Axinellida (Porifera :Demospongiae)
from the New Caledonia Lagoon
John N. A. ~ o o p e r *and Claude ~ P v i ~
Queensland Museum, P.O. Box 3300, South Brisbane, Qld 4101, Australia.
Laboratoire de Biologie des InvertCbrCs marins et Malacologie,
Museum national d'Histoire naturelle, 57, rue Cuvier, 75005 Paris, France.
A
Abstract
Sixteen species of axinellid demosponges, including seven new species and seven new locality records,
are described from the shallow-water New Caledonian lagoon and reefs [families Axinellidae (12 species,
Cyrnbastela, Reniochalina, Axinella, Phakellia, Stylissa, Ptilocaulis, Pseudaxinella, Rhaphoxya) and
Desmoxyidae (4 species, Myrrnekioderrna, Higginsia)],bringing the total number of described axinellid
species in this region to 25. Brief revisions are provided for several of these genera, based primarily
on the Indo-west Pacific fauna, in order to place these New Caledonian species. Non-endemic New
Caledonian axinellids belong predominantly to the north-eastern Australian (Solanderian province) and
Indo-Malay fauna, usually representing the easternmost extent of these species' distributions in the
Indo-west Pacific. Two species, Axinella carteri (Dendy) and Astrosclera willeyana Lister, were found
to be truly widely distributed throughout the Indo-Pacific, typically associated with coral reefs; other
previously suspected widely distributed species were generally found to be allopatric, cryptic sibling
species related to the tropical Australasian fauna.
Introduction
Prior to the present work, only 11 species belonging to the (polyphyletic) order Axinellida
had been described from the New Caledonian region: five in the family Axinellidae (four
of which are apparently endemic and one also known from the Indo-Malay archipelago),
one in Desmoxyidae (also known in northern New Zealand), one in Trachycladidae (also
known from New Zealand and south-eastern and south-western Australia), two in Agelasidae
[now considered t o be a separate order: Hartman 1982 (one of which is apparently endemic)],
and two hypercalcified 'sclerosponges' described by Vacelet (1981) which have also been
suggested as having possible affinities with the Agelasida (e.g. Vacelet 1981; van Soest
1984). However, these species represent only a small proportion of the axinellid fauna known
t o live in this region; furthermore, 6 of these 11 described species are strictly deeper-water
taxa, collected from the shelf and slope to the south of New Caledonia. Thus, the shallowwater axinellid sponge fauna is virtually undescribed, although many species have already
been collected by ORSTOM.
The present paper describes 16 species of shallow-water sponges in the families Axinellidae
and Desmoxyidae living in the lagoon and coral reef habitats surrounding the island.
These species represent some of the more visible axinellids in New Caledonian shallow-water
habitats, although there are still many more taxa yet to be described from this region (LCvi,
unpublished data).
J. N. A. Hooper and C. LCvi
1396
This is the second of a series of papers on the taxonomy of demosponges in the lagoon
and outer reefs of New Caledonia (Hooper and LCvi 1993); a third contribution will examine
the biogeographical relationships of the New Caledonian Microcionidae, Raspailiidae and
Axinellidae (Hooper and LCvi, unpublished data). These papers represent the results of a
collaborative investigation amongst several sponge biologists (C. Battershill, P. Bergquist,
J. Fromont, J. Hooper, M. Kelly-Borges, C. LCvi, J. Vacelet, C. Wilkinson), to document
the major components of this sponge fauna, arising from a series of workshops jointly
funded by ORSTOM and DITAC. The eventual aim of this investigation is to produce
a comprehensive taxonomic inventory of the islands' shallow-water sponges, and an
authoritative lay field guide to this fauna.
Unlike the previous study on Poecilosclerida (Hooper and LCvi 1993) for which there were
good contemporary revisions for most of the higher taxa, the axinellids have not been
substantially revised, and hence some level of revision was necessary for many of the genera
in order to place the New Caledonian species. These revisions, however, are certainly
incomplete and relatively cursory due to the unavailability of much of the relevant type
material required to undertake this considerable task.
Methods
Methods used to prepare and examine sponges for both light microscopy and scanning electron
microscopy follow the procedures described in Hooper (1991). Measurements are based on examination
of 25 random samples of each spicule category for each specimen, and are cited as range (and mean
in parentheses). Abbreviations used in the text are as follows: AIMS, Australian Institute of Marine
Science, Townsville; AM, Australian Museum, Sydney; BMNH, The Natural History Museum, London;
CMMI, Canterbury Museum, Marine Invertebrate collection, New Zealand; CSIRO, Commonwealth
Scientific and Industrial Scientific Organisation, Division of Fisheries, Hobart; DITAC, Commonwealth
of Australia Department of Industry, Technology and Commerce, Canberra; ICZN, International
Commission for Zoological Nomenclature (Anon. 1985); LFM, Merseyside County Museums, Liverpool
(formerly Liverpool Free Museum); MMBS, Mukaishima Marine Biological Station, Faculty of Science,
Hiroshima University, Onomichi; MNHN, MusCum National dYHistoireNaturelle, Paris; MHNG,
MusCum d'Histoire Naturelle de Genkve, Geneva; NCI, U.S. National Cancer Institute shallow-water
collection contract (Australian Institute of Marine Science, Townsville; primary collections in AIMS,
NTM and QM, duplicates and fragments in USNM); NMNZ, National Museum of Natural History,
Wellington; NMV, Museum of Victoria, Melbourne; N.S.W., New South Wales; NTM, Northern
Territory Museum of Arts and Sciences, Darwin; ORSTOM, Institut Frangais de Recherche Scientifique
pour le Developpement en Cooperation, Centre de Noumea; QFS, Queensland Fisheries Service; Qld,
Queensland; QM, Queensland Museum, Brisbane; SM, Strasbourg Museum, France; SMF, NaturMuseum und Forschungsinstitut Senckenberg, Frankfurt; USNM, National Museum of Natural History,
Smithsonian Institution, Washinton D.C.; ZMB, Zoologisches Museum fiir Naturkunde an der
Humboldt-Universitat zu Berlin, Berlin; ZMC, Zoologisk Museum, Copenhagen.
Systematics
Order AXINELLIDA Carter
Family AXINELLIDAE Ridley & Dendy
Definition
Axinellid sponges which generally lack microscleres, although several genera (Dragmaxia,
Dragmacidon, Dragmatella, Dragmatyle, Rhaphoxya, Raspaigella and Tragosia) may have
raphides, occurring singly or grouped into bundles (trichodragmata). Skeleton typically
divided into distinct axial and extra-axial components; main skeletal tracts, composed of
spongin fibres enclosing megascleres, usually condensed in axis; extra-axial region becoming
plumose or plumoreticulate towards surface. However, many exceptions to this pattern
shown in this family, and affinities of many species are still contested but poorly understood. Megascleres are monactinal, diactinal, or both, and sinuous spicules (usually
strongyles or oxeas) reappear throughout many genera. Axinellidae frequently have branching
growth form but funnel-shaped, flabellate, tubular, and massive forms also occur (modified
from Wiedenmayer 1989).
New Caledonian Axinellids
1397
Remarks
This family contains a large and heterogeneous assemblage of taxa, with about 50
nominal genera included, and is in urgent need of an extensive, detailed revision; hence, a
comprehensive definition of the family is provided here. The heterogeneity of the family
has been indicated by several authors (e.g. Bergquist and Hartman 1969; van Soest et al.
1990), and Hooper and Bergquist (1992) questioned whether the family was in fact monophyletic. Hooper et al. (1992) suggested that the Axinellidae should probably be subdivided
further, although they did not propose how this should be done. Many species show a
number of similarities with members of the order Halichondrida (skeletal structure, spicule
geometry), and indeed a recent revision of that order (van Soest et al. 1990) indicated that
the Axinellidae should be placed there. However, this hypothesis is premature and uncorroborated by any other evidence, having no greater or lesser support than the existing
arrangement. Thus, the higher systematic relationships of the family are still not clear and,
for the time being, the family is retained in a polyphyletic order, Axinellida.
Members of this large family occur from polar regions to the tropics, and from tidal
habitats to at least 4400 m depth (Hartman 1982). Many species of Axinellidae are known
to live in the New Caledonian region (LCvi, unpublished data), but so far only five of these
have been published from this region. These are: Ptilocaulis fusiformis LCvi, 1967, from the
Bay St Vincent (apparently endemic); Cymbastela cantharella (LCvi, 1983) from outer reef
slopes of New Caledonia (endemic); Axinella lifouensis LCvi & LCvi, 1983, from deeperwaters (> 350 m) to the south of the island (endemic); Phakellia columnata (Burton, 1928)
also from deeper-waters (>300 m) (also known from the Andaman Sea); and Reniochalina
plumosa (LCvi & LCvi, 1983), a new combination here, from deeper-waters (>400 m)
(endemic).
Genus Cymbastela Hooper & Bergquist
Pseudaxinyssa (in part).-Bergquist and Tizard, 1967: 190; Lkvi, 1983: 719.
Cymbastela Hooper & Bergquist, 1992: 103 [type species Pseudaxinyssa stipitata Bergquist & Tizard,
1967: 189 (holotype AM Z3101)].
Diagnosis
Typically stalked, cup-shaped, thinly lamellate growth form. Ectosome with or without
specialised skeleton of smaller oxeas. Choanosomal skeleton with compressed reticulate
axial region, in which major tracts run longitudinally through lamellae, and with gradually
ascending, diverging, radial, plumose or plumo-reticulate extra-axial region, in which tracts
become plumose and project through surface. Spongin fibres well developed, cored by oxeas,
frequently with telescoped points. Predominantly autotrophic, most species containing
symbiotic cyanobacteria. Oviparous (from Hooper and Bergquist 1992).
Remarks
This genus was recently erected for atypical species previously referrable to Pseudaxinyssa
Burton, including thinly lamellate, frequently cup-shaped, predominately coral reef species.
The genus is known only from Australasia, with seven species described including one
endemic to New Caledonia, C. cantharella. A second species from this region is described
below.
Cymbastela cantharella (LCvi)
(Figs 1-2)
Pseudaxinyssa cantharella Levi, 1983: 719-22, fig. 1, pl. 1.
Cymbastela cantharel1a.-Hooper and Bergquist, 1992: 119-20, figs 12-14, table 1 .
Material Examined
Holotype. MNHN DCL3141: outer reef, SW. coast New Caledonia, 22°20'S.,166013'E.,40 m
depth, coll. ORSTOM.
J. N. A. Hooper and C. LCvi
1398
New Caledonian material. ORSTOM R1261 (fragment QM G30001.5): stn 198, pinnacle S. of
Canyon Central, Chenal des Cinq Milles, 22"30.4'S.,166"45.1'E., 25 m depth, 15.ii.1978, coll.
G. Bargibant, SCUBA; ORSTOM R163: stn 116, SW. pointe, Baie de Prony, 22°21.8'S.,166049.9'E.,
40 m depth, 30.vi.1976, coll. P. Laboute, SCUBA.
Comparative material. Refer to Hooper and Bergquist (1992) for additional material from New
Caledonia.
Description
Colour. Pale orange-brown alive (Munsell 2.5-7.5 YR 8/10), beige in ethanol.
Shape. Short, erect cup-shaped or vasiform sponges, up to 150 mm high, 170 mm
maximum diameter, with relatively thick lamellae, up to 6 mm diameter, usually with
convoluted margins, occasionally with secondary cups or lamellae growing inside primary
cup, often with buttresses and exterior secondary projections, and with a short, cylindrical,
basal stalk, up to 40 rnm long, 17 mm diameter.
Surface. Predominantly smooth, with distinct interior (inhalant, osculiferous) and
exterior (smooth but uneven, buttressed) faces of lamellae. Oscules small, up to 2 mm
diameter, about 2 mm apart, each surrounded by slightly raised membraneous lip. Texture
firm, flexible, slightly compressible.
Ectosome. Membraneous, with heavy collagen, through which choanosomal oxeas
protrude, individually or in paucispicular plumose bundles, arising from ascending radial
tracts in peripheral skeleton.
Choanosome. Choanosomal skeleton plumo-reticulate, without axial compression or
axial and extra-axial differentiation. Two components of choanosomal skeleton present:
longitudinal (radial) spongin fibres run through lamellae, cored by multispicular tracts of
oxeas, plumose near periphery; transverse uni- or paucispicular tracts of oxeas interconnecting radial fibres; overall skeleton appears nearly disorganised, almost halichondroid.
Mesohyl with heavy collagen.
Megascleres. Oxeas short, slender, slightly curved, symmetrical, occasionally asymmetrical, tapering, fusiform, usually with slightly telescoped points. Length 143-(207.1)245 pm, width 2.5-(7.6)-12 pm.
Microscleres. Absent.
Distribution
Known only from the New Caledonia region.
Remarks
The description given above is mainly condensed from Hooper and Bergquist's (1992)
more comprehensive redescription of the species, as the present study includes only one
Fig. 1. Cymbastela cantharella (Lkvi): specimen ORSTOM R1261,
structural oxeas.
New Caledonian Axinellids
1399
previously unpublished specimen from the south-west New Caledonia lagoon. This species
has also been illustrated in more detail in this earlier work, and present illustrations are
provided for comparative purposes.
Cymbastela cantharella differs from other tropical species of the genus by its atypical
orange pigmentation in life (most other species are green and mauve), prominent surface
sculpturing on the inner (oscular) face of lamellae, having a dense radial-plumose skeleton,
with a secondary paucispicular secondary reticulate skeleton, together producing a nearly
halichondroid appearance, a radial-plumose ectosomal skeleton, and specific dimensions of
its oxeas (refer to Hooper and Bergquist 1992: table 1). The species is most similar to
C. stipitata, both having thinly flabellate, irregular, buttressed surface processes, and similar
geometry of oxea megascleres, but the two species differ markedly in most other features.
Fig. 2. Cymbastela cantharefla (Lkvi): A, specimen ORSTOM R1261 in situ
(photo C . Debitus); B, specimen ORSTOM unregistered (stn 163/1) in situ (photo
P . Laboute); C, skeleton [specimen QM G300015 (ORSTOM R1261)l (scale=
500 pm); D, peripheral skeleton (scale=200 pm); E, SEM skeletal structure
(specimen QM G300004) (scale = 500 pm).
1400
J. N. A. Hooper and C. LCvi
Cymbastela concentrica (Lendenfeld)
(Figs 3-5, Table 1)
Antherochalina concentrica Lendenfeld, 1887: 788, pl. 22, fig. 42.
Cymbastela concentrica. -Hooper and Bergquist, 1992: 114-19, figs 9-1 1, table 1.
Material Examined
Lectotype.
known.
AM 21993: Port Molle (now Airlee Beach), Qld, 20°13'S.,148049'E., no other details
New Caledonian material. QM G301229 (ORSTOM R180): stn 109, Baie des Citrons, Noumea,
22°18~0'S.,1660251'E., 10 m depth, 31.v.1976, coll. P. Laboute, SCUBA; QM G301230, G301264,
G301266 (ORSTOM R153): stn 180, S. of sand cay, Ilat l'Areignere, 22°20~0'S.,1660191'E., 12 m
depth, 4.v.1977, coll. P. Laboute, SCUBA; ORSTOM R363: stn 150, S. of entrance, Baie St Vincent,
22"02.2'S.,165"59.5'E., 18 m depth, 9.ix. 1976, coll. P. Laboute, SCUBA; ORSTOM 'cfR363': stn 148,
NE. point, Ilat Mboa, 22"08.3'S.,166"09.3'E., 13 m depth, 9.ix.1976, coll. P. Laboute, SCUBA;
ORSTOM 'cfR363': stn 154, channel between Ilat Puen and RCcif, 21°59~0'S.,165"57l1E., 14 m
depth, 15.ix.1976, coll. P. Laboute, SCUBA; QM G301272, G301268, (ORSTOM RHO), ORSTOM
R181: stn 110, SE. Il6t RCdika, 22°31~1'S.,166036.0'E., 15-19 m depth, 3-8.vi.1976, coll. P. Laboute,
SCUBA; QM G301330: Croisant-Larigriere, Ilat Maitre, off Noumea, 22"20.2'S.,166"225'E., 20 m
depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA.
Comparative material. QM G301233: Davies Reef, Great Barrier Reef, Qld, 18"501S.,147"3g1E.,
22 m depth, coll. C. R. Wilkinson, AIMS, 24.iii.1982, SCUBA; QM G301234: same locality, 15 m
depth, coll. S. Seddon, 13.ix.1991, SCUBA; AIMS R.442-PS (NTM 22729): Myrmidon Reef, Great
Barrier Reef, Qld, 18°10'S.,147"23'E., 15 m depth, l.i.1985, coll. C. R. Wilkinson, AIMS, SCUBA;
NTM 23170: Blue Lagoon, Lizard I., Great Barrier Reef, Qld, 14°40'S.,145028'E., 10-20 m depth,
1.i.1987, coll. A. W. D. Larkum, SCUBA. [Refer to Hooper and Bergquist (1992) for additional
material from eastern Australia.]
Description
Colour. Pale beige, olive-brown or reddish brown alive (Munsell 2.5Y 8/6-2.5YR
4/4); chlorophyll pigments present.
Shape. Growth form predominantly vasiform, but varying from more-or-less symmetrical
cup-shaped with small basal stalk, to vasiform with symmetrical or asymmetrical lamellae,
Table 1. Comparison in spicule dimensions between known specimens of
Cymbastela concentrica
All measurements given in micrometres, and expressed as minimum-(mean)maximum range of measurement (comparative data from Hooper and
Bergquist 1992). N = 2 5 for each specimen
Material
(Locality)
AM 21993 (holotype)
(Airlee Beach, 20"s.)
