a single core of
sinuous megascleres (frequently strongyles), echinated
either by single spicules or by secondary tracts of a second
class of megascleres (frequently styles).
Phakellia tropicalis is the only species of Phakellia
recorded in this study and as far as we know the irst one
recorded from warm waters and shallow depths. Note that
some species currently accepted under Phakellia from
similar habitats are likely to be misidentiications (see
Carvalho et al. 2007 for an extensive review of Phakellia
species). Other species of Phakellia recorded from northern
Australia and adjacent areas are currently accepted under
Acanthella (Van Soest et al. 2008) or Axinella (i.e. Phakellia
aruensis Hentschel, 1912, see above). Yet other species
described under Phakellia in the Indian Ocean are not
typical of the genus: P. ceylonensis Dendy, 1905: 192 is an
Axinella (see above); P. crassistylifera Dendy, 1905: 192
is likely to belong in Stylissa and Phakellia ridleyi Dendy,
1887: 159 is currently accepted as a species of Phakettia.
Distribution. This species is found along the NT coast
and is very common in both Darwin and Bynoe harbours.
It is also recorded for Papua New Guinea and Malaysia. It
is found between 5–20 m depth
Etymology. Referring to its tropical habitat. It is
intended as a noun in apposition.
Genus Reniochalina Lendenfeld, 1888
Gender feminine. Type species, by subsequent
designation of Hallmann (1914), Reniochalina stalagmitis
Lendenfeld, 1888. Recent, Western Australia.
30
Reniochalina stalagmitis Lendenfeld, 1888
(Figs 7G–H, 13A–F, 14 A–E, 15)
Reniochalina stalagmitis Lendenfeld, 1888: 82;
Whitelegge 1902: 283; Hallmann 1914: 346; Hooper and
Wiedenmayer 1994: 81; Hooper and Lévi 1993: 1404;
Alvarez, et al. 2000: 197; Alvarez and Hooper 2002: 746;
Holmes and Blanch 2007.
Axinella echidnaea. – Ridley 1884: 462; Kieschnick
1896: 533; Hentschel 1912: 419 [misidentiication; not
Ridley and Dendy 1887: 183]
Reniochalina lamella Lendenfeld, 1888: 83; Whitelegge
1902: 283; Hallmann 1914: 346.
Axiamon folium Hallmann, 1914: 441 [objective
synonym, see Wiedenmayer (1989: 49) and Hooper and
Lévi (1993: 1403)]
Material examined. type mAteriAl – Reniochalina
stalagmitis: Lectotype, BMNH 1887.4.27.122, Western
Australia, Fig. 13A; paralectotype, AM G9004, wet, West
Australia, Fig. 13B [also holotype of Axiamon folium].
Reniochalina lamella: AM B5478, syntype, wet, no locality
data [also paratype of A. folium]. AdditionAl specimens –
Ridley’s (1884) material: BMNH 1882.2.23.261, Prince
of Wales Channel, Torres strait, North Queensland, coll.
HMS Alert; BMNH 1881.10.21.259, Thursday I., Torres
strait, North Queensland, coll. HMS Alert. Hentschel’s
Axinellidae from northern Australia
Table 5. Comparison of spicule dimensions among specimens of Phakellia tropicalis. Measurements in micrometres.
Specimen
Locality
Z.5847
Holotype, Stevens Rock
Z.4488
Bynoe Harbour
Z.4463
Wessel Is
G312926
Papua New Guinea
Strongyles
Styles
284.1–651.2 (480.4±105.3)
x 3.8–8.4 (5.7±1.2)
222.8–670.7 (435.5±135)
x 3.7–9.5 (7±1.6)
293.1–800 (553.4±134.6)
x 4.4–8.4 (6.6±1.1)
277.8–696.3 (476.4±117.6) [24]
x 4.2–8.4 (6.3±1.1)
231.9–549.2 (385.7±82.5)
x 7.3–430.9 (29.8±83.6)
353.9–703.6 (484.6±102.2)
x 9–16.4 (13.8±2)
273.6–658.2 (439.6±111.1)
x 8.3–16 (11.2±2.2)
239.6–490.6 (343.5±69.5)
x 5.9–11.5 (8.7±1.8)
(1912) specimen, SMF 1687, Aru-Inseln, bei Pulu Bambu,
Indonesia, 10 m depth, 3 April 1908, coll. H. Merton,
dredge. NW Shelf, WA: NTM Z.2358, Z.2361, Z.2273,
Z.724, Z.738. Ashmore reef, WA: QM G301093, G301112,
G301139. NE Joseph Bonaparte Gulf, QM G301202.
Fog Bay, NT, QM G303548. Bynoe Harbour, NT: NTM,
Z.4462 (0M9H2388-N), Z.5074 (0M9H2451-H), Z.5853.
Darwin Harbour, NT: QM G303329, G303362, G303374,
G303579, NTM Z.227, Z.285, Z.474, Z.483, Z.815, Z.1107,
Z.1989, Z.2686, Z.4448 (0M9H2005-V), Z.5854, Z.5855,
Gunn Point, NT, QM G303535. Parry Shoals, NT, Z.525.