NTM 23169
(Lizard I., 14"s.)
QM G301233
(Davies Reef, 18"s.)
QM G301234
(Davies Reef, 18"s.)
ORSTOM R180
(Noumea lagoon, 22's.)
ORSTOM R153
(Noumea lagoon, 22's.)
QM G300003
(Moreton I., 26"s.)
Oxeas
Length
Width
172-(239.6)-305
2.5-(9.5)-16
103-(145.6)-172
2.5-(3.3-6
138-(165.0)-184
4.0-(5.2)-6.0
92-(118.6)-146
3.5-(5.2)-8.0
107-(126.4)-142
3.0-(4.1)-5.0
96-(115.2)-132
3.0-(4.9)-6.5
67-(86.8)-104
2.5-(3.4)-4.5
New Caledonian Axinellids
LATITUDE
( O S )
Fig. 4. Cymbastela concentrica (Lendenfeld), comparison of spicule
dimensions with latitudinal distribution of samples: A, mean length;
B, mean width.
to thickly encrusting plate-like, attached directly to substrate. Size up to 150 mm high,
140 mm maximum width. Lamella thickness variable, ranging from card thin to thick and
rubbery, 1.0-3.5 mm thick.
Surface. Typically with convoluted, multiple lamellae inside cups or with digitate
projections on exterior surface, but some specimens lack any surface ornamentation.
Lamellae smooth, even or irregular. Texture flexible, compressible, velvet-like.
1402
J. N. A. Hooper and C. LCvi
Ectosome. Membraneous without specialised skeleton, but microscopically villose from
protruding spicules from peripheral skeleton, usually forming multispicular plumose brushes.
Choanosome. Reticulate to plumo-reticulate mineral and fibre skeleton, with poorly
differentiated axial and extra-axial regions. Reticulate skeleton predominant over plumose
portion. Fibres in axial region only slightly condensed, forming an open reticulation, cored
by uni- or paucispicular tracts of spicules. Extra-axial fibres reticulate, slightly plumose,
paucispicular, whereas peripheral skeleton clearly diverges into plumose multispicular spicule
bundles. Spongin fibres well developed; collagen abundant in mesohyl.
Fig. 5 . Cymbastela concentrica (Lendenfeld): A , peripheral skeleton [specimen QM
G301229 (ORSTOM R180)] (scale=200 pm); B, specimen ORSTOM R180 in situ (photo
P . Laboute); C , specimen QM G301230 (ORSTOM R153) in situ (photo G. Bargibant);
D, specimen ORSTOM 'cfR153' in situ (photo G. Bargibant); E, preserved specimen
QM G301229; F, SEM skeletal structure (QM G301230) (scale=200 pm).
New Caledonian Axinellids
1403
Megascleres (refer to Table 1 for dimensions). Oxeas vary considerably in size between
specimens, usually slender, fusiform, straight or slightly curved, symmetrical, typically with
very faintly telescoped points.
Microscleres. Absent.
Distribution
Northern, central and southern Queensland, and south-west New Caledonia, found in the
lagoon, inshore fringing reef or platform coral reef fauna, 10-22 m depth.
Remarks
This is a new locality record for this species in the New Caledonian region. Although
C. concentrica has recently been comprehensively redescribed and illustrated (Hooper and
Bergquist 1992), the description given above is based on a number of previously undescribed
specimens from both the Great Barrier Reef and New Caledonia. The present records,
together with material described by Hooper and Bergquist (1992), show that the species is
widespread throughout all sections of the Great Barrier Reef, and it is possible that the
species will eventually be found to inhabit many other islands and reefs throughout the
Coral Sea.
This species is remarkable for the heterogeneity in its spicule dimensions, unlike any
of the other known species of the genus. Hooper and Bergquist (1992) suggested that this
variability may correspond to latitudinal gradients in distribution of the species, although
re-examination of spicule dimensions from all known material (Fig. 4) shows no such trend.
It is possible that we have two or more sympatric sibling species presently included in
C. concentrica but present morphometric characters cannot clearly distinguish these
morphotypes.
Genus Reniochalina Lendenfeld
Reniochalina Lendenfeld, 1888: 82.-Hallmann, 1914b: 346; Wiedenmayer, 1989: 48 [type species
Reniochalina stalagmitis Lendenfeld, 1888: 82 (lectotype BMNH 1887.4.27.122)l.
Axiamon Hallmann, 1914~:440. -de Laubenfels, 1936: 130 [type species Axiamon folium Hallmann,
1914c: 440 ('syntypes' AM G9004, B5478)I.
Diagnosis
Arborescent, frondose, ramose or lobate growth forms; typically with tubercular, ridged
or closely conulose surface; with plumo-reticulate skeletal structure, mainly in the form of
ascending skeletal bundles, and without condensation of axial skeleton or differentiation
between axial and extra-axial components; spiculation with interchangeable styles, oxeas
and anisoxeas in approximately equal proportions and similar size, without differential
distribution within the skeleton (modified from Wiedenmayer 1989).
Remarks
The above diagnosis is for the nominal genus Axiamon, since Lendenfeld's diagnosis of
Reniochalina is grossly inadequate. Furthermore, this genus is best known under its junior
synonym of Axiamon, and further comment is required on that synonymy. Hallmann
(1914b: 346) noted that the primary description of ~endenfeld'sReniochalina stalagmitis
was erroneous; consequently, Reniochalina was considered to be a genus dubium. Hallmann
(1914~:440-1) subsequently described a specimen under the name of Axiamon folium (from
Western Australia), which became the type species of the genus Axiamon, whilst at the same
time he recognised that it was probably identical to Lendenfeld's R. stalagmitis (from the
Illawarra region, N.S.W.). De Laubenfels (1936: 47) concurred with Hallmann. However,
although Lendenfeld's (1888) definition of R. stalagmitis was not entirely accurate, differing
significantly from the actual characteristics of the species (Whitelegge 1902: 283), it is still
true that Lendenfeld's published name has priority since it is the earliest available for the
species (Wiedenmayer 1989): hence, Axiamon is an objective synonym of Reniochalina.
1404
J . N. A. Hooper and C. Ltvi
Prior to the present study, only four nominal species of the genus were known, but of
these species only two are valid. Two other species are also referred to the genus in the
present study. The genus now contains four valid species, all from the Indo-west Pacific
region: R. condylia, sp. nov., from the shallow-water New Caledonian fauna (see below)
(Figs 7-8); R. plumosa (LCvi & LCvi, 1983), comb. nov. (holotype MNHN DCL2972), from
the deeper-water New Caledonian fauna (Fig. 6C-D); R. sectilis Wiedenmayer, 1989: 49,
pl. 4, fig. 1, pl. 23, fig. 1 (holotype NMV F51962), from south-east Australia (not figured
here); and the type species R. stalagmitis Lendenfeld, 1888, from north-west and north-east
Australia (Fig. 6A-B) [with synonyms R. lamella Lendenfeld, 1888 (holotype AM B5478
from Western Australia), and Axinella echidnaea sensu Ridley (1884; specimen BMNH
1882.2.23.261 from north-west Australia) and Hentschel (1912; specimen SMF1687 from
southern Indonesia) not Spongia echidnaea Lamarck, 1814 (Wiedenmayer 1989)]. Recent
collections from tropical Australasia also discovered another six undescribed species referable
to this genus, which will be described elsewhere in the future.
Reniochalina is characterised by its plumo-reticulate skeletal architecture, and in having
anisoxeas, sometimes with spinose extremities. The skeletal architecture of R. stalagmitis is
certainly atypical of Axinellidae, and the existence of such structures makes the task of
satisfactorily diagnosing the family a difficult one, but that feature is certainly not unique
in the family (e.g. Phycopsis). Reniochalina should also be compared to Axinosia in skeletal
construction, and to Ptilocaulis, from which it differs mainly in lacking axial skeletal
condensation, or differentiation between axial and extra-axial skeletons, and also in lacking
telescoped endings on spicules. Wiedenmayer (1989) suggested that the variability in spiculation known to occur in Reniochalina (e.g. Hallmann 1914c: 443), and the confused skeletal
architecture in some species (e.g. Wiedenmayer 1989), raises doubts about which characters
should be treated as most important in the systematics of axinellid genera (spicule geometry
or skeletal structure) since these appear to occur in all combinations (cf. Axinosia and
Pseudaxinella, for example). It is clear that an evaluation of those characters in the
taxonomy of these genera is central to a revision of Axinellidae, but a satisfactory resolution
of these systematics will probably not be possible without additional (non-skeletal) evidence.
Fig. 6. Reniochalina spp.: A, Reniochalina stalagmitis Lendenfeld, peripheral skeleton
(leitotype BMNH 1887.4.27.122) (scale= 500 pn);B, specimen NTM 21107 in situ (photo
J. N. A. Hooper); C, Phakellia plumosa Ltvi and LCvi, peripheral skeleton (paratype
MNHN DCL2974) (scale =200 pm); D, paratype.
New Caledonian Axinellids
1405
If the evidence cited for Reniochalina were to be carried over to other genera, (viz. the
unstable nature of axial and extra-axial differentiation, and the presence or absence of
certain categories of megascleres), then many of the nominal axinellid genera would have
to be merged.
Reniochalina condylia, sp. nov.
(Figs 7-8)
Material Examined
Holotype. QM G300020 (ORSTOM R1223): stn 184, SE. of Ilbt Ua, New Caledonia lagoon,
22'42. l'S., 166"49+WE.,16 m depth, 8.vi.1977, coll. G. Bargibant, SCUBA.
Description
Colour. Pale orange alive (Munsell 7.5YR 8/10), pale grey-brown in ethanol.
Shape. Thickly encrusting plate, 260 mm diameter, 10-35 mm thick, with prominent
low, conical-bulbous, digitate projections on upper surface; bulbous digits 14-23 mm high,
10-14 mm maximum diameter, each with a single, large osculum on apex, 3-5 mm diameter.
Surface. Membraneous (where intact in preserved condition), even, porous upper
surface, without ornamentation other than large bulbous digits. Subdermal spicule bundles
clearly visible below translucent dermal membrane.
Ectosome. Without specialised spiculation; with plumose brushes composed of choanosoma1 megascleres protruding slightly through surface, up to about 80 pm from surface, but
presumably these are confined to within the dermal membrane when intact. Mesohyl
in peripheral skeleton with heavier, granular collagen as compared with the deeper
choanosomal region.
Choanosome. Skeletal structure plumo-reticulate, without axial compression or differentiation between axial and extra-axial regions. Skeleton divided into ascending, multispicular, diverging primary skeletal tracts, up to 150 pm diameter, with well-developed
plumose brushes at surface, and transverse, paucispicular, secondary spicule tracts, 2040 pm diameter, more-or-less interconnecting primary tracts. Skeletal meshes 150-300 pm
diameter, with few loose spicules between tracts. Spongin fibres very lightly invested with
Fig. 7. Reniochalina condylia, sp. nov., holotype QM G300020: A, oxeas, styles and
modifications; B, section through peripheral skeleton.
1406
J. N. A. Hooper and C. Levi
spongin, seen clearly only between major spicule tracts. Choanocyte chambers oval, 80170 pm diameter. Mesohyl with light collagen.
Megascleres. Single category of structural megasclere present, varying from oxeas to
styles, with various intermediate forms also present. Spicules consist of predominantly
slightly curved, slightly asymmetrical oxeas; less frequently styles with evenly rounded bases
and slightly curved towards basal end; rarely asymmetrical anisoxeas. Points mostly fusiform,
tapering, sharply pointed; occasionally fusiform rounded. Length 208-(259.8)-289 pm,
width 10-(12.2)-14 pm.
Microscleres. Absent.
Distribution
Known only from the south-west lagoon of New Caledonia, 16 m depth, on coral
rubble substrate.
Remarks
The present species is most similar to R. stalagmitis in its skeletal structure (plumose
spicule bundles) and spicule geometry (oxeas, styles and anisoxeas), although in
R. stalagmitis many spicules have spinous extremities, particularly the anisoxeas.
Reniochalina condylia, sp. nov., is also very similar to two undescribed species of
Reniochalina from western and north-western Australia (Reniochalina provisional species
numbers 353, 798), having virtually identical skeletal architecture and spicule geometry, but
in both cases these undescribed species have quite different external morphology and surface
ornamentation, and there are also differences in the specific dimensions of megascleres
(broader and shorter in the two undescribed species). These new species will be contrasted
further with other described species in a future revision of Reniochalina.
The deeper-water R. plumosa from New Caledonia has a plumo-reticulate skeleton
composed mostly of ascending discrete skeletal columns, cored by sharply pointed
oxeas incompletely separated into two size categories, 112-(284.21-372 x 9-(11.0)-15 pm
(Fig. 6C).
Etymology
Named for the conical digitate surface projections; Lat. condylus, knob, prominence.
Fig. 8. Reniochalina condylia, sp. nov.: A, SEM skeletal structure [holotype QM G300020 (ORSTOM
R1223)l (scale=500 pm); B, holotype; C, holotypeh situ (photo P. Laboute).
New Caledonian Axinellids
1407
Genus Axinella Schmidt
Axinella Schmidt, 1862: 60.-Gray, 1867: 513; Ridley and Dendy, 1887: 178; Dendy, 1905: 193;
Dendy, 1922: 114; Vosmaer, 1912: 308,318; Topsent, 1928: 37-8; Bergquist, 1970: 14; Bergquist,
1978: 167, 192; Lbvi, 1973: 605; Pansini, 1983: 79-98 [type species Axinellapolypoides Schmidt,
1862: 62 (holotype possibly SM, 'schizotypes' MNHN DCL1148L, BMNH 1867.7.26.81) (de
Laubenfels 1936)l.
Chalinissa Lendenfeld, 1887: 771.-Burton, 1927: 502 (type species Isodictya dissirnilis Bowerbank,
1866) (de Laubenfels 1936).
Astrospongia Gray, 1867: 514 (objective synonym of Axinella).
Diagnosis
Variable growth form, ranging from digitate to flabellate; surface typically hispid,
conulose; choanosome always with some axial compression o f spongin fibres and spicules;
with or without differentiated primary and secondary fibre elements; extra-axial skeleton
plumose or plumo-reticulate, diverging single or bundles o f spicules; ectosome without
specialised spiculation, although extra-axial megascleres usually protrude through surface;
megascleres oxeas, styles, occasionally strongyles, in various combinations; microscleres may
include raphides or microraphides, although these do not appear to be widely distributed
within the genus [compiled from Vosmaer (1912) and Donadey et al. (1990)l.
Remarks
Schmidt (1862) characterised A. polypoides in having a peculiar distribution o f oscules,
for which Gray (1867) erected Astrospongia, but Vosmaer (1933, 1935a, 1935b) correctly
surmised that the character was unlikely to be o f much systematic importance, and in any
case he noted that it also occurred in other axinellid genera. Other authors who have dealt
with this genus (e.g. Ridley and Dendy 1887; Thiele 1903; Vosmaer 1912, 1935a; Dendy
1922; Babic 1922; Topsent 1934; Vacelet 1961, 1969; Bergquist 1970; Pansini 1983) agree
that it is still problematic, with no completely satisfactory or clearly discriminatory diagnosis
yet developed. Part o f the problem arises from the existence o f species with predominantly
stylote spiculation (e.g. Dendy 1922), whereas typically the genus has both oxeas and styles
o f equal size and proportion (e.g. Vosmaer 1935a; Wiedenmayer 1989). Vosmaer (1935a)
and Pansini (1983) suggest further that intra-specific variability is known for the type
and other nominal Axinella species, whereby both oxeas and styles may be modified to
anisoxeote or anisostrongyle forms, in some cases almost completely. Consequently, the
placement o f species with a reticulate axially condensed skeleton, and reticulate to plumoreticulate extra-axial skeleton o f oxeas and styles in Axinella is relatively easy (e.g.
A . polypoides; Fig. lOC), whereas species such as A . spiculifera (Lamarck), A . profunda
Ridley and Dendy, and A . erecta Carter have greatly modified spiculation, and could be
easily placed in genera such as Phakellia or Reniochalina.
The literature contains records o f 39 species from the Australasian region (including
New Zealand, New Caledonia, Papua New Guinea and southern Indonesia), that have been
referred to Axinella at one time or another but, o f these, only 18 appear to be correctly
placed here. The type material o f many o f these species has not yet been located, so a
specific revision o f the Indo-west Pacific fauna in this genus is not yet possible. However,
valid species o f Axinella from this region are presently thought to be: Phakellia aruensis
Hentschel, 1912 (holotype SMF 953; Fig. 9A-B), from southern Indonesia, north-west
Australia around to the Great Barrier Reef (Hentschel 1912; Bergquist et al. 1980; Hooper
et al. 1992); Axinella australiensis Bergquist, 1970 (holotype NMNZ Por .18 [not seen]),
from New Zealand; Axinella brondstedi Bergquist, 1970 (holotype probably ZMC [not
seen]),with synonym Axinella verrucosa sensu Brondsted, 1923 (preocc.) from New Zealand;
Acanthella carteri Dendy, 1889, from New Caledonia (LCvi 1979) (Fig. 11-12; see below);
Axinella clathrata Dendy, 1897a (holotype NMV G2330 [not seen]), from south-east
Australia; Phakellia crassa Carter, 1885 (holotype BMNH 1886.12.15.129), from southeast Australia (Fig. 9C-D); Acanthella euctimena Hentschel, 1912 (holotype SMF 1012;
Fig. 9E-F) from southern Indonesia; Axinella Iifouensis LBvi & LBvi, 1983 (holotype
1408
J. N. A. Hooper and C. Levi
MNHN DCL2943; Fig. 9G-H), from deeper-waters off New Caledonia; Axinella globula
Brondsted, 1924 (holotype probably ZMC [not seen]), from New Zealand (Bergquist 1970);
Axinella kirkii Dendy, 1897a (lectotype NMV G2370 [not seen]), from south-east Australia;
Axinella meloniformis Carter, 1885 (holotype BMNH 1886.12.15.117; Fig. 91-J), from
south-east Australia (considered atypical of the genus [Vosmaer 19121, but skeletal structure
is most similar to Axinella of all other genera); Axinella retepora (Lendenfeld, 1887)
(lectotype BMNH 1886.8.27.417; Fig. 10A-B) from Torres Strait, Qld (not Port Phillip,
Vic., as published); Axinella richardsoni Bergquist, 1970 (holotype NMNZ Por. l9 [not
seen]), from New Zealand; Axinella sinclarii (Gray) (holotype unknown) from New Zealand
(Dendy 1897b; Bergquist 1970); Axinella torquata Brondsted, 1923 (holotype probably ZMC
Fig. 9 . Axinella spp.: A , Phakellia aruensis Hentschel, in situ (specimen NTM 22156)
(photo J. N. A. Hooper); B, skeleton (holotype SMF 953) (scale=500 pm); C, Phakellia
crassa Carter (holotype BMNH 1886.12.15.129); D, skeleton (scale = 500 pm); E, Acanthella
euctimena Hentschel (holotype SMF 1012); F, skeleton (scale = 500 pm); G, Axinella
lifouensis Ldvi and L6vi (holotype MNHN DCL2943); H, skeleton (scale=500 pm); I,
Axinella meloniformis Carter (holotype BMNH 1886.12.15.117); J, skeleton (scale = 500 pm).
New Caledonian Axinellids
1409
[not seen]), from New Zealand (Bergquist 1970); Axinella tricalyciforrnis Bergquist, 1970
(holotype CMMI 3/63 [not seen]), with synonym Axinella larnellata sensu Bergquist, 1961,
from New Zealand; and Axinella villosa Carter, 1885 ('syntype' BMNH 1886.12.15.396;
Fig. lOD), from south-east Australia (Dendy 1897a). There are another 28 possibly
undescribed species collected from tropical Australasia (Hooper, unpublished data), which
are also probably most appropriately placed in this genus. These will be described in future
revision of the tropical Australasian axinellids.
Other Australasian species described in Axinella but better placed elsewhere include:
Axinella aurantiaca Lendenfeld (1888, 'syntypes' AM G9171,Z468) from south-east Australia
belongs with Bubaris (Bubaridae) (Whitelegge 1889, 1901; Hallmann 1914a; Burton 1928);
Axinella arborescens Ridley & Dendy, 1886 (holotype BMNH 1887.5.2.64), from south-east
Australia belongs in Hornaxinella (Whitelegge 1889, 1907; Hallmann 1914~;Vacelet et al.
1976); Axinella atropurpurea Carter, 1885 (holotype BMNH 1886.12.15.I), from southeast Australia is a species of Raspailia (Raspailiidae) (Hooper 1991); Axinella axifera
Hentschel, 1912 ('syntype' SMF 1666), from southern Indonesia and north-west Australia
is a species of Ceratopsion (Raspailiidae) (Hooper 1991); Axinella colvilli Brondsted, 1924
('syntypes' probably in ZMC), from New Zealand belongs to Ciocalypta (Halichondriidae)
(Bergquist 1970); Axinella chalinoides et varr. Carter, 1885 (lectotype BMNH 1886.12.15.401),
and Axinella cladoflagellata Carter, 1886a (holotype BMNH 1886.12.15.407), from southeast Australia, are synonymous and belong to Echinoclathria (Microcionidae) (comb. nov.,
determined from re-examination of the type material); Axinella ? coccinea Carter, 1886a
(holotype BMNH 1886.12.15.8), from south-east Australia is an Zotrochota (Desmacididae)
(comb. nov.); Axinellaflabellata Carter, 1885 ('syntypes' BMNH 1886.12.15.471, 143), from
south-east Australia belongs to Sigrnaxia (Desmacellidae) (Hooper 1984); Axinella frondula
Whitelegge, 1907 (holotype AM G4349), from south-east Australia is a species of Raspailia
(Hooper 1991); Axinella hispida var. gracilis Lendenfeld, 1888 (holotype AM G9083), and
var. tenella Lendenfeld, 1888 (lectotype AM G9074), are species of Raspailia (Hooper 1991);
Axinella inflata Lendenfeld, 1888 (holotype AM G9081), from south-east Australia is
virtually unrecognisable (but possibly belonging to Dictyodendrillidae; F. Wiedenmayer,
Fig. 10. Axinella spp.: A ,
Chalinopora retepora Lendenfeld
(lectotype BMNH 1886.8.27.417);
B, skeleton (scale =500 pm);
C, Axinella polypoides Schmidt,
skeleton (specimen BMNH 1925.
1 1.1.1003 from Naples) (scale =
1 mm); D, Axinella villosa
Carter, skeleton ('syntype' BMNH
1886.12.15.396) (scale = 1 mm).
1410
J. N. A. Hooper and C. LCvi
personal communication);Axinella labyrinthica Dendy, 1889 (holotype BMNH 1889.1.21.3),
from the Indian Ocean and north-west Australia belongs to Teichaxinella (comb. nov.);
Axinella obtusa Lendenfeld, 1888 (holotype AM G9082, presently missing), from south-east
Australia is unrecognisable;Axinella pilifera Carter, 1885 (holotype unknown), from southeast Australia is virtually unrecognisable; Axinella setacea Carter, 1885 (holotype BMNH
1886.12.15.61) belongs to Raspailia and is a synonym o f R. pinnatifda (Carter) (Hooper
1991); Axinella solida Carter, 1885 (lectotype BMNH 1887.7.11.24), from south-east
Australia (Dendy 1897a) is a species o f Rhaphoxya (comb. nov.; Fig. 32C-D); Axinella
stelliderma Carter, 1885 (lectotype BMNH 1886.12.15.33), and var. acerata Carter, 1885
('holotype' BMNH 1886.12.15.63), from south-east Australia (Dendy 1897a) are species o f
Raspailia (Hooper 1991);Axinella symbiotica Whitelegge, 1907 (holotype AM G4350), from
south-east Australia is the type species o f Axinosia (Hallmann 1914b);Axinella vermiculata
Whitelegge, 1907 (holotype AM G4360), from south-east Australia belongs in Teichaxinella
(de Laubenfels 1936); Axinella virgultosa var. massa Carter, 1886b (holotype BMNH
1889.6.9.4; Fig. 20F-G), from the central Indian Ocean, south-east Asia and north-west
Australia, with synonym Suberites mollis Kieschnick (1898, 1900), belongs to Stylissa (see
below). Also, re-examination o f material described as 'Axinella echidnaea' by Ridley (1884)
and Hentschel (l912), from north-west Australia and southern Indonesia respectively, shows
that they refer to Reniochalina stalagmitis (see above).
Axinella carteri (Dendy)
(Figs 11-12, Table 2)
Acanthella carteri Dendy, 1889: 93-4, pl. 4, fig. 6; Dendy, 1905: 193, pl. 8, fig. 6; Dendy, 1922:
119, pl. 5, fig. 5; Vacelet et al., 1976: 43-4, pl. 2, fig. b; van Soest, 1989: 223-4, fig. 14.
Axinella carteri. -Burton, 1959: 258-9; LCvi, 1979: 311.
Acanthella aurantiaca Keller, 1889: 396.-Topsent, 1906: 562; Row, 1911: 356; Dendy, 1922: 119;
Vacelet and Vasseur, 1971: 80.
Material Examined
Holotype.
BMNH not seen; holotype of A. aurantiaca, ZMB 2921.
New Caledonian material. ORSTOM R1056: stn 241, Pointe Nind'Hio, Hienghene, 20'41 .OIS.,
164"57.2'E., 10 m depth, 6.ix.1978, coll. P. Laboute, SCUBA; ORSTOM R01: stn 153, barrier reef,
Ilat Taenia, Passe de St Vincent, 22'00. 11S.,165"56.llE., 7 m depth, 14.ix.1976, coll. P. Laboute,
SCUBA; ORSTOM 'cfRO1': stn 253, barrier reef S. of pass, YatC, 22°11~2'S.,167002~1'E.,25 m
depth, 26.ix.1979, coll. G. Bargibant, SCUBA; ORSTOM R1429: stn 332, W. entrance Canal Woodin,
Table 2.
Comparison in spicule dimensions between published and present
records of Axinella carteri
All measurements given in micrometres, and expressed as minimum-(mean)maximum range of measurement. N=25 for each specimen
MateriaVauthor
(Locality)
Holotype (Dendy 1889)
(Gulf of Manaar)
Keller 1889
(Red Sea)
Vacelet et al. 1976
(Madagascar)
Present study
(Comores) N = 1
(Papua New Guinea) N = 1
(Great Barrier Reef) N = 1
(New Caledonia) N = 4
Styles
Length
Width
New Caledonian Axinellids
1411
22°23.1'S.,166046.1'E., 18 m depth, 9.ii.1983, coll. G. Bargibant, SCUBA; NTM 23883 (ORSTOM
R170): stn 136, channel between I1Bt Canard and IlBt Maitre, 22°19~2'S.,166021.1'E., 12 m depth,
l 6 . i ~198
. 1, coll. J.-L.Menou, SCUBA; NTM 23890: Ile aux Canards, Noumea lagoon, 22Ol9.2'S.,
166"26'E., 22 m depth, 26.ix.1990, coll. J. N. A. Hooper, SCUBA; QM (3301323: CroisantLarkgritre, Ilat Maitre, off Noumea, 22°202'S.,166"22.5tE., 20 m depth, 13.x.1992, coll. J. N. A.
Hooper, SCUBA.
Comparative material. Great Barrier Reef, Qld: QM G300291 (fragment NTM 24027): E. side
of Magra I., Cockburn Is, Cape York Region, 11°52.0'S.,143017~O'E.,12.5 m depth, 10.xii.1990,
coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', SCUBA, (stn JH-90-051). Papua New Guinea:
NCI Q66C-4329-V (fragment QM G300355): Cement Mixer Reef, E. of Christiansen Research Institute,
Madang, 6°40.0'S.,145049~0'E., 6 m depth, 3.ix.1990, coll. NCI, SCUBA. Western Indian Ocean:
QM G301000: Comoros Is, 12"'S.,44"'E., 37 m depth, 1991, coll. G. R. Pettit et al., SCUBA (ref.
no. M5351).
Description
Colour. Bright orange-brown alive (Munsell2.5YR 7/8), pale orange-brown in ethanol.
Surface slightly darker than interior of sponge.
Shape. Flabellate growth form, 110-400 mm high, with massive, lobate, irregularly
planar or globular branching, up to 350 mm wide, and branches consist of relatively thick,
flattened planar or buttressed lamellae, 4-11 mm thick, with irregular margins; sponge
attached to substrate directly or by small basal stalk, 20-90 mm long, up to 40 mm
diameter.
Surface. Fleshy, conulose, rough surface; conules irregular, 3-5 mm high, solitary
or joined to form meandering surface ridges; texture rubbery, compressible, easily torn.
Small oscules, 2-5 mm diameter, occur predominantly on the margins of lamellae, usually
between surface conules.
Ectosome. Membraneous band of heavy collagen, 100-200 pn wide, slightly more
darkly pigmented than choanosomal region, with extra-axial spicules protruding only
sparsely.
A
Fig. 11. Axinella carteri Dendy, specimen QM G300291: A , structural
styles; B, section through peripheral skeleton.
1412
J . N. A. Hooper and C. LCvi
Choanosome. Axial skeleton condensed into several multispicular bundles, 100-250 pm
wide, running more-or-less longitudinally through lamellae, fully cored by long slender styles,
bound together by very light spongin fibre, and interconnected at irregular angles by vaguely
plumose, ascending, paucispicular extra-axial tracts of styles or individual spicules. Fibre
reticulation relatively close meshed, with lacunae forming elongate oval chambers, up to
300 x 70 pm. Collagen in mesohyl relatively light compared with ectosomal region.
Megascleres (refer to Table 2 for dimensions). Single category of style present, although
variable in thickness, occasionally strongylote; styles relatively long, slender or robust,
usually slightly curved symmetrically or towards basal end, sharply pointed, fusiform, with
evenly rounded base.
Microscleres. Absent.
Distribution
Red Sea (Keller 1889; Row 1911), Arabian Gulf (Burton 1959), western Indian Ocean:
Cargados Carajos, Diego Garcia, Amirante, Salomon, Seychelles Is, Comores, Madagascar
(Dendy 1922; Vacelet et al. 1976; present study), Gulf of Manaar, Sri Lanka (Dendy 1889,
1905), Lesser Sumba region, Indonesia (van Soest 1989), Papua New Guinea and the Great
Barrier Reef (present study), and New Caledonia (LCvi 1979; present study). This species is
a common component of Indo-Pacific coral reefs, usually found in both the lagoon and
Fig. 12. Axinella carteri Dendy: A , specimen ORSTOM R170 in situ
(photo P . Laboute); B, specimen QM G300291;C, SEM skeleton (scale=
500 pm).
New Caledonian Axinellids
1413
outer reef slope; usually associated with living and dead coral. The species appears to be
most common in areas of strong current, attached to coral rubble or rock, in sand and sea
grass beds. Known depth range is 6-37 m.
Remarks
None of the material described above had a second category of longer style present, as
described in Dendy's (1889) original description (cited as 1200x 11 pm, compared with
typical spicules of 400 x 21 pm), although Burton (1959) suggests that this longer spicule is
present in few of the many known specimens of A. carteri. Apart from this feature, the
species appears to be fairly homogeneous in skeletal structure, spicule morphology and
size across its wide geographic range, although spicule dimensions vary slightly between
material (Table 2). The species is readily recognisable in the field by its bright orange-brown
coloration, fleshy surface and thick irregular lamellae.
Axinella carteri is difficult to place with absolute confidence in any single genus, given
the relatively unclear boundaries between several structurally similar axinellid genera (e.g.
Phakellia, Acanthella, Axinella, Teichaxinella). Vosmaer's (1912) criteria for differentiation
of these taxa are not entirely clear cut, because many more species have been described since
that early revision, showing that many intermediate (transformation) states are possible
between the skeleton plans he initially proposed. Nevertheless, under his scheme for differentiating such genera, the present species falls closest to Axinella in having some axial
compression and sparsely diverging, plumose extra-axial spicules.
Genus Phakellia Bowerbank
Phakellia Bowerbank, 1864: 186.-Bowerbank, 1866: 122; Schmidt, 1866: 15; Gray, 1867: 516;
Ridley and Dendy, 1887: 169; Dendy, 1905: 190; Dendy, 1922: 116; Vosmaer, 1912: 310; Topsent,
1928: 37; de Laubenfels, 1936: 130; Bergquist, 1970: 17 [type species Spongia ventilabra Linnaeus,
1767: 1296, which is a junior synonym of Phakellia strigosa (Pallas, 1766) (Vosmaer 1912: 310)l.
Querciclona de Laubenfels, 1936: 46 [comb. nov.] (type species Antherochalina quercifolia Keller,
1889: 383; holotype ZMB 429).
Diagnosis
Compressed flabellate or cup-like growth forms predominant; surface smooth or
microconulose; oscules frequently surrounded by subdermal drainage canals; ectosome
membraneous without specialised skeleton, often lightly hispid from protruding choanosomal
megascleres; choanosomal axial skeleton dense, typically with only interwoven styles, or may
include sinuous strongyles and styles, or only strongyles, organised into multispicular
ascending and paucispicular transverse tracts, together forming compressed axial reticulation;
extra-axial skeleton comparatively sparse, with plumose bundles or single styles or oxeas
perpendicular to axis, with or without transverse connecting megascleres; megascleres styles,
sinuous strongyles or oxeas; microscleres absent (modified from Vosmaer 1912).
Remarks
Phakellia is similar to, and often confused with, Acanthella, differing primarily in
having a well-differentiated axial and extra-axial skeleton (Vosmaer 1912; Bergquist 1970).
Querciclona de Laubenfels (1936: 46) is an obvious synonym of Phakellia, although
de Laubenfels initially placed the taxon in Microcionidae. Pararhaphoxya Burton (1934)
may also eventually be merged with Phakellia, but is presently maintained separately
and distinguished only by the possession of asymmetrical megascleres. However, from
examination of nearly 40 species of both Acantheua and Phakellia from the Indo-Pacific
faunas, consisting of both described and undescribed taxa, it is tentatively suggested that
the structure of axial and extra-axial skeletons is the only feature that consistently differentiates these genera, lending support to Vosmaer's (1912) original proposal. By comparison,
the presence or absence of sinuous axial megascleres seems to be much less stable and
appears to occur indiscriminantly in species of both genera, and these features must be used
with more caution.
1414
J. N. A. Hooper and C. Ldvi
In total, 21 species from the Australasian region (including New Zealand, New Caledonia
and southern Indonesia) have been described in, or subsequently referred to, Phakellia,
but examination of relevant type material suggests that only seven of these are most
appropriately placed here. Approximately 20 other, undescribed Phakellia species have also
been collected from tropical Australia, but these will be described elsewhere at a later date.