Melville I., NT, NTM Z.608. Cobourg Peninsula, NT:
NTM Z.67, Z.135, Z.537, Z.565, Z.1335, Z.2527. Groote
Eylandt, NT, G313555. Wessel Is, Z.5075 (0M9H2647-T).
Gulf of Carpenteria, NT, QM G300817. Torres Strait, QM
G316882.
Description. Shape (Figs 7G–H, 13 A–B). Arborescent,
branching or fan-shaped, generally stalked, and erect with
specimens up to 60 cm high. Branches, lat to cylindrical,
dichotomous or fused, 1–2 cm diameter, generally with
pointed tips, dividing and anastomosing irregularly in
different planes.
Colour. Two colour forms, orange-red and beige-yellow.
Always brown in alcohol.
Oscula. Small, less than 5 mm diameter, with elevated
rims thin, membranous and transparent, irregularly
distributed through the branches.
Surface. Long surface processes or conules with
projecting spicules, up to 5 mm long, evenly distributed and
separated by more-or less longitudinal and parallel channels,
1 mm apart, 1 mm deep.
Skeleton (Fig. 13C–F). Specialised ectosomal skeleton
absent; choanosomal skeleton differentiated into axial and
extra-axial regions. Extra-axial skeleton reticulated with
ascending spongin ibres, 50–100 µm interconnected at all
angles by single spicules or short ibres, or anastomosing
and forming oval to round meshes up to 200 µm in
diameter; spongin fibres slightly developed and cored
with paucispicular tracts of spicules, sometimes plumose;
projecting into surface processes and becoming dense and
disorganised at tips. Axial skeleton slightly condensed and
reticulated as in extra-axial region.
Spicules (Fig. 14A–E; Table 6). Oxeas or anisoxeas,
occasionally modiied to styles, straight, bent or slightly
sinuous; with tips surmounted by microspines, which might
be rudimentary or absent; microspines at one end might be
half the compared size to those at other end (175–450 x 6–20
µm). Thin, sinuous styles or oxeas with smooth or slightly
spined ends, 156–288 x 3–6 µm (Fig. 14A–C), scattered
through the choanosomal skeleton, rare in most examined
specimens. Few long styles, projecting through ectosome,
present in some specimens but extremely rare.
Remarks. The thin and sinuous styles (Fig. 14) were
irst mentioned by Hallmann (1914) in his description
of Axiamon folium. The presence of these spicules was
conirmed in all the specimens examined here and point
out relationships of Reniochalina with other members of
the family Raspailiidae (order Poecilosclerida). Long and
slightly sinuous styles, most of which were broken, were
also observed in the type material of Axiamon folium (AM
G9004) and in some of the specimens examined. These
were extremely rare and scattered throughout the extraaxial skeleton and projecting through the surface, indicating
further afinities with raspailid taxa and challenging the
position of the genus within Axinellidae (see below).
Distribution. Reniochalina stalagmitis is one of the
most abundant sponge species of northern Australia. It is
found from the intertidal to depths of 60 m. Its distribution
extends well beyond the boundaries of the studied region
(Fig. 15) with validated records from the northern region of
WA (down to W Buccaneer Archipelago) and QLD (down
to the Howick Is region). It also occurs in Indonesia.
Notes on Reniochalina. Reniochalina was deined by
Alvarez and Hooper (2002) as ‘Axinellidae with extra-axial
spongin ibres projecting into surface processes and cored
with paucispicular tracts of oxeas, anisoxeas and styles.
Oxeas with tips surmounted by micro-spines’.
The genus was considered to be closely related to
other axinellid genera (i.e. Ptilocaulis and Phycopsis) by
Alvarez and Hooper (2002), based on the shared presence
of conspicuous, long, ilamentous surface processes and
their skeletal features. The close afinities with Ptilocaulis
were further conirmed by molecular phylogenetic analyses
(Alvarez et al. 2000). Recent molecular studies (Erpenbeck
et al. 2007; Holmes and Blanch 2007) also showed strong
afinities of Reniochalina stalagmitis with the raspaillid
species Axechina raspailoides. As previously indicated
(Hooper 1991; Hooper 2002b), these species have similar
growth forms and choanosomal skeletons, and it is now
confirmed that they also share the presence of styles
31
B. Alvarez and J. N. A. Hooper
Fig. 13. Reniochalina stalagmitis: A, lectotype, BMNH 1887.4.27.122; B, paralectotype, AM G9004; C, lectotype, BMNH1887.4.27.122,
light microphotograph of skeleton; and D, SEM of skeleton; E, paralectotype, AM G9004, light microphotograph of skeleton and F, SEM
of skeleton. Scale bars: A, B, 2 cm; C–F, 500 µm.
32
Axinellidae from northern Australia
with spined tips and identical shape (Fig. 14B, C), a fact
overlooked by previous authors with the exception of
Hallmann (1914).