Valid Australasian species of Phakellia include: Phakellia carduus (Lamarck, 1814;
holotype MNHN LBIM DT533) from north-west and north-east Australia (Ridley 1884;
Fig. 13. Phakellia spp.: A , Acanthella cavernosa Dendy (holotype BMNH
1921.11.7.100);B, specimen on deck (NTM 22736); C, skeleton (holotype) (scale =
500 pm); D, peripheral skeleton (scale=200 pm); E, Spongia carduus Lamarck
(specimen on deck NTM 21837); F, skeleton (paralectotype MNHN DT3447)
(scale=500 pm); G, paralectotype MNHN DT533); H, specimen on deck (QM
G300431).
New Caledonian Axinellids
1415
Topsent 1930; Bergquist 1970; Hooper, unpublished data) (Fig. 13E-H); Phakellia cavernosa
(Dendy, 1922; holotype BMNH 1921.11.7.100) [with synonyms Acanthella stipitata, in part
(variety o f Ridley and Dendy, 1887; not Acanthella stipitata Carter, 1881; 'holotype'
BMNH 1887.5.2.73), and Burton, 1934 (specimen BMNH l930.8.13.142), and Acanthella
klethra Pulitzer-Finali, 1982 (holotype MHNG 46927), from the Indian Ocean, north-west
and north-east Australia, and the Great Barrier Reef (Burton 1934; Bergquist 1970; Hooper,
unpublished data) (Fig. 13A-D)]; Phakellia columnata (Burton, 1928; holotype BMNH
[not found]; 'representative specimen' MNHN DCL2934), known from the Andaman Sea
and deeper waters o f f New Caledonia (LCvi and LCvi 1983) (Fig. 14A-B); Phakellia conulosa
Dendy, 1922 (holotype BMNH [not found]; 'representative specimen' from the Philippines
QM G300315), from the Indian Ocean and north-west Australia (Dendy 1922; Hooper,
unpublished data) (Fig. 14C-D); Phakellia dendyi Bergquist (1970;holotype NMNZ Porif .24)
from north-west, north-east and south-east Australia, and New Zealand (Dendy 1897b;
Bergquist 1970; Hooper, unpublished data) (Fig. 14E-F); Phakelliapulcherrima (Ridley and
Dendy, 1886; holotype BMNH 1887.5.2.23) (Figs 14G-H, 15-16) (see below); and Phakellia
stipitata (Carter, 1881; holotype BMNH [confirmeddestroyed];neotype QM G300874, here
designated) from eastern and south-east Australia (see below) (Fig. 17-18).
Other Australasian species described in Phakellia but better placed elsewhere include:
Phakellia aruensis Hentschel, 1912 (holotype SMF 953), from southern Indonesia is assigned
to Axinella; Phakellia brassicata Carter, 1885 (holotype BMNH 1886.12.15.75), from southeast and north-east Australia belongs to Cribrochalina (Niphatidae, order Haplosclerida);
Phakellia crassa Carter, 1885 (holotype BMNH 1886.12.15.129), from south-east Australia
(Fig. 9C-D) is an Axinella; Phakellia flabellata Carter, 1885 (holotype BMNH 1886.
12.15.471), and Phakellia villosa Carter, 1886a (lectotype BMNH 1886.12.15.78), both from
south-east Australia, are both synonyms o f Teichaxinellaflabellata (Carter, 1885);Phakellia
inj7exa Pulitzer-Finali, 1982 (holotype MHNG 46931 [not seen by the authors]), from the
southern Great Barrier Reef is an Acanthella; Phakellia jacksoniana Dendy, 1897a (lectotype
BMNH 1887.5.2.9), is a synonym o f Clathria (Isociella)jacksoniana (Dendy)(Microcionidae,
order Poecilosclerida);Phakellia multiformis Whitelegge, 1907 ('syntype' AM G4358), from
south-east Australia belongs to Axinosia; Phakellia papyracea Carter, 1886a (lectotype
BMNH 1886.12.15.231), from south-east Australia is a synonym o f Echinoclathria leporina
(Lamarck, 1814) (Microcionidae, order Poecilosclerida); Phakellia plumosa LCvi & LCvi,
1983 (holotype MNHN DCL2972), from deeper waters o f f New Caledonia belongs in
Reniochalina (Fig. 6C-D); Phakellia ramosa Carter, 1883 (holotype BMNH 1884.4.14.2),
from south-east Australia belongs to Sigmaxinella (Desmacellidae, order Poecilosclerida);
Phakellia tumida Dendy, 1897a (holotype NMV G2464), from south-east Australia belongs
to Pseudaxinella; and Phakellia ventilabrum australiensis Carter, 1886a (holotype BMNH
1886.12.15.422), from south-east Australia is a synonym o f Clathria (Thalysias) cactiformis
(Lamarck, 18 14) (Microcionidae, order Poecilosclerida).
Phakellia pulcherrima (Ridley & Dendy), comb. nov.
(Figs 14G-H, 15-16, Table 3)
Acanthella sp. -Ridley, 1884: 463.
Acanthella pulcherrima Ridley & Dendy, 1886: 218, 479.-Ridley and Dendy, 1887: 177, pl. 32,
fig. 3.
Not Acanthella pulcherrima. -Capon and MacLeod, 1988: 979-83.
Acanthella pulcherrima calyx Dendy, 1922: 120, pl. 5, fig. 6.
Material Examined
Holotype. BMNH 1921.11.7.100 (fragment MNHN DCL209L): Prince of Wales Channel, Torres
Strait, 14 m depth, HMS 'Challenger'.
New Caledonian material. Q M G300019 (ORSTOM R1321): stn 102, mid-channel, Canal Woodin,
22"23.1'S.,166°48.1'E.,28.iv.1976, 33 m depth, coll. P. Laboute, SCUBA; ORSTOM R1052: stn 136,
channel between Il6t Canard and Ilat Maitre, 22°19~2'S.,166021.1'E.,20 m depth, 16.iv.1981, coll.
P. Laboute, SCUBA.
J . N. A. Hooper and C. Ltvi
1416
Description
Colour. Pale orange-brown or brown alive (Munsell 5YR 7/10), pale beige in ethanol.
Shape. Small club-shaped sponge (75-120 mm long, 58-75 mm maximum breadth),
with short cylindrical stalk (12-22 mm long, 7-10 mm wide), enlarged basal holdfast, and
several very thin, leaf-like, flattened branches (10-30 mm maximum width, 4-8 mm thick),
with even margins.
Fig. 14. Phakellia spp.: A , Bubaris columnata Burton (specimen from New Caledonia
MNHN DCL2934); B, skeleton (scale= 500 pm); C, Phakellia conulosa Dendy, in situ
(specimen from the Philippines, QM G300315) (photo J . N. A. Hooper); D, skeleton
(scale=200 pm); E, Phakellia dendyi Bergquist (specimen on deck NTM 21318);
F, skeleton (scale=500 pm); G, Acanthella pulcherrima Ridley and Dendy (holotype
BMNH 1887.5.2.23); H, skeleton (scale = 500 pm).
New Caledonian Axinellids
1417
Table 3. Comparison in spicule dimensions between published and present records of
Phakellia pulcherrima
All measurements given in micrometres, and expressed as minimum-(mean)-maximum
range of measurement. N=25 for each specimen
MateriaYauthor
(Locality)
Holotype (BMNH 1887.5.2.23)
Dendy (1922) (Cargados Carajos)
Present material (New Caledonia)
L
W
L
W
L
W
Extra-axial
styles
Axial
strongyles
294-(331.2)-392
~ 6 - ( 1 1.O)-15
'Larger'
360-(577 1)-940
x 2-(5.2)-9
Up to 1300
x 8.6
449-(488.4-552
x 2-(4.2)-6
253-(360.3)-413
x 2-(5.4)-9
Fig. 15. Phakellia pulcherrima (Ridley and Dendy), specimen QM G300019: A , extra-axial
styles; B, axial strongyles; C , anisoxeas frequent in holotype BMNH 1887.5.2.23; D, section
through peripheral skeleton of specimen QM G300019.
1418
J. N. A. Hooper and C. L&i
Surface. Evenly conulose; conules rounded or pointed, more-or-less arranged in ridges,
up to 5 mm high, running longitudinally along branches, with ridges producing an almost
striated pattern. Oscules and pores not seen in preserved material. Texture firm, flexible.
Ectosome. Membraneous, fleshy, without speciaiised spicules; points of extra-axial styles
protrude 100-250 pm from surface conules, whereas between/conules ectosome is merely
fleshy, with more darkly pigmented granular collagen than in choanosomal region.
Choanosome. Skeleton divided into distinct axial and extra-axial regions. Axis tightly
compressed, occupying only about 30% of branch diameter, running longitudinally through
branches, cored by closely reticulate sinuous strongyles, more-or-less interlocked and crisscrossed within axis. Extra-axial skeleton consists of radial tracts of individual, or loosely
plumose bundles of styles, embedded in and standing perpendicular to the axis. Axial
spicules bound by moderately heavy, close-set spongin fibres, up to 30 pm diameter, with
fibre reticulation producing elongate meshes, 20-70 pm diameter. Extra-axial spicules free
within mesohyl, not associated with spongin fibres except where embedded into axis.
No choanocyte chambers observed in peripheral skeleton.
Megascleres (refer to Table 3 for dimensions). Extra-axial styles relatively thick, straight
or slightly curved near basal end, with tapering fusiform points and with evenly rounded or
slightly constricted bases. Axial strongyles usually sinuous, occasionally completely straight
or vermiform, relatively thick, with evenly rounded bases.
Microscleres. Absent.
Distribution
New Caledonian lagoon, Cape York, Torres Strait, Great Barrier Reef, Cargados Carajos,
Indian Ocean; 5-54 m depth; coral rubble.
Fig. 16. Phakellia pulcherrima (Ridley and Dendy): A , specimen in situ (ORSTOM R1052) (photo
P. Laboute); B, specimen QM G300019; C, SEM skeleton (magnified 80 x); D, SEM transverse section
(magnified 94 x ) .
New Caledonian Axinellids
1419
Remarks
The New Caledonian population of P. pulcherrima is a new locality record for the
species, and is the first validated record (i.e. confirmed by voucher specimen) since Dendy
(1922) rediscovered the species from the western Indian Ocean. A major difference between
the New Caledonian specimen and the two previously known records of the species is that
many of the extra-axial styles have modified oxeote bases in the older specimens, whereas
in the New Caledonian specimen these spicules are invariably styloid. Spicule dimensions also
vary slightly, particularly in the maximum lengths of strongyles (Table 3), but considering
that these spicules break up easily during histological preparation it is possible that they
may actually be larger in New Caledonian material than described here. In other details
(growth form, surface sculpturing, skeletal structure), all known specimens are similar and
undoubtedly conspecific.
This species is contrasted with other Australasian Phakellia in Figs 13-14.
Phakellia stipitata (Carter)
(Figs 17-18)
Acanthella stipitata Carter, 1881: 380, pl. 18, fig. 8.
Not Acanthella stipitata.-Ridley and Dendy, 1887: 178; Burton, 1934: 565.
Material Examined
Neotype. QM G300874: Peel I., Lazarette Gutter, Moreton Bay, Qld, 27"28.9'S.,l53"21.4'E.,
8 m depth, 14.iv.1992, coll. J . N. A. Hooper and S. D. Cook, SCUBA.
New Caledonian material. QM G 300280 (fragment NTM 23888): Ile aux Canards, Noumea
lagoon, 22°19.0'S.,166026.0'E.,22 m depth, 26.ix.1990, coll. J. N. A. Hooper and J . Vacelet, SCUBA,
sand, rubble, sea grass beds; ORSTOM R1429 (fragment NTM 23875): stn 332, W. entrance Canal
Woodin, 22"23.1'S.,166"46.1'E., 18 m depth, 9.ii.1983, coll. J.-L. Menou, SCUBA.
Comparative material. QM G301235: same locality as neotype.
Description
Colour. Bright orange-brown alive (Munsell 5YR 7/10), orange-yellow in ethanol.
Shape. Flabellate sponge, 90-130 mm long, 70-110 mm wide, 40-90 mm thick, with
one or more fans aligned face-to-face, composed of irregularly fused and reticulated
branches, with excavated wide meshes between reticulations, producing thick, nearly bulbous
branching; small basal stalk, up to 30 mm long, 15-20 mm diameter (often detached from
specimen), with a broad basal holdfast.
Surface. Clathrous, excavated surface, with well-developed, regularly spaced conules,
10-20 mm apart, 5-20 mm high; conules have rounded tips, usually joined together by
low ridges, surrounding large excavations through the sponge (4-15 mm diameter) (i.e.
producing the surface meshes); remainder of membraneous surface with 'goose-flesh'
appearance, and covered with small ostia (< 1 mm diameter). Large oscules on margins of
fans, 15-25 mm diameter, slightly raised above surface, and slightly more darkly pigmented
than rest of surface. Texture firm, rubbery, difficult to tear, usually requires cutting off
substrate.
Ectosome. Membraneous, heavily collagenous, darkly pigmented, without special
spicules and only points of extra-axial styles barely protruding through surface (these usually
only on ends of surface conules).
Choanosome. Skeleton clearly differentiated into axial and extra-axial regions. Axis
moderately compressed, with short, heavy, reticulated spongin fibres, 20-45 pm diameter,
producing oval or elongate meshes, 60-110 pm diameter; fibres only partially cored by
styles in uni- or multispicular tracts. Extra-axial skeleton not well formed (as in some other
species of Phakellia), consisting of radially arranged styles, as individuals or multispicular
brushes, standing perpendicular or at acute angles to axis, ascending to and usually
1420
J . N. A. Hooper and C. Lkvi
protruding through ectosome. Styles in axial and extra-axial skeletons do not appear to be
differentiated. Choanocyte chambers small, elongate, up to 40 pm diameter.
Megascleres. Axial and extra-axial megascleres are exclusively styles, long, slender,
straight or slightly curved, with abrupt points, sharp or slightly stepped tips, and with
evenly rounded or occasionally oxeote bases. Length (neotype): 172-(345.5)-494 pm; (New
Caledonian material): 301-(412.7)-545 pm. Width (neotype): 3-(7-3-12 pm; (New
Caledonian material): 3-(8.1)-15 pm.
Microscleres. Absent.
Distribution
Gulf of Manaar (original record), south-east Queensland, and New Caledonia; associated
with living and dead coral, coral reef lagoon.
Fig. 17. Phakellia stipitata (Carter):A, neotype QM G300874, structural styles and anisoxeas;
B, specimen QM G300280 from New Caledonia, styles and anisoxeas; C, section through
peripheral skeleton of neotype.
New Caledonian Axinellids
1421
Remarks
There is no extant type material of this species in the BMNH or LFM (S. Stone, BMNH,
personal communication, and personal observation), nor has the species been recorded since
it was first described. Records cited by Ridley and Dendy (1887) and Burton (1934) refer
to Phakellia cavernosa (Dendy, 1922), and this is confirmed by the presence of sinuous
strongyles in their material (BMNH 1887.5.2.73 and BMNH 1930.8.13.142, respectively).
However, some distinctive features of the species are obvious from Carter's (1881) original
description, notably the characteristic clathrous, excavated gross morphology, and spicule
morphology (particularly spicule size, geometry and presence of abrupt points). These
agree closely with the present material from Moreton Bay and the Noumea lagoon. New
Caledonian specimens have marginally larger styles than does the neotype from Moreton
Bay, but this difference is relatively trivial, and the species are similar in all other features.
Fig. 18. Phakellia stipitata (Carter): A , specimen in situ (ORSTOM R1429) (photo
J.-L. Menou); B, preserved specimen (QM (3300280); C, three specimens on deck (neotype
QM G300874 top left, indicated by arrow; other specimens QM G301235); D, peripheral
skeleton (neotype QM G300874) (scale=200 pm); E, SEM transverse section (specimen
QM G300280) (scale = 200 pm).
1422
J. N. A. Hooper and C . L6vi
Genus Stylissa Hallmann
Stylissa Hallmann, 1914b: 349 [type species Stylotellaflabelliformis Hentschel, 1912: 355, holotype
SMF 15871.
Dragmaxia Hallmann, 1916a: 543.-Hallmann, 1916b: 674; de Laubenfels 1936: 130 [type species
Spongosorites variabilis Whitelegge, 1907: 513, holotype AM (presently missing)].
Diagnosis
Fan, cup-shaped or massive foliose growth forms; surface shaggy, often with small
papillae or grooved ridges; ectosome fleshy, without specialised spiculation, but with brushes
of peripheral choanosomal styles protruding through surface; choanosomal skeleton disorganised plumo-reticulate, with a slightly condensed axis and slight differentiation between
axial and extra-axial skeletons; fibres cored by parallel tracts of styles, of 1 or 2 sizes,
more-or-less ascending, and diverging towards the periphery; peripheral styles often slightly
larger than those in axis. Microscleres absent or present (raphides, trichodragmata) (modified
from Hallmann 1914b).
Remarks
Whitelegge (1907) characterised Spongosorites variabilis partly by the presence of raphides
in the dermal layer, and Hallmann (1916~)erected Dragmaxia to emphasise this feature,
which he generally considered to be diagnostically important throughout his taxonomy.