Additional molecular analysis based on the CO1
fragment (Erpenbeck 2007) also indicated afinities of the
Caribbean species Ptilocaulis marquezi (Duchassaing and
Michelotti, 1864) with other raspailiid species (i.e. Pandaros
acanthifolium (Duchassaing and Michelotti, 1864) and
Ecyoplasia ferox (Duchassaing and Michelotti, 1864)
suggesting strongly that both Ptilocaulis and Reniochalina
are closely related to the Raspailiidae. These relationships
however, remain unresolved as neither Reniochalina nor
Ptilocaulis have the typical raspailiid ectosomal skeleton
which is clearly present in Axechina and other raspailiid
taxa. Unfortunately, the range of taxa sampled in the
molecular analyses mentioned above was inadequate to
conclude further on the afinities of the axinellid genera
Reniochalina and Ptilocaulis (and likely Phycopsis too)
with the family Raspailiidae. Therefore, if more evidence
from molecular analyses becomes available to support
the current results, these genera might be relocated to the
Raspailiidae.
Other species of Reniochalina reported in the literature
do not agree with the current deinition of the genus and
are referred here to more appropriate genera: Reniochalina
condylia Hooper and Lévi, 1993, to Dragmacidon;
R. plumosa Lévi and Lévi, 1983 to Axinella and, R. sectilis
Wiedenmayer, 1989 to Rhaphoxya in Dictyonellidae.
Additional species of Reniochalina including Reniochalina
sp., reported in Alvarez et al. (2000), remain to be described
from other areas of Australia (Hooper, pers. obs.). It is likely
also that some species of Reniochalina are misidentiied as
species of Ptilocaulis (e.g. P. rigidus Carter, 1883:322).
initial stages of this revision indicated that the Axinellidae
was represented in the area by a larger number of species.
This observation turned out to be contrary to what was found
after a thorough examination of recorded material. Many
of the species thought to belong to the Axinellidae were
in fact found to be members of other families and orders,
particularly Raspailiidae (Poecilosclerida).This relects
that separation of species within this group is extremely
subjective and could be erroneous if is taken in an isolated
context. Examination of the taxonomic characters across a
large number of specimens is critical to detect the variability
and plasticity of morphological characters present in this
group and to avoid incorrect splitting of taxa. It is possible
that cryptic species or hybrid forms are hidden within the
continuum of variability commonly observed in species of
Axinellidae; but only results of population genetic studies
can reveal such cryptic species, and indeed verify the
occurrence of hybridisation among the Porifera, as has been
demonstrated for the Cnidaria (e.g. Veron 1995, Van Oppen
2000 and references within).
The ive remaining genera of the Axinellidae (i.e. Auletta,
Dragmaxia, Pipestela, Ptilocaulis and Phycopsis) are
curiously not represented in the studied material despite
the fact that they do include tropical species.
Axinella loribellae, Cymbastela stipitata and
C. vespertina are the only species of axinellids reported
here with distributions restricted to northern Australia,
reflecting some degree of endemism in the area. The
latter sibling species pair represent western components
of east-west coast species pairs of the genus Cymbastela,
with C. coralliophila and C. concentrica of the east
coast, presumed remnants of Pleistocene separation of
northern Australian faunas during low strand sea levels
(e.g. Hooper and Ekins 2004). Axinella sinoxea displays a
similar distribution but can be found in deep waters of WA.
The remaining species seem to have a more widespread
distribution throughout Indonesia, Papua New Guinea and
Malaysia. Axinella aruensis and Reniochalina stalagmitis
are widely distributed in the northern region of Australia and
Indonesia. Dragmacidon australe has a disjunct distribution
with isolated records from New Zealand, GBR and northern
DISCUSSION
The results of this revision indicate that the Axinellidae
is represented in the area covered by this study by only ive
genera (i.e. Axinella, Cymbastela, Dragmacidon, Phakellia
and Reniochalina) and nine species, three of which are new.
Data gathered from NTM and QM collections during the
Table 6. Comparison of spicule dimensions among specimens of Reniochalina stalagmitis. Measurements in micrometres.
Specimen
Locality
Lectotype BMNH1887.4.27.122
WA
AM G9004
WA
AM B5478
WA
G303362
Darwin
Z.4462
Bynoe
Harbour
Wessel Is
Z.5075
Oxeas
Thin styles/oxeas
190–315 (248.6±27.9)
x 8–15.2 (12.1±2.4)
175–450 (243.9±61.8)
x 7.9–20 (10.7±3)
197–376.6 (248.3±49.5)
x 10.1–19 (14.4±1.9)
209.1–356.6 (253.3±43.4)
x 9.2–17 (13±2.2)
195.1–379.1 (293.5±39.1)
x 7.1–18.2 (13.9±2.6)
199.6–380.2 (286.4±44.6)
x 6.3–18 (12.4±2.8)
157.1–287.6 (240.2±35.3) [18]
x 2.6–5.8 (4.5±0.9) [18]
211.7–268 (244±22.7) [9]
x 2.6–5 (3.8±0.8) [9]
156.4–235 (203.7±25.3) [12]
x 3.5–6.3 (4.9±0.8) [12]
206.5–232.6 (219.5±18.4) [2]
x 3.9–4 (3.9±0) [2]
158.2–244.4 (213.2±40.2) [4]
x 2.6–5.2 (4.2±1.1) [4]
205.8x3.7 [1]
33
B. Alvarez and J. N. A. Hooper
A
C
B
D
E
Fig. 14. Reniochalina stalagmitis: A, diagram of spicules; B, C, SEM of sinuous style with spined tip (AM G9004); D, E, tip of oxeas,
(AM G9004). Scale bars: A, 50 µm; B, 5 µm; C, 20 µm; D, E, 2 µm.
Australia. Dramacidon durissimum is a species common
from Indian Ocean but its distribution is now extended to
the Ashmore Reef, WA of northern Australia.
The position of Reniochalina within the Axinellidae
is now debatable. The presence of sinuous styles with
spiny tips in R. stalagmitis and the recent evidence based
on molecular data (see above) suggest strongly that the
species is closely related to the raspailiid species Axechina
raspailioides (Poecilosclerida) and challenges the position
of Reniochalina stalagmitis within the family. The typical
ectosomal skeleton present in raspailiid species and
considered a synapomorphy for that family is absent in
R. stalagmitis, thus no deinitive conclusions can be made
at this stage. New studies as suggested above are needed
120˚ E
130˚
140˚
5˚
15˚ S
Fig. 15. Distribution of Reniochalina stalagmitis in northern Australia,
based on conirmed records from QM and NTM. Species distribution
extends along the Queensland coast (Hooper, unpublished data).
34
Axinellidae from northern Australia
to decide whether Reniochalina should remain classiied
under the Axinellidae.
Phylogenetic relationships within the family Axinellidae
and other related groups have been explored previously,
using either morphological, molecular or chemical characters
(Alvarez et al. 2000; Erpenbeck et al. 2002; Erpenbeck et al.
2005; Erpenbeck et al. 2006; Erpenbeck et al. 2007). The
relationships shown in those studies remain inconclusive
at this stage but suggest strongly that the Axinellidae is a
polyphyletic taxon with uncertain afinities. The taxonomic
revision of species of the Axinellidae and its sister groups
is critical to support conclusions derived from those studies
and is currently the focus of ongoing studies undertaken at
regional basis (e.g. Indonesia, Eastern Australia, CentralWest Paciic, Southern Australia and New Zealand).
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ACKNOWLEDGEMENTS
This work was funded by an Australian Biological
Research Studies (ABRS) research grant (Grant No 20510) and by the ‘Collection and Taxonomy of Shallow Water
Marine Organisms’ program for the US National Cancer
Institute (Contract N02-CM-27003) subcontracted to NTM
through CRRF.
We specially thank Michael Browne and Huy Nguyen,
for their invaluable assistance during 2002–2004 NTM ield
collections; Dr Pat Colin (CRRF) and Don DeMaria, for
their assistance and photographic work during NTM ield
collections in the 2004 Wessel Is; Terry Yumbuluy, Wessel
Is, to allow collections in his home-land area; Merrick
Erins (QM) for his assistance in interrogating the QM
database and making specimens available for study; Ellie
Hayward (Charles Darwin University) for her assistance
with SEM preparations; Drs Rob W.M. Van Soest (ZMA),
Richard Willan and especially Chris Glasby (NTM) for their
continuous advice and suggestions during the preparation
of this manuscript; and the two referees of this paper for
their valuable suggestions.
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Accepted 21 October 2009
37
B. Alvarez and J. N. A. Hooper
APPENDIX
Collection and locality data of material examined in the collections of QM and NTM.
QM material
G300181 Passage West I., outer reef, Ashmore Reef, WA, 12°14´S, 122°56´E, 15.5 m, 27 Jul 1986, coll. Hooper, JNA
G300295 Snake Reef, Howick Group, GBR, QLD, 14°27´S, 145°1´E, 12.5 m, 14 Dec 1990, coll. Hooper, JNA
G300609 N side of Cumberland Strait, Wessel Is, Gove, NT, 11°28´S, 136°29´E, 13 m, 14 Nov 1990, coll. NCI, AIMS
G300759 Marinbar I, SE Cape Wessel, Wessel Is, NT, 11°1.13´S, 136°46.04´E, 20 m, 17 Nov 1990, coll. NCI, AIMS
G300768 Gugari Rip 100m NE, E Guluwuru I, Wessel Is, NT, 11°34´S, 136°22.12´E, 8 m, 13 Nov 1990, coll. NCI, AIMS
G300817 Duyfken Point, W Gulf of Carpentaria, QLD, 12°34´S, 141°0´E, 58 m, 26 Nov 1991, coll. Cook, SD. on CSIRO RV
Southern Surveyor
G301089 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 14 m, 6 May 1992, coll. Hooper, JNA
G301092 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 22 m, 7 May 1992, coll. Hooper, JNA
G301093 Cartier I, outer reef slope, N side reef, WA, 12°31.07´S, 123°33.05´E, 22 m, 7 May 1992, coll. Hooper, JNA
G301112 Cartier I, outer reef slope, S side of reef, WA, 12°32.15´S, 123°33.12´E, 23 m, 8 May 1992, coll. Hooper, JNA
G301139 Hibernia Reef, entrance to lagoon, NE side reef, WA, 11°57.13´S, 123°22.06´E, 23 m, 10 May 1992, coll. Hooper, JNA
G301197 Flattop Bank, NE Joseph Bonaparte Gulf, NT, 12°16´S, 129°15´E, 32 m, 17 May 1992, coll. Hooper, JNA
G301202 Flattop Bank, NE Joseph Bonaparte Gulf, NT, 12°16´S, 129°15´E, 32 m, 17 May 1992, coll. Hooper, JNA
G303262 South Shell I., reef N of boat ramp, East Arm, Darwin Harbour, NT, 12°29.1334´S, 130°53.09´E, 0 m, 19 Sep 1993, coll.