Hallmann also noted that these raphides occurred throughout the mesohyl, not necessarily
confined to the ectosomal region, as supposed by Whitelegge (1907). In addition, Hallmann
(1916~)noted that although both Dragmaxia and Stylissa have comparable architecture and
only styles for megascleres, those of the former were differentiated into two size categories,
with some evidence of spicule localisation (the larger apparently found closer to the
periphery). However, in other Stylissa spicule size may extend over a large size range,
and often the peripheral spicules are marginally larger than those in the axial skeleton.
Whitelegge's holotype of S. variabilis is missing from AM collections, and these features
cannot be accurately verified, but the species is perfectly recognisable from Hallmann's
(1916~:544) redescription, which includes comprehensive illustrations. It is concluded from
this available evidence that Dragmaxia differs in no substantial respect from Stylissa and the
two are here merged, as was suggested by de Laubenfels (1936).
Stylissa presently contains only three valid species, all from the Indo-west Pacific region:
the type species S. flabelliformis (Hentschel, 1912), previously known only from the Arafura
Sea but now known from the Indo-west Pacific region in general (Fig. 19-20); S. variabilis
(Whitelegge, 1907), known only from the Crookhaven River, N.S. W. (Hallmann 1916a)
(holotype AM, missing); and another widely distributed species, S. massa (Carter, 1886b)
[for Axinella virgultosa var. massa Carter (holotype BMNH 1889.6.9.4; Fig. 20F-G)
(Burton 1959), with synonyms Suberites mollis Kieschnick, 1898, and Stylotella conulosa
Topsent, 1897 (lectotype MHNG C-12/45, paralectotype MNHN LBIM DT1775, here
designated)], extending from the south Arabian coast (Burton 1959), the Mergui Archipelago,
Andaman Sea (Carter 1886b), Torres Strait, Qld (Kieschnick, 1898, 1900), Ambon, Moluccas
(Topsent 1897; Desqueyroux-Faundez 1981), Ternate, Moluccas (Kieschnick 1898, 1900),
Java Sea (Lindgren 1897, 1898), and Christmas Island, western Indian Ocean (Kirkpatrick
1900).
Stylissa flabelliformis (Hentschel)
(Figs 19-20, Table 4)
Stylotella flabelllformis Hentschel, 1912: 355-6, pl. 19, fig. 26.
Stylissa flabelliformis. -Hallmann, 1914b: 349.
Teichaxinella Iabyrinthica.-Hooper et al., 1992: 265.
Material Examined
Holotype. SMF 1587 (schizotype MNHN DCL2314): Meriri, Aru I., Arafura Sea, 6"S.,134"501E.,
6-10 m depth.
New Caledonian Axinellids
1423
New Caledonian material. QM G300017 (ORSTOM R1257): stn 166, N. pass Toemo, Port de
Goro, 22°20.0'S.,167001.OIE., 18 m depth, 28.x.1976, coll. A. IntCs, SCUBA; QM (3300689 (fragment
NTM 23874): Baie des Citrons, Noumea lagoon, 22°18'S.,166025'E., 3 m depth, 25.ix.1990, coll.
J. N. A. Hooper, snorkel; ORSTOM R565: stn 113, L'epave du 'Bonhomme', Grand RCcif Mbttre,
22°21~O'S.,166014~0'E.,25 m depth, 21.vi.1976, coll. P. Laboute, SCUBA.
Comparative material. Seychelles Is: QM G300068: E. of Curieuse I., 4"15'S.,55"47'E., depth and
date of collection unknown, coll. Pettit, G. R. et al., SCUBA (ref. no. M5281). Japan: QM G301236:
Yonaguni I., E. of Taiwan, 24"29'N.,123"00'E., 30 m depth, 1992, coll. T. Higa (ref. K-1). Sahul
Shelf, W.A.: QM G300183 (fragment NTM 22817): West I. passage, outer reef, Ashmore Reef,
12"15'S.,122"55'E., 16.5 m depth, 28.vii.1986, coll. C. Johnston, SCUBA; NTM 22797, 2798: West I.
passage, outer reef edge, Ashmore Reef, 12°14.3'S.,123056.0'E., 15.5 m depth, 27.vii.1986, coll.
A. M. Mussig, SCUBA; QM G301081, G301082: Cartier I., outer reef slope, N. side reef, 12"31.4'S.,
123"33.3'E., 14 m depth, 06.v.1992, coll. J. N. A. Hooper, SCUBA. Northwest Shelf, W.A.: NTM
23374: 2 nm from shore, N. of Barrow I., 20°38.8'S.,11548.8'E., 22 m depth, 26.viii.1988, coll.
D. Low Choy, SCUBA; NTM 2664: NW. of Yampi Sound, 15"27.04'S.,121°49.00'E.,76 m depth,
29.iv.1982, coll. CSIRO R.V. 'Sprightly', dredge; QM G300108 (fragment NTM 21220), NTM 21236:
40 m depth, 26.iv.1983, coll.
N. of Bedout I., W. of Port Hedland, 19°28~05'S.,118055~30'E.,
J. N. A. Hooper, R.V. 'Soela', S02/83, stn B9, trawl; NTM 21471: W. of Port Hedland, 1g056'S.,
117"57'E., 40 m depth, O5.xii.1985, coll. Ward, T., trawl; NTM 21807, 21829: W. of Port Hedland,
19°26~09'S.,118054~02'E.,
50 m depth, 30.viii.1983, coll. Ward, T. R.V. 'Soela', trawl; NTM 2684:
N. of Port Hedland, 19"16'S.,118"50'E., 70 m depth, 4.v.1982, coll. CSIRO R.V. 'Sprightly', dredge;
NTM 22321: NW. of Lacepede Is, 16°31'S.,121028'E., 38-40 m depth, 17.iv.1985, coll. Russell, B. C.,
pair trawl; NTM 2731: N. of Adele I., Collier Bay, 15°58.03'S.,122039.07'E., 59 m depth, 21.iv.1982,
coll. CSIRO R.V. 'Sprightly', dredge; NTM 22351: NW. of Lacepede Is, 16"34'S.,121°27'E., 40-46 m
depth, 17.iv.1985, coll. B. C. Russell, pair trawl; NTM 23021: N. of Amphinome Shoals, 19"19.7'S.,
119O08.8'E., 50 m depth, 19.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', trawl.
Arafura Sea, N.T.: NTM 2607: Cootamundra Shoals, N. of Melville I., 10"49~07'S.,129"12~09'E.,
31 m depth, 6.v.1982, coll. B. Thom, R. Lockyer, SCUBA; NTM 2616: same locality, 10°50.22'S.,
129"13.17'E., 22 m depth, 10.v.1982, coll. R. Lockyer, SCUBA; NTM 23081: Parry Shoals,
11°11~41'S.,129043~01'E.,
18 m depth, 13.viii.1987, coll. A. M. Mussig, SCUBA.
Description
Colour. Live coloration dark orange-brown (Munsell 5YR 7-6/10), with a paler
membraneous ectosome, and brighter orange interior (5YR 7/12); red-brown on deck;
produces an orange exudate upon collection.
Shape. Growth form generally thickly flabellate, 120-450 mm long, 70-180 mm wide,
with flabellate-digitate branches growing in more than 1 plane, with 140 mm maximum span,
with even or uneven digitate margins on branches, up to 30 mm thick, attached to substrate
by small thickly cylindrical basal stalk, 20-75 mm long, 11-45 mm diameter.
Surface. Characteristically rough, striated, conulose, shaggy surface, with either longitudinal striations in larger specimens, or irregularly conulose, sculptured surface in younger
material. Surface soft, fleshy in life, contracting in preserved material to produce harsh
texture. Oscules visible on apex of surface ridges and margins of branches, up to 14 mm
diameter, with a large membraneous lip surrounding each exhalant pore. Fleshy parts of
surface porous, with evenly dispersed ostia, predominantly between conules and ridges.
Ectosome. Fleshy, darkly pigmented, heavily collagenous ectosome, with points of styles
from peripheral skeleton protruding up to 200 pm from the surface. In vicinity of surface
conules peripheral (subectosomal) skeleton extends directly into subdermal region, whereas
in fleshy area between conules there are only few spicules present.
Choanosome. Skeletal structure plumo-reticulate, although it appears disorganised due
to proportionally large size of megascleres in relation to spongin fibres, with only slightly
differentiated axial and extra-axial regions. Axial skeleton reticulate, with heavy spongin
fibres (80-170 pm diameter), forming rectangular meshes (80-170 pm diameter), cored by
multispicular tracts of styles. Extra-axial skeleton vaguely plumo-reticulate, with multispicular ascending tracts of styles interconnected at irregular intervals by uni- or pauci-
1424
J. N. A. Hooper and C. LCvi
Table 4. Comparison in spicule dimensions between published and present records of Stylissa
flabellifonnis and other members of the genus
All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of
measurement. N=25 for each specimen
Population
Styles
Length
Stylissa flabelliformis (Hentschel)
Indonesia (holotype)
New Caledonia (N= 2)
Seychelles (N= 1)
Japan (N= 1)
North-west Australia (N= 13)
Atypical specimen, Northwest Shelf, W.A. (QM G300108)
Stylissa massa (Carter)
Mergui Archipelago (holotype)
Stylissa variabilis (Hallmann)
Southern New South Wales (Hallmann 1916a)
Width
348-(447.1)-522
339-(433.5)-516
343-(423 '6)-482
396-(486.5)-582
341-(444.6)-554
272-(333.9)-380
9-(16.9)-23
6-(15.8)-22
13-(19.7)-27
8-(20.5)-28
7-(17.4)-28
7-(20.1)-32
504-(564.4)-648
11-(17.8)-22
350-900
1000-1 500
33-45
18
Fig. 19. Stylissaflabelliformis (Hentschel): A, specimen QM G300017, styles and strongylote spicules;
B, schizotype MNHN DCL2314, structural spicules; C, section through peripheral skeleton of New
Caledonian specimen QM G300017.
New Caledonian Axinellids
1425
spicular transverse tracts. Fibre reticulation in extra-axial region slightly more cavernous
than axis, with meshes up to 300 pm diameter. Collagen abundant; choanocyte chambers
small, elongate, 70-90 pm diameter.
Megascleres (refer to Table 4 for dimensions). Styles of a single size class, although
great variability in thickness; styles predominantly robust, slightly curved near basal end,
rarely straight, usually with evenly rounded bases, rarely rhabdose, tapering to fusiform
points, and occasionally modified to strongyles.
Microscleres. Absent.
Distribution
Indian Ocean and Indo-west Pacific, known from the Seychelles, south-eastern Indonesia,
Arafura Sea, Timor Sea, mid-Western Australia, Japan and New Caledonia. Recorded from
Fig. 20. Stylissa flabelliforinis (Hentschel): A , fragment of (?) paratype SMF1690
(schizotype MNHN DCL2314); B, specimen from New Caledonia [ORSTOM R1257
(QM G300017)l; C, specimen on deck from NW. Australia (NTM 21807); D, specimen
from Ashmore Reef (QM G300183); E, SEM skeleton (scale = 500 pm); F, Stylissa massa
(Carter), skeleton of holotype BMNH 1889.6.9.4 (scale = 500 pm); G, holotype.
J. N. A. Hooper and C. L6vi
1426
coral reefs, in lagoon, fringing and patch reefs and outer reef slope, sand and coral rubble,
3-76 m depth.
Remarks
The New Caledonian population of S. flabelliformis is currently the most easterly known
record of the species. The species is probably a prominent member of the Indo-west Pacific
fauna, although its known distribution is patchy. Certainly, in the eastern Indian Ocean,
from eastern Indonesia to the vicinity of Northwest Cape, Western Australia, the species is
relatively abundant in the subtidal to about 50 m depth, whereas its relative abundance
outside this zone is unknown. In New Caledonia the species is also moderately common
within the lagoon, less abundant on the outer reefs, and from the restricted material
examined there were no obvious differences detected between any of the regional populations
(see also Table 4). However, one specimen trawled from the Northwest Shelf of Western
Australia (QM G300108) was atypical in growth form (being much larger, with even margins
on the lobate branches, and having distinct longitudinal surface ridges), and in spicule
dimensions (having generally shorter and thicker spicules; Table 4), but these differences are
not considered to be significant and in other features it agreed closely with all other known
material of this species.
Stylissa flabelliformis is closely related to its sympatric sibling species, S. massa, which,
from re-examination of the holotype, appears to differ only in growth form (the latter being
massive; Fig. 20G), skeletal structure (S. massa has an even more disorganised reticulate
skeleton than S. flabelliformis; Fig. 20F), and larger spicule dimensions (Table 4). These
features appear to be consistent in all the descriptions of S. massa, and the two species are
retained here, but morphologically they are certainly closely related. From Hallmann's
(1916~:544) comprehensive redescription of S. variabilis, it appears to be similar in growth
form to S. flabelliformis, whereas the skeleton is much more organised, plumose (plumoreticulate, resembling Teichaxinella more than Stylissa), styles are larger and more-or-less
divisible into two size classes, and raphides are also present.
Genus Ptilocaulis Carter
Ptilocaulis Carter, 1883: 321.-Carter, 1884: 130; Topsent, 1928: 37, 172; de Laubenfels, 1936: 127;
Wiedenmayer, 1977: 152 [type species Ptilocaulis gracilis Carter, 1883: 321 (holotype BMNH
1845.12.30.1) (de Laubenfels, 1936) (Fig. 21D)l.
Plicatella Schmidt, 1864: 39 [preocc.]. -de Laubenfels, 1936: 132 [type species Reniera labyrinthica
Schmidt, 1864: 391.
Diagnosis
Typically erect, cylindrical, digitate, clavate to bushy, occasionally lamellate growth
form; surface conulose or with numerous elongate and overlapping papilliform projections,
often dividing at the apex. Choanosomal skeleton plumo-reticulate, with clearly differentiated
axial and extra-axial components; axial portion of skeleton condensed, composed of
irregularly anastomosing, close-set spongin fibres and spicules; extra-axial skeleton plumoreticulate or plumose, with heavy fibres cored by ascending multispicular tracts of megascleres, interconnected by paucispicular transverse tracts forming subisodictyal reticulation
(s.s.), or without transverse spicule skeleton and simply with meandering, plumose extraaxial spicule tracts; ectosome fleshy, without specialised spiculation, although plumose
brushes of spicules may protrude through surface. Megascleres styles, subtylostyles, anisoxeas
or strongyles (usually asymmetrical), sometimes including sinuous forms. Microscleres absent
(modified from Wiedenmayer 1977).
Remarks
De Laubenfels (1936: 127) suggested that Ptilocaulis was characterised by having superficial surface processes, a condensed and plumo-reticulate skeleton rich in spongin, and only
styles for megascleres, but these features barely define the taxon and in any case are not
completely accurate (i.e. spicules usually include styles, anisoxeas, strongyles, and oxeas,
New Caledonian Axinellids
1427
some or all with telescoped ends). Wiedenmayer (1977: 152) suggested that longer spicules
in the extra-axial skeleton, larger than those found in the choanosome, could also be used
as a discriminatory character for the genus, but this feature is not present in the type species
nor in many other described species. Wiedenmayer's observations were based on two species
that he placed in Ptilocaulis, one of which was Spongia spiculifera Lamarck (1814) from
southern Australia, following Topsent (1932), which had clearly differentiated size classes
of ectosomal and choanosomal megascleres but was similar in other characters to other
members of the genus, but this feature cannot be construed as being typical of Ptilocaulis.
Fig. 21. Ptilocaulis spp.: A , Spongia echidnaea Lamarck (holotype MNHN DT640); B, specimen
(MNHN DT3347); C, Topsent's spicule preparation of holotype (scale = 200 ~ m )D,
; Ptilocaulis gracilis
Carter, skeleton of holotype (BMNH 1845.12.30.1) (scale=500 pm); E, Spongia spiculifera Lamarck
(lectotype MNHN DT3345); F, Topsent's spicule preparation of lectotype (scale = 200 pm); G, section
of peripheral skeleton (scale = 500 pm).
J. N. A. Hooper and C. Ltvi
1428
The presence of anisoxeas and a plumo-reticulate extra-axial skeleton in some species of
Ptilocaulis is also similar to Reniochalina, as noted above.
Prior to the present study only four species of Ptilocaulis had been described from the
Australasian region (including New Caledonia), although several others are known worldwide
(e.g. Wiedenmayer 1977). Australasian species include: Spongia echidnaea Lamarck, 1814,
probably from southern Australia (holotype MNHN DT640; Fig. 21A-C) (Topsent 1932;
not Ridley 1884); Ptilocaulis fusiformis Ltvi, 1967, from New Caledonia (see below;
Figs 22-23); Ptilocaulis rigidus Carter, 1883, from southern Australia (?holotype BMNH
1936.5.16.1) (Thiele 1899; Hallmann 1914~;de Laubenfels 1936); and Spongia spiculifera
Lamarck, 1814, from Bass Strait, Vic. (holotype MNHN DT638; Fig. 21E-G) (Ridley 1884;
Dendy 1922; Topsent 1932, 1933; de Laubenfels 1936; Wiedenmayer 1977).
Ptilocaulis fusiformis Ltvi
(Figs 22-23, Table 5)
Ptilocaulis fusiformis Levi, 1967: 21, pl. 1 , fig. b, text-fig. 4.
Material Examined
Holotype. MNHN DCL818: Baie St Vincent, Grand Tenia, New Caledonia, 40 m depth, l0.xii.
1962, coll. Mission Singer-Polignac.
New Caledonian material. QM G300719 (ORSTOM R1547): stn 503, Pointe des Pins, Canal
Woodin, 22"23.4'S.,166"49.6'E.,25-35 m depth, 11.x.1991, coll. G. Bargibant, SCUBA; QM (3301262
(ORSTOM R854): stn 198, pinnacle S. of Canyon Central, Chenal des Cinq Milles, 22"30.4'S.,
166"45.11E.,35 m depth, 15.ii.1978, coll. G. Bargibant, SCUBA; QM (3301324, G301327, (3301335,
G301341: Croisant-Laregritre, IlBt Maitre, off Noumea, 22"20.2'S.,166"22.5'E., 20 m depth, 13.x.
1992, coll. J. N. A. Hooper, SCUBA.
Description
Colour. Live coloration pale orange, yellow-brown (Munsell7.5YR 8/10), pale orangebrown in ethanol.
Shape. Digitate or arborescent digitate, 42-110 mm long, with cylindrical bifurcate
branches, 23-55 mm long, up to 10 mm diameter, tapering towards ends, and with short
basal stalk, 15-19 mm long, 7-10 mm diameter, and broad basal holdfast, 12-21 mm
diameter.
Surface. Highly conulose, with more-or-less evenly distributed conules, up to 5 mm high,
usually forming meandering ridges running longitudinally along branches. Conules interconnected by fleshy surface membrane, pierced by small ostia (visible only between conules),
about 500 pm diameter. Oscules small, 1.5-2 mm diameter, rarely observed, near apex of
branches. Texture firm, flexible.
Ectosome. Fleshy, mostly membraneous surface, with sparse plumose brushes of longer,
usually sinuous megascleres barely protruding through ectosome, restricted to tips of conules.
Ectosomal membrane highly collagenous, more darkly pigmented than choanosomal mesohyl,
and with small quantities of detritus embedded.
Choanosome. Skeletal structure plumo-reticulate, with clearly differentiated axial and
extra-axial regions. Axial skeleton compressed, composed of a heavy spongin fibre skeleton
with individual fibres no more than 150 pm long, up to 50 pm diameter, forming a closeset reticulation producing with oval meshes, 30-90 pm diameter, and cored by plumose,
paucispicular tracts of mostly shorter anisoxeas and fewer sinuous strongyles. Extra-axial
skeleton corresponds with distribution of surface conules. Extra-axial fibres plumo-reticulate,
noticeably more cavernous than in axis, with individual fibres extending for up to 300 pm,
25-35 pm diameter, producing elongate meshes, and extra-axial fibre system runs predominantly laterally through branch cross-section. Fibres cored by multispicular plumose
tracts of both sinuous strongyles and anisoxeas ascending towards surface. Extra-axid
skeletal columns are separated by large cavernous areas (canals, up to 650 pm diameter),
New Caledonian Axinellids
1429
covered by an external collagenous layer (up to 550 pm wide, extending for 750-1250 pm
between surface conules.
Megascleres (refer to Table 5 for dimensions). Two length-classes of megascleres distinguished here, although these clearly intergrade in size and morphology. Longer megascleres
are thin, curved or sinuous strongyles, predominantly in extra-axial region of choanosome
and at surface, although also coring axial fibres, with asymmetrical (styloid), or symmetrical
rounded ends.
Shorter megascleres are slightly curved, thin anisoxeas, found predominantly in the axial
skeleton, although also dispersed in peripheral skeleton, usually with symmetrical rounded
or pointed, usually telescoped ends ('oxeas'), or less often with asymmetrical ends (hastate
points and evenly rounded bases; 'styles').
Microscleres. Absent.
Distribution
Known only from the coral reefs of the New Caledonian lagoon, 25-40 m depth.
Remarks
Although two size classes of megascleres (longer sinuous strongyles and shorter curved
anisoxeas) are differentiated in the above description, these intergrade to some extent and
intermediate 'oxeas' and 'anisoxeas' could fit into either category. There was no observed
regional localisation of these two categories of spicules to any particular region of the
Table 5. Comparison in spicule dimensions between species of Ptilocaulis
All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of
measurement. N = 25 for each specimen
-
Species
(material)
Ptilocaulis echidnaeus (Lamarck) (holotype MNHN DT640)
Ptilocaulis epakros, sp. nov. (holotype QM G300016)
Ptilocaulis fusiformis Levi
(holotype MNHN DCL818)
(QM G300719)
(holotype MNHN DCL818)
(QM G300719)
Ptilocaulis gracilis (Carter) (holotype BMNH 1845.12.30.1)
Ptilocaulis papillatus, sp, nov. (holotype QM (3300748)
Ptilocaulis rigidus Carter (?holotype BMNH 1936.5.16.1)
Ptilocaulis spiculifera (Lamarck) (holotype MNHN DT638)
-
Megascleres
Length
Width
493-(647.8)-816
18-(22.4-29
Styles anisoxeas, oxeas
424-(448.2)-448
1.5-(1.8)-2
Vestigial strongyles
134-(268.4)-328
2.5-(3.4-5
Styles, styloids
414-(664.0)-900
2.5-(3.5)-6
516-(649.3)-743
2-(2.3-3.5
Sinuous strongyles
224-(267.5)-350
6-(8.2)-11
196-(269.9)-339
2,543.3-6
Styles, anisoxeas, oxeas with
telescoped ends
183-(290.6)-562
5-(6.2)-8
Styles, sinuous styles
140-(190.1)-243
1.5-(2.4-4
Vestigial styles, styloid, rarely
strongyloid
252-(364.0)-481
13-(17.6)-21
Styles, oxeas, anisoxeas
242-(291.6)-334
11-(16.9-20
Short robust curved styles
11-(12.2)-14
492-(599.6)-704
Long slender, curved styles
1430
J. N. A. Hooper and C. L h i
skeleton, and thus we cannot confirm Wiedenmayer's (1977) 'typical' diagnosis of these
characters for the genus. Similarly, unlike most other species of Ptilocaulis, it is difficult to
classify the spicules in P. fusiformis as either styles, oxeas or strongyles, due to the great
range in spicule terminations (abrupt points, telescoped endings, evenly rounded bases,
tapering points). LCvi (1967) suggested that the spicules in the holotype were mostly styles,
but re-examination of this material confirmed that a diversity of spicule terminations is
characteristic of the species, although 'true styles' also occur in low numbers.
In its spiculation, the present species is most similar to Rhaphoxya (e.g. R. typica
Hallmann, 1916b: 645, text-fig. 17), and thus the species is virtually intermediate between
Ptilocaulis and Rhaphoxya. However, its characteristic growth form, surface sculpturing
(papillae), well-differentiated axial and extra-axial skeletal structure (including a compressed
axial skeleton) indicate that P. fusiformis is most appropriately placed in Ptilocaulis.
The species is contrasted further with other Australasian Ptilocaulis below, and in Table 5.
Fig. 22. Ptilocaulis fusiforrnis L M , specimen Q M G300719: A, shorter
predominantly axial anisoxeas; B, longer predominantly extra-axial strongyles;
C, section through peripheral skeleton.
New Caledonian Axinellids
Fig. 23. Ptilocaulis fusCformis LCvi: A , holotype (MNHN DCL818); B, specimen
;
[QM G300719 (ORSTOM R1547)l; C, SEM axial fibres (magnified 4 0 0 ~ ) D,
skeleton of holotype (scale= 500 pm); E, SEM skeleton (QM G300719) (magnified
40x).
Ptilocaulis epakros, sp. nov.
(Figs 24-25, Table 5)
Material Examined
Holotype. QM G300016 (ORSTOM R1232): stn 247, Baie Kouo, Canal Woodin, New Caledonia
lagoon, 22°23.5'S.,166049.2'E., 40 m depth, 29.xi.1978, coll. P. Laboute, SCUBA.
Description
Colour. Live coloration pale yellow-brown (Munsell 2.5YR 8/8), beige in ethanol.
1432
J. N. A. Hooper and C. Ltvi
Shape. Arborescent, bifurcate branching, 200 mm long, 70 mm maximum lateral branch
span, with thin, cylindrical branches, 27-60 mm long, 5-17 mm wide including papillae,
tapering towards pointed branch tips, with long, unornamented stalk, 75 mm long, 4 mm
diameter, and expanded basal attachment, 13 mm diameter.
Surface. Heavily ornamented, papillose surface, with long, close-set, sharply pointed,
soft papillae, 2-4 mm long, 0-5-1 mm diameter, up to 2 mm apart; tips of papillae
bifurcate and/or hispid; bases of adjacent papillae interconnected by membraneous ridges
running longitudinally along branches, slightly elevated above surface of sponge. Oscules
not observed; minute ostia, about 200 pm diameter, scattered between papillae.
Ectosome. Fleshy, membraneous ectosome, without specialised spicules, with sparse
detritus embedded in and on surface, with heavy collagenous, aspicular matrix, 150-200 pm
wide, lying between papillae (=surface ridges) and on sides of papillae; apex of each papilla
has plumose brushes of choanosomal styles, in small multispicular bundles, protruding for
short distances, up to 200 pm from surface.
Choanosome. Skeleton structure plumo-reticulate, with clearly differentiated axial and
extra-axial regions, and a compressed axial skeleton. Compressed region of skeleton occupies
only about half (2-3 mm) of branch diameter (3-5 mm), composed of heavy, bulbous
spongin fibres, with individual fibres only about 100 pm long, 50-70 pm diameter, together
Fig. 24. Ptilocaulis epakros, sp. nov., holotype QM G300016:A, styles and
styloids, comprising the majority of spicules; B, sinuous strongyle, relatively
rare; C , section through peripheral skeleton.
New Caledonian Axinellids
1433
forming a close-set reticulation and producing small oval meshes, 50-90 pm diameter; axial
fibres cored by uni-, pauci- or multispicular tracts of thin megascleres, occupying only a
small proportion of fibre diameter. Abundant, lightly pigmented collagen in mesohyl of
axial skeleton. Extra-axial skeleton extensive, including area immediately surrounding axis
of branches (1-2 mm diameter) as well as elongated, slender skeletal columns (=papillae;
up to 4 mm long). Extra-axial skeleton composed of primary and secondary fibre systems,
differentiated mainly by presence or absence of coring spicules; both fibre systems composed
of heavy spongin fibres, with individual fibres up to 300 pm long, 40-60 pm diameter,
producing a relatively wide, elongate mesh reticulation, 110-170 pm mesh diameter;
ascending extra-axial fibres cored by multispicular, plumose columns of choanosomal styles,
with spicule tracts becoming heavier towards peripheral skeleton [eventually terminating in
plumose brushes which may or may not protrude through surface (tips of papillae, versus
between papillae respectively)]. Many (but not all) transverse, connecting fibres in extraaxial skeleton uni- or aspicular, with long exceedingly slender strongyles (although these were
invariably broken in situ, and consequently were not observed in spicule preparations).
Collagen abundant and slightly more darkly pigmented in extra-axial region; choanocyte
chambers not observed.
Megascleres (refer to Table 5 for dimensions). Two categories of spicules present,
clearly differentiated in morphology but not obviously localised to any particular region of
skeleton. Few transverse, connecting fibres contain a single, long, thin, sinuous strongyle,
but these spicules are rare.
Majority of spicules are styles or styloids, short or long, slender, straight or slightly
curved asymmetrically, with evenly rounded or tapering mucronate bases, and hastate,
fusiform or telescoped points.
Fig. 25. Ptilocaulis epakros, sp. nov.: A, SEM skeletal structure (scale= 1 mm);
B, holotype [QM G300016 (ORSTOM R1232)]; C, holotype in situ (photo
P . Laboute).
J. N. A. Hooper and C. LCvi
Microscleres. Absent.
Distribution
Known only from the New Caledonian lagoon, inter-reef region, 40 m depth.
Remarks
This species is obviously a closely related, sibling species of Ptilocaulis fusiformis. In its
external morphology, including shape and size of surface papillae, it is easily differentiated
from P. fusiformis, whereas there are only subtle differences in the geometry and size of
megascleres (Table 5), and in the structure of the skeleton between the two species (Figs 22,
23 cf. Figs 24, 25). In fact, all three sympatric sibling species, P. fusiformis, P. epakros
and P. papillosus, are most visibly differentiated only by these features. It is conceivable
that all three nominal species are members of a single, extremely polymorphic species, but
gross external morphology and more subtle skeletal differences do not presently support
this hypothesis, and the three taxa are retained here. Only molecular evidence will support
or refute this hypothesis.
In its arborescent growth form, with thinly cylindrical bifurcate branches and long,
unornamented stalk, and its long, tapering, sharply pointed surface conules, P. epakros
shows remarkable superficial resemblance to Thrinacophora funiformis Ridley & Dendy
(1887: pl. 24, fig. 1) (Raspailiidae), although obviously in spiculation and skeletal architecture the two species are otherwise quite different.
Etymology
Named for the long, attenuated, sharply pointed surface papillae, from epakros (Gk),
pointed at the end.
Ptilocaulis papillatus, sp. nov.
(Figs 26-27, Table 5)
Material Examined
Holotype. QM G300748 (ORSTOM R564): stn 113, L'epave du 'Bonhomme', Grand RCcif Mbkre,
22°21.0'S.,166014~(YE.,25 m depth, 21.vi.1976, coll. P. Laboute, SCUBA.
Description
Colour. Live coloration unknown, white in ethanol.
Shape. Elongate, cylindrical digitate, 120 mm long, 14 mm wide at widest point, 4 mm
wide at end of branch, single branching and bifurcate tips at end of branches, with short
(unornamented) stalk, 17 mm long, 5 mm diameter, and enlarged basal attachment, 16 mm
diameter.
Surface. Prominently sculptured surface composed of large discrete papillae, each papilla
formed by extra-axial skeletal columns, up to 6 mm long, 2.5 mm diameter, standing
perpendicular to axial core, dispersed approximately 1.5-3 mm apart, enlarged, flattened
and bifurcate at apex, usually interconnected by translucent dermal membrane, but cavernous
below membrane.
Ectosome. Translucent dermal membrane, collagenous, aspiculose except at ends of
extra-axial skeletal columns (papillae) where tufts of megascleres protrude slightly through
surface.
Choanosome. Skeletal structure plumo-reticulate, with very well differentiated axial and
extra-axial skeletons, and heavily compressed axial skeleton. Spongin fibres dominate both
sections of skeleton; in axial region fibres heavy, close-set, up to 150 pm long, 50 pm
diameter, bulbous, meandering, producing oval, elongate or irregular meshes, 30-140 pm
diameter, with heavy collagenous mesohyl; in extra-axial skeleton fibres less heavily compacted, with individual fibres up to 250 pm long, 30 pm diameter, producing elongated
New Caledonian Axinellids
Fig. 26. Ptilocaulis papillatus,
sp. nov., holotype QM G300748:
A , vestigial styles, styloids and
strongylote spicules; B, section
through peripheral skeleton.
Fig. 27. Ptilocaulis papillatus, sp. nov.: A , holotype (QM G300748), SEM skeletal
structure (scale = 1 mm); B, SEM axial fibres (scale = 200 pm); C, SEM fibre structure
(scale = 100 pm); D, holotype.
J. N. A. Hooper and C. LCvi
1436
meshes up to 170 pm diameter, more-or-less directed towards surface, and fibres become
thinner and more widely spaced near ectosome. Spicule skeleton nearly vestigial, with very
few megascleres in axial region, but becoming increasingly abundant towards surface; extraaxial spicule skeleton plumose, ending with plumose bundles of megascleres protruding
slightly through surface, at apex of skeletal columns (papillae). Area between skeletal
columns usually cavernous, without collagen, fibres or mineral skeleton, measuring up to
2 mm between adjacent columns. Collagen in extra-axial skeletal columns abundant but
lightly pigmented; choanocyte chambers small, oval, 20-40 pm diameter.
Megascleres (refer to Table 5 for dimensions). Most spicules vestigial, with blackened
axial canals, exclusively styles or styloids, rarely strongylote, usually slightly curved, sometimes sinuous, with evenly rounded or tapering bases and hastate points, not telescoped.
Microscleres. Absent.
Distribution
Known only from the New Caledonian lagoon, in the inter-reef region on sandhubble
substrate.
Remarks
This species is most similar to the sympatric P. fusiformis in its skeletal structure, growth
form and spiculation, except that all these characters are greatly exaggerated in P. papillatus.
Surface papillae are exceptionally large and bifurcate, and rejoin with adjacent papillae at
their apex by a translucent dermal membrane; spongin fibre compression is relatively greater
in both axial and extra-axial regions; spicules are virtually vestigial in most of the skeleton,
except towards the surface where they become exclusively plumose; and spicules are
exclusively styloid, without any apparent differentiation in size (Table 5). These features
clearly differentiate this species from P. fusiformis, whereas differences between
P. fusiformis and P. epakros are more subtle.
Etymology
Named for the unusual surface sculpturing, from papillatus (Lat.), bud-like, papillose.
Genus Pseudaxinella Schmidt
Pseudaxinella Schmidt, 1875: 120.-Thiele, 1903: 378; Bergquist, 1970: 20; LCvi, 1973: 606;
Wiedenmayer, 1977: 155 [type species Pseudaxinella sulcata Schmidt, 1875: 1201.
Diagnosis
Usually massive, subspherical, cushion-shaped, unbranched or lobate growth forms, with
finely conulose or tuberculate, corrugated surface. Skeleton typically plumo-reticulate,
without axial compression or differentiation between axial and extra-axial regions; skeletal
tracts consist of oxeas and styles, often in crowded, ascending tracts. Ectosome fleshy, and
megascleres in this region may be thinner than choanosomal spicules, but not definitely so.