Hooper, JNA and Hobbs, LJ
G303322 East Point Bommies, Darwin Harbour, NT, 12°24.08´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs,
LJ
G303329 East Point Bommies, Darwin Harbour, NT, 12°24.08´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and Hobbs, LJ
G303332 East Point Bommies, Darwin Harbour, NT, 12°24.0834´S, 130°48.14´E, 10 m, 23 Sep 1993, coll. Hooper, JNA and
Hobbs, LJ
G303362 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and
Hobbs, LJ
G303365 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and
Hobbs, LJ
G303374 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 23 Sep 1993, coll. Hooper, JNA and
Hobbs, LJ
G303383 Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 19 m, 24 Sep 1993, coll. Hooper, JNA and
Hobbs, LJ
G303388 Stevens Rock, West Arm, Darwin Harbour, 12°29.1667´S, 130°47.19´E, NT, 19 m depth, 24 Sep 1993, coll. Hooper, JNA
and Hobbs, LJ
G303444 Fish Reef, west side, Bynoe Harbour, NT, 12°26.01´S, 130°26.09´E, 11 m, 26 Sep 1993, coll. Hooper, JNA and Hobbs, LJ
G303535 Shoal Bay, W Gunn Point, NT, 12°9.15´S, 130°56.02´E, 14 m, 11 Oct 1993
G303548 Fog Bay, 1 nml E Point Blaze, NT, 12°54.15´S, 130°7.16´E, 7 m, 4 Oct 1993
G303579 Lee Point near Anglers Reef, Darwin Harbour, NT, 12°18.13´S, 130°52.14´E, 10 m, 11 Oct 1993
G304182 Granite Bluff, Lizard I., S headland Mermaid Cove, QLD, 14°39´S, 145°27´E, 18 m, 4 Apr 1994, coll. Hooper, JNA and
party
G304246 Cobia Hole, Mrs Watson´s Bay, Lizard I., QLD, 14°39.03´S, 145°26.15´E, 18 m, 5 Apr 1994, coll. Hooper, JNA and
party
G304253 Palfrey I., W side, Lizard I., QLD, 14°42.03´S, 145°26.09´E, 16 m, 6 Apr 1994, coll. Hooper, JNA and party
G310136 Parry Shoals 35nm W Bathurst I., NT, 11°7.03´S, 129°25.9´E, 16 m, 12 Aug 1987
G311873 100m NE Gugari Rip, East side Guluwuru IS, Wessel Is, NT, 11°20.4´S, 136°13.63´E, 8 m, 13 Nov 1990
G312926 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.05´S, 148°10.14´E, 20 m, 15 Dec 1996, coll.
Hooper, JNA
G312913 Coutance Islet, Kupiano, SE. Papuan Barrier Reef, Papua New Guinea, 10°14.0167´S, 148°6.14´E, 41 m, 14 Dec 1996,
coll. Hooper, JNA
G312935 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.0501´S, 148°10.14´E, 20 m, 15 Dec 1996,
coll. Hooper, JNA
G312937 12 mile sandbank, Kupiano, SE Papuan Lagoon, Papua New Guinea, 10°11.05´S, 148°10.14´E, 20 m, 15 Dec 1996, coll.
Hooper, JNA
G313555 S Groote Eylandt, NT, 14°27.1801´S, 136°14.29´E, 22.5 m, 12 Oct 1997, coll. Cook, SD. on CSIRO RV Southern
Surveyor
G316882 Torres Strait, QLD, 10°46.8´S, 142°15´E, 16.4 m, 19 Jan 2004, coll. TSMap_GM_01_2004 Gwendoline May
G320664 Munro Reef, Coral Sea, QLD, 14°18.15´S, 144°48.82´E, 23 m, 2 Jul 2003, coll. Hooper, JNA and party
38
Axinellidae from northern Australia
APPENDIX (continued)
Collection and locality data of material examined in the collections of QM and NTM.