Megascleres typically include only (anis-)oxeas and styles in equal proportions, but some
species also have long flexuous styles or strongyles confined to the surface. Microscleres
absent (modified from Wiedenmayer 1989).
Remarks
The presence, absence or modification of megascleres to long flexuous diacts (strongyles)
probably has questionable diagnostic value in the Axinellidae. Vacelet (1969) and Pansini
(1983) have shown that the traditional distinction between Axinella and PhakeNia (cf.
Vosmaer 1912), based on external form and the presence of flexuous diactines in the axial
skeleton, is not reliable. Consequently, the importance of those spicules in diagnosing
Pseudaxinella may also be of doubtful systematic value (Wiedenmayer 1989). Nevertheless,
in the absence of a reliable, comprehensive revision of the Axinellidae, incorporating all taxa
that have sinuous strongylote modifications, it is not possible to evaluate whether or not
New Caledonian Axinellids
1437
those megascleres occur consistently or at what level they have systematic value. Consequently, definitions of Pseudaxinella provided by Ridley and Dendy (1887), Vosmaer (1912,
1935a), Babic (1922) and Topsent (1934) must be treated with circumspection.
Pseudaxinella was originally erected for species 'like Axinella' (i.e. in spiculation), but
lacking axial compression (de Laubenfels 1950); but Wiedenmayer (1989) noted that some
species [e.g. P. convexa (Hoshino, 1981), P. decipiens Wiedenmayer, 19891 have nearly
confused skeletons, atypical of the genus, although agreeing in all other respects. Thus, in
Wiedenmayer's (1989) opinion, skeletal organisation may be a poor diagnostic character for
these groups, whereas features such as the absence of a special axial skeleton and external
morphology (massive subspherical in Pseudaxinella, thinly flabellate in Axinosia/Teichaxinella)
might be more reliable diagnostic features. However, rightly or wrongly, the present scheme
of classification for the Axinellidae differentiates all constituent genera at least partly on
the basis of their skeletal construction, and without revising this current basis for the
classification, which is unrealistic in the present work, we propose to retain this character
[i.e. the non-compressed, plumose (plumo-reticulate) spicule skeleton] for now. The genus
is discussed further at length by Wiedenrnayer (1989: 48).
Prior to the present work, only three species had been recorded from the Indo-west
Pacific: Pseudaxinella australis Bergquist, 1970, from northern New Zealand (holotype
NMNZ Por.26 [not seen]), also recorded recently from northern Australia (Hooper, unpublished; Fig. 30, Table 6); Pseudaxinella decipiens Wiedenmayer, 1989, from Bass Strait, Vic.
(holotype NMV F51961 [not seen]); and Phakellia tumida Dendy, 1897a, from Port Phillip,
Vic. (holotype NMV G2464 [not seen]) (Vosmaer 1912). A fourth species is also known from
the neighbouring province, Ariake Sea, Japan: Axinella convexa Hoshino, 1981 (holotype
MMBS AR-1-11 [not seen]).
Pseudaxinella debitusae, sp. nov.
(Figs 28-29, Table 6)
Material Examined
Holotype. QM G300725 (ORSTOM 'cfR806'): stn 124, I1Bt Maitre, 22°20.1'S.,166025. l1E., 25 m
depth, 1.x.1991, coll. G. Bargibant, SCUBA.
Paratypes. QM G300722 (ORSTOM 'cfR806'): stn 181, Ilat Maitre, 22°20.0'S.,166025.0'E.,10 m
depth, 1.x.1991, coll. J.-L. Menou, SCUBA; NTM 23887, QM G300695: Baie des Citrons, off
Noumea, 22°20'S.,166027'E., 3 m depth, 25.ix.1990, coll. J . N. A. Hooper, snorkel, stn JH-90-019.
Other New Caledonian material. QM G301263 (ORSTOM R1227), ORSTOM 'cfR1221': stn 261,
SW. IlBt Nda, Lagon Sud, 21°52.5'S.,166"51.2'E.,33 m depth, 4.xii.1979, coll. P. Laboute, ORSTOM,
SCUBA; QM G301319, (3301328, G301332: Croisant-Laregribre, IlBt Maitre, off Noumea, 22"20.2'S.,
166'22.5'E., 20 m depth, 13.x.1992, coll. J. N. A. Hooper, SCUBA.
Description
Colour. Orange to orange-yellow alive (Munsell 10R 6/10-2.5YR 7/8), beige or light brown
in ethanol.
Shape. Massive, irregularly or regularly subspherical, cushion shaped, 55-80 mm
diameter, 32-40 mm maximum height above substrate, loosely attached to large pieces of
detritus (e.g. dead coral, pelecypod valve), or occasionally rolling free on the substrate (i.e.
'tumbleweed' effect).
Surface. Evenly microconulose, goose-flesh appearance, covered by small conules,
1-2 mm diameter, less than 0.5 mm high, scattered over entire surface, interconnected by
semi-translucent dermal membrane. Oscules scattered over 'upper' surface, large in life
(5-10 mm diameter), contracted in ethanol (1-2 mm diameter), located in slight depressions
on surface and surrounded by slightly raised membraneous lip. Texture soft, compressible,
relatively easy to tear.
Ectosome. Membraneous, fleshy surface, with tips of choanosomal spicules protruding
for short distances, up to 150 pm, from surface in sparse plumose brushes. Heavy, more
darkly pigmented collagen clearly marks peripheral region, whereas in choanosomal mesohyl
J. N. A. Hooper and C. L h i
1438
collagen is only lightly pigmented. Oval choanocyte chambers in peripheral skeleton, 70190 pm diameter, also clearly outlined by more darkly pigmented, granular spongin.
Choanosome. Skeleton plumo-reticulate, without axial compression or any noticeable
difference between axial and extra-axial regions. Spongin fibre skeleton reticulate, with
predominantly ascending primary fibres, up to 70 pm diameter, interconnected by shorter,
thinner secondary fibres, 40-60 pm diameter, together producing oval meshes 70-140 pm
diameter. Spicule skeleton plumo-reticulate, with clearly differentiated primary, ascending,
multispicular tracts, cored by spicules in plumose bundles, interconnected by secondary, unior paucispicular, more-or-less transverse spicule tracts, and spicule reticulation producing a
vaguely subrenieroid reticulate skeleton, although the plumose component is emphasised over
the reticulate component. Mesohyl moderately light, lightly pigmented.
Megascleres (refer to Table 6 for dimensions). Spicules predominantly oxeas, with
rare styles and strongyloxeas also present; all spicules relatively long, slender, usually
asymmetrically curved (but not rhabdose), sometimes straight, mostly sharply pointed,
fusiform, although telescoped and bifurcate points also observed.
Microscleres. Absent.
Table 6.
Comparison in spicule dimensions between similar species of Pseudaxinella
All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of
measurement. N = 2 5 for each specimen
-
Species (Locality)
(material)
Pseudaxinella australis Bergquist (New Zealand)
(holotype; Bergquist 1970: 20)
(Great Barrier Reef) (QM G300295, G301089, G300880;
Hooper, unpublished)
Pseudaxinella debitusae, sp. nov. (New Caledonia)
(holotype)
(paratypes)
Pseudaxinella convexa (Hoshino, 1981: 207)
Megascleres
Length
Width
203-(402)-560
9-(15.0)-22
Styles, slightly rhabdose bases
320-(367)-406
3-(4.0)-6
Thinner ectosomal styles
217-(260)-339
8-(9.6)-10
Centrally curved oxeas
172-(274.0)-312
10-(13.2)-16
Styles, slightly rhabdose bases
252-(289.2)-315
8-(11.8)-15
Centrally curved oxeas
243-(358.5)-503
4-(9.6)-15
223-(369.9)-483
2-(7.8)-12
Predominantly asymmetrical oxeas,
rare styles or strongyloxeas
550-(760)-920
10-(18)-26
Oxeas, occasionally styles
330-950
9-20
Sinuous oxeas, strongyloxeas
Pseudaxinella decipiens Wiedenmayer (1989: 48)
(holotype)
Pseudaxinella rosacea (Verill) (de Laubenfels, 1950)
Pseudaxinella tumida (Dendy, 1897a: 237)
Oxeas, styles, strongyloxeas,
anisoxeas
542-770
5-9
Sinuous strongyles
235-400
8-1 1
Styles
300-320
8
Oxeas
About 180
About 6
Styles only
New Caledonian Axinellids
1439
Distribution
Known only from the New Caledonian lagoon, subtidal fringing coral reefs, coral rubble
substrate, 3-33 m depth.
Remarks
This species is closely related to Pseudaxinella australis Bergquist, to which it was initially
assigned. However, a detailed comparison between the New Caledonian material described
above, the New Zealand holotype of P. australis (Bergquist 1970: 20), and another three
specimens of P. australis from northern Australia (QM G300880, Moreton Bay, Qld;
G300295, Snake Reef, northern Great Barrier Reef, Qld; G301089, Cartier I., Sahul Shelf,
W.A.; Fig. 30, Table 6), showed quite a number of differences.
The New Caledonian material was pale orange in life, and this colour is probably truly
representative of the population as it was observed in all three specimens from the lagoon.
In contrast, both New Zealand and Great Barrier Reef material was consistently bright red
in life, and all known material was observed to exude clear mucus, whereas this trait was
not observed in New Caledonian specimens. Pseudaxinella debitusae has a plumo-reticulate
skeleton, with adjacent plumose, ascending skeletal columns interconnected by a paucispicular subreneiroid skeleton, whereas in P. australis the ascending, plumose spicule tracts
are more or less discrete with few interconnections (cf. Figs 28B, 29D-E and 30B). In New
Caledonian specimens, the plumose spicule brushes protruding through the ectosome were
not noticeably thinner than those in the choanosome, unlike the holotype of P. australis
(although these 'ectosomal spicules' were not observed in Great Barrier Reef material
-
Fig. 28. Pseudaxinella debitusae, sp. nov., paratype QM G300722: A , oxeas, styles
and strongyloxea structural spicules; B, section through peripheral skeleton.
1440
J. N. A. Hooper and C. LCvi
either). Similarly, there was no apparent localisation of different spicule morphologies in
P. debitusae (either oxeas, or the rarer styles or strongyloxeas) to any particular region of
the skeleton, whereas the plumose spicule columns of P. australis are usually made up
of a central core of oxeas surrounded by plumose brushes of styles (the styles verging on
echinating). Bergquist (1970) also noted that, in the holotype, oxeas become more abundant
towards the peripheral skeleton, but this was not confirmed in northern Australian material
of P. australis. Most megascleres observed in P. debitusae were strictly oxeote, with far
fewer 'abnormal terminations' (styloid, strongyloid) than seen in P. australis, in which
Fig. 29. Pseudaxinella debitusae, sp. nov.: A , paratypes (NTM 23887, QM
G300695); B, paratype (QM (3300722); C, specimen in situ (ORSTOM R1227)
(photo P. Laboute); D, peripheral skeleton (QM G300722) (scale=500 pm); E,
SEM skeleton (magnified 50 x ) .
New Caledonian Axinellids
Fig. 30. Pseudaxinella australis
Bergquist: A , specimens from
northern Australia on deck (QM
G300880); B, skeletal structure o f
specimen Q M G300295 (scale = 500
oxeas and styles occur in about equal proportions. Bergquist (1970) also notes that the
texture of the holotype was firm, compressible and brittle, which was similar for the
northern Australian material of P. australis, whereas P. debitusae is distinctly soft and
compressible, easily torn, probably reflecting the relatively higher spongin fibre content of
New Caledonian sponge.
Bergquist (1970) contrasts P. australis further with a similar species from Bermuda,
P. rosacea, and other Pseudaxinella described from the Indo-west Pacific are also compared
in Table 6 . It is our opinion that these subtle and more gross differences between the New
Caledonian species and other species of Pseudaxinella justify the creation of a new taxon
for this population, although we recognise that the allopatric populations of northern New
Zealand, south-east Queensland, Great Barrier Reef, and the Sahul Shelf are closely allied
species of this New Caledonian form.
Etymology
Named for Dr CCcile Debitus, ORSTOM Noumea, in appreciation for facilitating our
access to the vast ORSTOM collections, and for her role in the organisation of the collaborative workshops, leading to the publication of the present series of papers on the
New Caledonian sponge fauna.
Genus Rhaphoxya Hallmann
Rhaphoxya Hallmann, 1916b: 641.-de Laubenfels, 1936: 136; Bergquist, 1970: 18 [type species
Rhaphoxya typica Hallmann, 1916b: 643 (holotype AM Z1595)l.
[Acnnthellina] Carter, 1885: 365 [nomen oblitum; ICZN 50 year rule].-de Laubenfels, 1936: 139;
Bergquist, 1970: 18 [type species Acanthellina rugolineata Carter, 1885: 139, holotype BMNH
1886.12.15.941.
Diagnosis
Massive growth form; surface with papilliform conules. Skeleton not axially condensed,
without axial and extra-axial differentiation, consisting of loose, irregularly reticulate, often
meandering tracts of spongin fibres and spicules; spicule tracts may (s.s) or may not
protrude through ectosome. Ectosome fleshy, lacking any specialised spiculation. Spicules
often sinuous or curved, slender, monactinal and/or diactinal (styles, oxeas and strongyles,
1442
J. N. A. Hooper and C. LCvi
of one size category, differing only in the character of their extremities). Microscleres
raphides, occurring singly or in bundles (trichodragmata) (modified from Hallmann 1916b).
Remarks
Under van Soest et al.'s (1990) scheme for the distribution of axinellid genera into four
families, based on skeletal architecture (i.e. Axinellidae with axially compressed and extraaxially plumoreticulate skeletons; Desmoxyidae with reticulate skeletons; Dictyonellidae with
dendritic skeletons; and Halichondriidae with 'halichondroid' disorganised skeletons),
Rhaphoxya would fall under Dictyonellidae, together with other genera with atypical, noncompressed, meandering skeletal structure, such as Acanthella. Although we do not currently
subscribe to this division of Axinellidae and sister-groups, it does illustrate the problem in
classifying axinellids, especially in relying mostly (or exclusively) on skeletal structure.
Rhaphoxya does not fit easily with other axinellid genera that have typical skeletal
structure (compressed reticulate axial skeleton and plumoreticulate extra-axial skeleton),
but it also shows many similarities to Ptilocaulis, as noted above, which does have a typical
Fig. 31. Rhaphoxya spp.: A, Acanthella cactiforrnis Carter, skeleton of lectotype
; lectotype; C, Rhaphoxyafelina Wiedenmayer,
(BMNH 1886.12.15.81) (scale =500 ~ m )B,
peripheral skeleton of specimen (NCI Q66C-3339-U) (photo NCI); D, specimen; E,
Rhaphoxya pallida Dendy, choanosomal skeleton of specimen (QM G300469) (scale =
500 pm); F, specimen from northern Queensland.
New Caledonian Axinellids
1443
Fig. 32. Rhaphoxya spp.: A , Acanthella rugolineata Carter (holotype BMNH
1886.12.15.24); B, peripheral skeleton (scale =SO0 pm); C, Axinella solida Carter
(holotype BMNH 1887.7.11.24); D, peripheral skeleton (scale = 500 pm).
axinellid skeleton. These similarities are especially seen by comparing spiculation of
Ptilocaulis fusiformis, P. papillatus and P. epakros with Rhaphoxya systremma, sp. nov.
(Figs 22, 24, 26 and 33).
Bergquist (1970) suggested that Rhaphoxya was also close to Desmoxya (Desmoxyidae)
in spiculation and architecture, but differed in having a more lax, less halichondroid
skeleton, and in lacking centrangulate spined microscleres. In fact, Bergquist (1970)
previously included Rhaphoxya with the Desmoxyidae, whereas Wiedenmayer (1989)
referred it to the Axinellidae, illustrating its closer relationships with other axinellids such
as Axinyssa.
Prior to the present study, six species were assigned to the genus, all recorded from
Australasian waters: Acanthella cactiformis Carter, 1885, from Port Phillip Heads, Vic.
(lectotype BMNH 1886.12.15.81; Fig. 31A-B) (Carter 1885; Dendy 1897a; Burton 1934;
Bergquist 1970; Wiedenmayer 1989); Rhaphoxya felina Wiedenmayer, 1989, from Bass
Strait, Vic. (holotype NMV F51964 [not seen]; specimen NCI Q66C-3339-U; Fig. 31C-D);
Rhaphisia pallida Dendy, 1897a: 257, from Bass Strait, Vic., and far north Queensland
(holotype NMV G2377 [not seen]; specimen QM G300469; Fig. 31E-F) (Hallmann (1916b:
646); Acanthella rugolineata Carter, 1885: 365, from Port Phillip Heads, Vic. (holotype
BMNH 1886.12.15.24; Fig. 32A-B); Axinella solida Carter, 1885, from south-east Australia
(lectotype BMNH 1887.7.11.24; Fig. 32C-D) (Dendy 1897a); and Rhaphoxya typica
Hallmann, 1916b: 643, also from Port Phillip Bay, Vic., and Tasmania (holotype AM 21595
[not seen]) (Guiler 1950: 9).
Rhaphoxya systremma, sp. nov.
(Figs 33-34, Table 7)
Material Examined
Holotype. QM G300013 (ORSTOM R1221): stn 261, SW. Ilbt Nda, Lagon Sud, 21°52.5'S.,
166'51.2'E., 33 m depth, 4.xii.1979, coll. P. Laboute, ORSTOM, SCUBA.