NTM material
Z.67
Z.135
Z.227
Z.285
Z.474
Z.483
Z.525
Z.537
Z.565
Z.608
Z.615
Z.619
Z.630
Z.631
Z.632
Z.665
Z.724
Z.738
Z.815
Z.822
Z.866
Z.868
Z.877
Z.1107
Z.1335
Z.1363
Z.1388
Z.1948
Z.1961
Z.1989
Z.2156
Z.2246
Z.2249
Z.2273
Z.2284
Z.2304
Z.2310
Z.2322
Z.2331
Z.2345
Z.2358
Z.2361
Z.2402
Z.2511
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.50´S, 132°3.01´E, 17 Oct 1981, coll. Hooper, JNA &
Alderslade, PN
Sandy I. No.2, Cobourg Peninsula, NT, 11°5.50´S, 132°17´E, 10 m, 21 Oct 1981, coll. Hooper, JNA & Alderslade, PN
Lee Point, Darwin, NT, 12°19.0167´S, 130°53´E, 14 Nov 1981, coll. Hooper, JNA
Dudley Point Reef, East Point, Darwin, NT, 12°25.00´S, 130°48.01´E, 1 m, 18 Sep 1981, coll. Hooper, JNA & Murray, P
Fannie Bay, Darwin, NT, 12°25.00´S, 130°50´E, 9 Feb 1982, coll. Hooper, JNA
Fannie Bay, Darwin, NT, 12°25.00´S, 130°50´E, 9 Feb 1982, coll. Hooper, JNA
Parry Shoals 35nm W Bathurst I., NT, 11°7.03´S, 129°25.9´E, 1 m, 30 Apr 1982, coll. Hooper, JNA & Alderslade, PN
Port Bremer, Cobourg Peninsula, NT, 11°8.5´S, 132°18.8´E, 1 May 1982, coll. Hooper, JNA & Alderslade, PN
Sandy I. No.2, Cobourg Peninsula, NT, 11°5´S, 132°16.51´E, 14 m, 2 May 1982, coll. Hooper, JNA
Cootamundra Shoals, North of Melville I., NT, 10°49.07´S, 129°12.09´E, 31 m, 6 May 1982, coll. Thom, B & Lockyer, R
Cootamundra Shoals,North of Melville I., NT, 10°50.22´S, 129°13.17´E, 22 m, 10 May 1982, coll. Lockyer, R
Unnamed shoal N Melville I, NT, 11°38.23´S, 129°51.00´E, 24 m, 17 May 1982, coll. Thom, B & Lockyer, R
Unnamed shoal N Melville I, NT, 11°32.58´S, 130°02.50´E, 18 m, 25 May 1982, coll. Lockyer, R
Unnamed shoal N Melville I, NT, 11°32.58´S, 130°02.50´E, 18 m, 25 May 1982, coll. Lockyer, R
Unnamed shoal N Melville I, NT, 11°32.57´S, 130°2.51´E, 18 m, 25 May 1982, coll. Lockyer, R
NW Yampi Sound, NW Shelf, WA, 15°27.0334´S, 121°49.01´E, 76 m, 29 Apr 1982, coll. CSIRO R.V. Sprightly
N Adele I.,Collier Bay, NW Shelf, WA, 15°58.02´S, 122°39.07´E, 59 m, 21 Apr 1982, coll. CSIRO R.V. Sprightly
N Adele I.,Collier Bay, NW Shelf, WA, 15°58.02´S, 122°39.07´E, 59 m, 21 Apr 1982, coll. CSIRO R.V. Sprightly
Channel I., Middle Arm, Darwin, NT, 12°32.02´S, 130°51.02´E, 11 m, 16 Jul 1982, coll. Scott Chidgey (Caldwell Connell
Ass
Channel I., Middle Arm, Darwin, NT, 12°33.08´S, 130°51.04´E, 20 m, 18 Jul 1982, coll. Scott Chidgey (Caldwell Connell
Ass)
Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN.
Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN.
Channel I., Middle Arm, Darwin, NT, 12°32.07´S, 130°52.04´E, 13 m, 20 Aug 1982, coll. Alderslade, PN.
Dudley Point Reef, East Point, Darwin, NT, 12°25.00´S, 130°48.01´E, 22 Dec 1982, coll. Hooper, JNA
Table Head, Port Essington, Cobourg Peninsula, NT, 11°13.5´S, 132°10.51´E, 11 May 1983, 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
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2´S, 130°47.1´E, 27 Apr 1984, coll. Hooper, JNA
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.2´S, 130°47.1´E, 27 Apr 1984, coll. Hooper, JNA
West side of Weed Reef, Darwin, NT, 12°29.2001´S, 130°47.1´E, m, 11 May 1984, coll. Hooper, JNA and party
Northern tip of Weed Reef, outer reef slope, Darwin Harbour, NT, 12°29.2´S, 130°37.61´E, 5 Oct 1984, 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, 10 m, 12 Apr 1985, coll. Hood, C and party
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°31.00´S, 121°28.01´E, 38–40 m, 17 Apr 1985, coll. Russell, BC (TRASH Fish
project)
NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC
NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC
NW Lacepede Is, NW Shelf, WA, 16°34´S, 121°27.01´E, 40–46 m, 17 Apr 1985, coll. Russell, BC
Dudley Point Reef, East Point, Darwin, NT, 12°24.5´S, 130°48.01´E, 8 m, 29 Jul 1985, coll. Hooper, JNA
Coral Bay, Port Essington, Cobourg Peninsula, NT, 11°11.3´S, 132°3.71´E, 15 Sep 1985, coll. Hooper, JNA
39
B. Alvarez and J. N. A. Hooper
APPENDIX (continued)
Collection and locality data of material examined in the collections of QM and NTM.
NTM material
Z.2526
Z.2527
Z.2529
Z.2632
Z.2686
Z.2719
Z.2727
Z.3062
Z.3068
Z.3137
Z.3141
Z.3922
Z.3925
Z.3935
Z.3936
Z.3938
Z.3946
Z.3956
Z.4078
Z.4104
Z.4131
Z.4197
Z.4198
Z.4425
Z.4428
Z.4435
Z.4448
Z.4462
Z.4463
Z.4465
Z.4486
Z.4488
Z.4490
Z.4491
Z.5053
Z.5054
Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 18–20 m, 16 Sep 1985, coll.
Hooper, JNA
Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 18–20 m, 16 Sep 1985, coll.
Hooper, JNA
Orontes Reef,mouth of Port Essington,Cobourg Peninsula, NT, 11°3.60´S, 132°5.41´E, 17 Sep 1985, coll. Hooper, JNA
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
Dudley Point Reef, East Point, Darwin, NT, 12°24.5´S, 130°48.01´E, 3 Apr 1986, coll. Hooper, JNA and party
Myrmidon Reef,GBR, QLD, 18°10.00´S, 147°23´E, 15 m, 1 Jan 1985, coll. Wilkinson,CR
Parry Shoals, Arafura Sea, NT, 11°11.72´S, 129°43.26´E, 16 m, 12 Aug 1987, coll. Mussig, AM and NCI team
Parry Shoals, Arafura Sea, NT, 11°11.72´S, 129°43.26´E, 16 m, 12 Aug 1987, coll. Mussig, AM and NCI team
Parry Shoals, Arafura Sea, NT, 11°12.27´S, 129°42.71´E, 16 m, 14 Aug 1987, coll. Mussig, AM and NCI team
Parry Shoals, Arafura Sea, NT, 11°12´S, 129°43.01´E, 16 m, 14 Aug 1987, coll. Mussig, A.M. and NCI (AIMS)
Cumberland Strait, northern bay, Wessel Is, Gove Peninsula, NT, 11°27.5´S, 136°28.8´E, 20 m, 14 Nov 1990, coll. Hooper,
JNA
Cumberland Strait, northern bay, Wessel Is, Gove Peninsula, NT, 11°27.5´S, 136°28.8´E, 20 m, 14 Nov 1990, coll. Hooper,
JNA
N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper,
JNA
N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper,
JNA
N side of Cumberland Strait, Wessel Is, Gove Peninsula, NT, 11°27.60´S, 136°28.7´E, 32 m, 15 Nov 1990, coll. Hooper,
JNA
S W headland, Rimbija I., Cape Wessel, Wessel Is, Gove Peninsula, NT, 11°0.5´S, 136°43.79´E, 15 m, 16 Nov 1990, coll.
Hooper, JNA
N side Pugh Shoal, reef slope, NE of Truant I., English Company IS, Gove Peninsula, NT, 11°36.57´S, 136°53.39´E, 20
m, 18 Nov 1990, coll. Hooper, JNA
Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand
Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand
Near boat ramp, East Arm Port, Darwin, NT, 12º29.8´S, 130º53.5´E, coll. B. Glasby & party, by hand
“Town Hall” hole, SW Channel I., Middle Arm of Darwin Harbour, NT, 12°33.74´S, 130°51.67´E, 19.5 m, 9 Sep 2004,
coll. Alvarez, B
Sand Island, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°35.291´S, 130°52.264´E, 7 m, 9 Sep 2004, coll.
Alvarez, B
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.09´S, 130°47.1´E, 5–19 m, 8 May
2002, coll. Alvarez, B and party
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.09´S, 130°47.1´E, 5–19 m, 8 May
2002, coll. Alvarez, B and party
Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°33.09´S,
130°52.43´E, 4 –8 m, 6 May 2002, coll. Alvarez, B and party
Channel Island, 100–400 m N of bridge, Middle Arm, Darwin Harbour, NT, Australia, Australia, 12°33.09´S,
130°52.43´E, 4 –8 m, 6 May 2002, coll. Alvarez, B and party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 5–10 m, 26 May 2003, coll.
Alvarez, B and party
Rimbija I., 2.8 km W of Cape Wessel, Wessel Is, eastern Arnhem Land, NT, 11°00.21´S, 136°43.84´E, 17–20 m, 1 Apr
2004, coll. Colin, P
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.57´S, 136°17.86´E, 11–20 m, 5 Apr 2004, coll.
Alvarez B and party
Raft Point, Bynoe Harbour, NT, 12°37.69´S, 130°32.16´E, 5–8 m, 26 Jun 2003, coll. Alvarez, B and party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 5 m, 1 Jun 2005, coll. Alvarez, B
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 5 m, 8 May 2006, coll. Alvarez, B
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B
South Shell I., East Arm, Darwin Harbour, NT, 12°29.87´S, 130°53.12´E, 4–11 m, 18 Aug 2002, coll. Alvarez, B and party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll.
Alvarez, B and party
40
Axinellidae from northern Australia
APPENDIX (continued)
Collection and locality data of material examined in the collections of QM and NTM.
NTM material
Z.5055
Z.5057
Z.5058
Z.5059
Z.5064
Z.5065
Z.5066
Z.5067
Z.5068
Z.5071
Z.5072
Z.5074
Z.5075
Z.5662
Z.5665
Z.5816
Z.5817
Z.5818
Z.5819
Z.5820
Z.5821
Z.5822
Z.5823
Z.5824
Z.5825
Z.5826
Z.5827
Z.5828
Z.5829
Z.5830
Z.5831
Z.5832
Z.5835
Z.5836
Z.5839
Z.5840
Z.5841
Z.5842
Z.5843
Z.5844
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll.
Alvarez, B and party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll.
Alvarez, B and party
Raragala I., 700 m off NE tip Wessel Is, eastern Arnhem Land, NT, 11°32.85´S, 136°21.28´E, 13–16 m, 31 Mar 2004, coll.
Alvarez, B
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.55´S, 136°17.96´E, 25–30 m, 5 Apr 2004, coll.
Alvarez, B and party
Weed Reef, entrance to West Arm, Darwin Harbour, NT, 12°29.25´S, 130°47.54´E, 9–15 m, 3 Aug 2002, coll. Alvarez, B
and party
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.24´S, 130°35.56´E, 5–10 m, 23 May 2003, coll.
Alvarez, B and party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll.
Alvarez, B and party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.28´S, 136°17.52´E, 13–14 m, 5 Apr 2004, coll.
Alvarez B. and party
“Town Hall” hole, SW Channel I., Middle Arm of Darwin Harbour, NT, 12°33.74´S, 130°51.67´E, 10–18 m, 17 Sep 2002,
coll. Alvarez, B and party
Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.35´S, 130°31.45´E, 6–8 m, 11 Jun 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. Alvarez, B
Spencer Point, Indian I., Bynoe Harbour, NT, 12°35.49´S, 130°31.29´E, 9–10 m, 11 Jun 2003, coll. Alvarez, B and party
Raragala I., bay on SW coast, Wessel Is, eastern Arnhem Land, NT, 11°38.6´S, 136°17.84´E, 17–20 m, 30 Mar 2004, coll.
Alvarez, B and party
East Point, Darwin Harbour, NT, 12º24.16´S, 130º47.66´E, 11 m depth, 8 November 2008, coll. Ayling, A
East Point, Darwin Harbour, NT, 12º24.16´S, 130º47.66´E, 11 m depth, 8 November 2008, coll. Ayling, A
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.46´E, 3–12 m, 1 Jun 2005, coll. Alvarez,
B
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.459´E, 6 m, 27 Apr 2007, coll. Alvarez,
B
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12°42.21´S, 130°35.459´E, 6 m, 27 Apr 2007, coll. Alvarez,
B
East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.484´S, 130º 48.471., 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43´E, 14 m, 25 May 2007, coll. Alvarez, B
South Shell I., East Arm, Darwin Harbour, NT, 12°29.87´S, 130°53.14´E, 7–11. m, 19 Aug 2002, coll. Alvarez, B and
party
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 17 m, 8 May 2006, coll. Alvarez, B
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B
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
Stevens Rock, 1.25 km SE Talc Head, off Cox Peninsula, Darwin Harbour, NT, 12°29.071´S, 130°47.103´E, 10–15 m, 9
May 2002, coll. Alvarez, B and party
East Point, Fannie Bay, Darwin, Australia, 12º 24.48´S, 130º 48.47´E, 11 m, 7 Jun 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B
East Point, Fannie Bay, Darwin, NT, 12º 24.49´S, 130º 48.43., 14 m, 25 May 2007, coll. Alvarez, B
Mengalum I., off Kota Kinabalu, Malaysia, 6 10.87´N, 115 35.97´E, 10–13 m, 24 Oct 2005, coll. Alvarez, B
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
41
B. Alvarez and J. N. A. Hooper
APPENDIX (continued)
Collection and locality data of material examined in the collections of QM and NTM.
NTM material
Z.5848
Z.5853
Z.5854
Z.5855
Stevens Rock, Weed Reef, Darwin Harbour, NT, 12°29.17´S, 130°47.19´E, 14 m, 8 May 2006, coll. Alvarez, B
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12º42.2´S, 130º35.459´E, 3–12 m depth, 1 June 2005, coll.
Alvarez, B
Dawson Rock, 3 km SSE Rankin Point, Bynoe Harbour, NT, 12º42.2´S, 130º35.459´E, 3–12 m depth, 1 June 2005, coll.
Alvarez, B
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
42