J. N. A. Hooper and C. L&i
1444
Paratype. QM G300442 (fragment NTM 21500): Euston Reef, NE. of Cairns, Great Barrier Reef,
Qld, 16°43.0'S.,146013.8'E., 42.5 m depth, 26.i.1981, coll. QFS, dredge (stn B6, Cairns ground
truth survey).
Description
Colour. Pale or dark orange-brown alive (Munsell 7.5 YR 7/10-5/6), beige (holotype)
t o dark brown (paratype) in ethanol.
Shape. Spherical o r subspherical, globular growth form, 32-75 m m high, 28-60 m m
diameter, consisting of aggregated, globular lamellae ('lacunae' of earlier authors), producing
a conglomerated, honeycombed, Echinoclathria-like reticulation with numerous, oval, celllike cavities and large canals excavating entire sponge. Sponges are only loosely attached
t o pieces of coral rubble or shell fragments, or occasionally rolling free o n substrate
('tumbleweed' sponges).
Surface. Membraneous, gelatinous, irregularly convoluted surface, with prominent,
rounded papillae, u p t o 3 m m high, 2 mm diameter, most abundant o n apical surface of
sponge; largest papillae near apex ('upper surface') surround 1 or more oscules, 2-4 m m
diameter, although oscules also occur in other places on the 'upper surface', such as in ridges
between surface papillae. Texture soft, compressible, difficult t o tear.
Ectosome. Fleshy, heavily collagenous, darkly pigmented ectosomal region, without
spongin fibres or spicules, but with a thick collagen layer (up t o 80-300 pm thick) between
surface and beginning of choanosomal spongin fibre skeleton; this collagenous layer is
thicker in between the surface ridges and papillae than o n top of these structures; sparse
Table 7. Comparison in spicule dimensions between Rhaphoxya species
All measurements given in micrometres, and expressed as range of measurement
Species (locality)
[source of information]
Rhaphoxya cactiforrnis (Carter) (Port Phillip Bay)
[lectotype BMNH 1886.12.15.811
Rhaphoxya felina Wiedenmayer (Bass Strait)
[specimen NCI Q66C-3339-U]
Rhaphoxya pallida (Dendy) (Port Phillip Head
and Bass Strait)
[specimen QM G3004691
Rhaphoxya rugolineata (Carter) (Port Phillip Bay)
[holotype BMNH 1886.12.15.241
Rhaphoxya solida (Carter) (Port Phillip Head
and Bass Strait)
[lectotype BMNH 1887.7.11.241
Megascleres
Length
Width
Raphides
167-432
4-12
Absent
Predominantly oxeas with telescoped
ends, some styles
192-415
5-12
110-245
Exclusively oxeas
211-430
4-9
110-370
Predominantly oxeas with telescoped
ends, some anisoxeas and
strongyloxeas
305-5 14
7-12
Absent
Predominantly oxeas with telescoped
ends, some styles
232-388
4-8
65-146
Predominantly styles with hastate or
teiescoped points, some oxeas and
anisoxeas
Rhaphoxya systrernrna, sp. nov. (New Caledonia
and northern Great Barrier Reef)
[holotype QM G3000131
[paratype QM G3004421
Rhaphoxya typica Hallmann (Port Phillip Bay
and Bass Strait)
[Hallmann 1916b: 6461
Absent
248-(304.6)-369 2-(2.8)-4
Absent
201-(299.8)-382 2-(3 3-5
Predominantly vestigial, sinuous
strongyles, rarer oxeas with
telescoped ends
100-700
2-9
55-400
Predominantly oxeas with telescoped
ends, fewer styles or strongyles
New Caledonian Axinellids
1445
plumose brushes of choanosomal spicules may also protrude through surface, up to 100 pm,
especially on tips of papillae, otherwise entire ectosomal skeleton is collagenous. Sparse
deposits of detritus also dispersed over surface and incorporated into ectosomal collagenous
layer, but this is not a prominent feature of the skeleton.
Choanosome. Skeleton plumose, slightly plumo-reticulate, without axial compression
or differentiation between axial and extra-axial skeletons; skeleton is dominated mainly by
diverging, meandering, sinuous spongin fibres and spicule tracts; spongin fibre system
composed of ascending, primary fibres, with individual fibres up to 350 pm long, 5070 pm diameter, cored by multispicular tracts of choanosomal megascleres, and interconnected by shorter, transverse, secondary fibres, up to 120 pm long, 20-40 pm wide,
usually aspicular or sometimes paucispicular; generally the reticulate, connecting, secondary
spicule tracts are greatly reduced in proportion to the primary, plumose, ascending skeleton.
Fibre reticulation produces elongate, often cavernous meshes, 70-230 pm long, up to
80 pm wide, becoming more cavernous in periphery than in axis, with some scattered,
sinuous spicules outside fibres; choanosomal mesohyl with abundant but only lightly
pigmented collagen, with only few, small choanocyte chambers seen, up to 30 pm diameter.
Megascleres (refer to Table 7 for dimensions). Single category of choanosomal spicule
present, relatively homogeneous in size, although terminations of spicules vary from
symmetrical, evenly rounded and hastate tapering rounded ends (strongyles), to sharply
pointed, telescoped ends (oxeas). Majority of spicules strongylote, sinuous, very slender.
Microscleres. Absent.
Distribution
New Caledonian lagoon and northern Great Barrier Reef, coral reef rubble and inter-reef
region, 30-43 m depth.
Fig. 33. Rhaphoxya systremma, sp. nov., holotype QM G300013: A,
strongyles and oxea structural spicules; B, section through pheripheral
skeleton.
1446
J. N. A. Hooper and C. Levi
Remarks
This is the first record of the genus outside southern Australian waters. Rhaphoxya
systremma differs from most other species of the genus in lacking raphide microscleres
(four of the six described species have raphides), having exceedingly thin megascleres with
predominantly strongylote endings (most other species have predominantly or exclusively
styles or oxeas with telescoped ends; Table 7), and having i?i sinuous, virtually non-reticulate
skeleton (similar to R. typica).
According to their published descriptions, all six previously described species of
Rhaphoxya are morphologically very similar and difficult to clearly distinguish on the basis
of their published gross characteristics, including growth form, surface features, skeletal
architecture and spiculation. However, examination of type material and other representatives
of these species shows a number of distinctive, cryptic characters which support their
differentiation (Figs 31-34). Rhaphoxya felina has a distinctive, perpendicular ectosomal
skeleton, a more confused plumo-reticulate choanosomal skeleton, and megascleres are
exclusively oxeas; R. cactiformis has a more confused plumo-reticulate skeleton, heavy
collagenous mesohyl, spicules are predominantly oxeas with telescoped ends, and raphides
are absent; R. rugolineata has a virtually plumose, diverging spicule skeleton in which the
outer layer of spicules on spicule tracts are inserted at acute angles (i.e. nearly echinating),
spicules are predominantly oxeas with telescoped ends, and raphides are absent; R. solida
has a skeleton similar to that of R. felina, with a perpendicular ectosomal skeleton and
compact, confused, plumo-reticulate skeletal tracts, and spicules are predominantly
Fig. 34. Rhaphoxya systremma, sp. nov.: A , cross-section through surface papilla (holotype
QM G300013) (scale=500 pm); B, holotype; C, specimen in situ (ORSTOM R1221) (photo
P. Laboute).
New Caledonian Axinellids
1447
asymmetrically curved styles with hastate or telescoped points, with less frequent anisoxeas
and oxeas; R. pallida has a meandering plumo-reticulate skeleton similar to that of the
present species, with plumose (plumo-reticulate) spicule tracts and a heavy collagenous
mesohyl, but spicule tracts are not sinuous and spicules are predominantly oxeas with
telescoped ends; R. typica also has a meandering, plumo-reticulate skeleton, similar to that
of R. systremma, and its growth form is similar in being massive, subspherical, with papillae
on the upper surface, but spicules are substantially larger and consist predominantly of
oxeas with telescoped ends, with only few strongyles or styloid modifications (which
are predominant in the present species), and raphide microscleres are also abundant.
A comparison of spicule dimensions between these species is given in Table 7. Altogether,
the present species appears to be most closely related to, and clearly a sibling species of,
the southern Australian R. typica.
Etymology
The species name refers to the subspherical conglomeration of lacunae, forming a honeycombed mass, from the Greek systremma, relating to anything aggregated, consolidated,
or twisted together into a round object.
Family DESMOXYIDAE Hallmann
Definition
Axinellid sponges with 'microscleres' in the form of smooth or spined microxeas, often
centrangulate or strongly bent centrally, sometimes also with raphides, occurring singly or
in bundles (trichodragmata), or acanthose cladotoxa and birotules in one genus. Megascleres
monactinal, diactinal or both. Skeleton consisting of widely spaced reticulate bundles of
multispicular fibres, with little spongin, with poorly developed or no axial compression, and
relatively poorly differentiated extra-axial skeleton (disorganised-plumose). Growth forms
encrusting, massive or ramose [compiled from Hartman (1982) and Wiedenmayer (1989)l.
Remarks
The presence of smooth or spined oxeas is characteristic of, and apomorphic for,
desmoxyid genera. These spicules are traditionally classed as microscleres (e.g. Hallmann
1916b, Wiedenmayer 1977), although in comparison to many other taxa they are generally
much too large to be considered as such (e.g. Myrmekioderma). Wiedenmayer (1977)
suggested that the Desmoxyidae had affinities with the Hemiasterellidae (e.g. Laonaenia
Hallmann), and the Hadromerida (e.g. Paratimea Hallmann). However, Topsent (1928) had
previously included these genera in the Hemiasterellidae, but he was not followed by subsequent workers. Although some similarities can be drawn between some taxa of both
families, the hemiasterellids are restricted to forms with asterose microscleres, whereas the
desmoxyids included forms with microxeote spicules. On that basis, several genera currently
assigned to Bubaridae (e.g. Rhabdoploca, Bubaropsis) also have inferred affinites with the
Desmoxyidae, having acanthose or smooth oxeote megascleres.
This family was established by Hallmann (1916b: 673; with synonym Higginsiinae de
Laubenfels, 1936: 132), initially for five genera, of which only four were correctly assigned
(Desmoxya, Higginsia, Holoxea and Halicnemia). Numerous other genera were subsequently
added or associated with the family, although affinities between them were not always
completely clear. Of these, the following genera are excluded from the family: Negombo
Dendy (type species N. tenuistellata Dendy) and Diacarnus Burton (type species Axos
spinipoculum Carter) are probably best assigned to the Latrunculiidae [both suggested as
possible synonyms according to Hooper (1986), but this has not been firmly established];
Allantella Hallmann (type species Trachytedania arborea Keller) is probably a hadromerid;
Laonaenia Hallmann (type species Hymeraphia verticillata Bowerbank), and Paratimea
Hallmann (type species Bubaris constellata Topsent) are aster-bearing taxa, both belonging
to Halicnemia Bowerbank according to Topsent (1928), but are probably hemiasterellids or
hadromerids (e.g. Hooper 1986). Ommatosa (sensu de Laubenfels 1936; type and only
J. N. A. Hooper and C. LCvi
1448
species Axinella rugosa Schmidt) is either a desmoxyid or a bubarid; its placement is
problematic, but it probably has closest affinities with Bubaris.
Eight genera are considered to be valid and presently included in Desmoxyidae (Acanthoclada Bergquist, Halicnemia Bowerbank, Heteroxya Topsent, Higginsia Higgin, Holoxea
Topsent, Microxistyla Topsent, Myrmekioderma Ehlers and Parahigginsia Dendy) but, as
van Soest et al. (1990) suggest, there is a pressing need for a complete revision of the family
and a re-evaluation of its constituent genera.
Only one species of Desmoxyidae has been recorded previously for the New Caledonian
region, Parahigginsia phakellioides Dendy, 1924, also known from northern New Zealand.
Genus Myrmekioderma Ehlers
Myrmekioderrna Ehlers, 1870: 28. - Bergquist, 1965: 177 [type species Alcyonium granulaturn Esper,
1830: 711.
Acanthoxifera Dendy, 1905: 156. -Dendy, 1922: 129; Bergquist, 1965: 177 [type species A. ceylonensis
Dendy, 1905: 1571.
Anacanthaea Row, 1911: 329.-van Soest et al., 1990: 31 [type species A . nivea Row, 1911: 3291.
Callistes Schmidt, 1868: 16.-van Soest et al., 1990: 31 [type species C. lacazii Schmidt, 1868: 161.
Neoprosypa de Laubenfels, 1954: 189 [type species N. atina de Laubenfels, 1954: 1901.
Diagnosis
Growth form massive or encrusting; surface hispid, with characteristic canals and grooves
forming polygonal tuberculate plates. Choanosome condensed, with confused mass of
acanthoxeas and oxeas, strongyles or less frequently styles, forming irregular, ascending,
multispicular tracts bound together with sparse collagen. Extra-axial skeleton dense paratangential layer of acanthoxeas, with larger choanosomal styles protruding. Ectosomal
skeleton without specialised spiculation, subectosomal acanthoxeas protrude, forming closely
adjacent brushes perpendicular to surface. Megascleres long, smooth oxeas, strongyles, or
more rarely styles, often sinuous, and centrally flexed or straight acanthoxeas. Microscleres
raphides, occurring singly or in bundles (trichodragmata) (modified from Bergquist 1965).
Remarks
Bergquist (1965) meticulously redescribed the type species from type and recent material,
and showed conclusively that the type species of both Acanthoxifera and Neoprosypa were
junior synonyms of M. granulata, despite their apparent differences according to their
published descriptions. Bergquist (1965) also noted that M, granulata had a wide geographical
distribution with a corresponding high degree of skeletal variability, particularly in the
presence, absence and size of certain spicule categories. The genus is similar to Anacanthaea
and Heteroxya, and also apparently related to Higginsia.
Van Soest et al. (1990) revised the higher systematics of Myrmekioderma, placing it in
a redefined order Halichondrida and family Halichondriidae, and suggested that it was most
closely associated with a genus-group also containing Didiscus (based on a synapomorphy
of one or more categories of trichodragmata, the larger sinuously curved). They suggested
that within this group, the smaller oxea 'microscleres' varied from an acanthose to a
completely smooth condition, and thus had a dubious systematic value. In contrast,
Higginsia was retained in Desmoxyidae by van Soest et al. (1990), implying a more distant
relationship with M~rmekiodermathan previously recognised, but this opinion was not
supported by a chemotaxonomic study (Hooper et al. 1992). Consequently, Myrmekioderma
and Higginsia are retained here in the same family.
Van Soest et al. (1990) included six species in the genus, of which only two live in
the Indo-west Pacific: M. dendyi (Burton, 1959) from the south Arabian coast and
Indonesia, M. granulata widespread throughout the Indo-west Pacific (Figs 35-36), M. rea
de Laubenfels, 1934, from the vicinity of Puerto Rico, M. spelea (Pulitzer-Finali, 1983)
from the Mediterranean, M. styx de Laubenfels, 1953, from the Gulf of Mexico, and
M. tulearensis (Vacelet et al., 1976) from south-west Madagascar.
New Caledonian Axinellids
Myrmekioderma granulata (Esper)
(Figs 35-36)
Alcyonium granulatum Esper, 1830: 71, pl. 24.
Myrmekioderma granu1ata.-Ehlers, 1870: 28; Burton, 1938: 39, pl. 7, fig. 42; de Laubenfels, 1954:
121, fig. 75; Lbvi, 1961: 14, fig. 17; Bergquist, 1965: 177, fig. 27a, b; van Soest et al., 1990: 29,
fig. 28; Hooper et al., 1992: 265.
Acanthoxifer ceylonensis Dendy, 1905: 157, pl. 9, fig. 5. -Dendy, 1922: 129.
Myrmekioderma tylota de Laubenfels, 1954: 119, fig. 74.
Neoprosypa atina de Laubenfels, 1954: 190, fig. 127.
Acanthoxifer fourmanoiri LCvi, 1956: 5.
Material Examined
New Caledonian region material. QM G300022 (ORSTOM R1347): stn 427, Caye de I'Observatoire,
Chesterfield Is, 21°24.6'S.,158050.3'E., 28 m depth, 26.vii. 1984, coll. G. Bargibant, SCUBA.
Comparative material. Darwin region, N.T.: NTM 2196: Dudley Pt, East Pt, 12°25'S.,130049.01'E.,
0-0.5 m depth, 13.ix.1981, coll. J. N. A. Hooper, by hand; NTM 22053: same locality, 12'25.01S.,
130°48.4'E., 6-10 m depth, 10.v.1984, SCUBA, coll. J. N. A. Hooper; NTM 2214: Lee Pt, 12O19.02'S.,
130°53.01'E., 0 m depth, 14.xi.1981, coll. J. N. A. Hooper, by hand; NTM 2430: same locality,
13.xii.1981. Arafura Sea, N.T.: NCI Q66C-0540-A (fragment NTM 23097): Parry Shoals, 1l011.411S.,
129"43.011E., 18 m depth, 14.viii.1987, coll. A.-M. Mussig and NCI, SCUBA. Cobourg Peninsula,
N.T.: NTM 22508: NW. Table Head, Port Essington, 11°13~5'S.,132010~5'E.,
3-5 m depth, 14.ix.
1985, SCUBA, coll. J. N. A. Hooper; NTM 23249: SW. Table Head, Port Essington, 11°13.5'S.,
132"105'E., 2-5 m depth, ll.ix.1986, coll. J. N. A. Hooper and C. Johnson, C., SCUBA. Wessel
Islands, N.T.: NCI Q66C-4769-Y (fragment QM (3300741): Bay N. side of Cumberland Strait,
11°28.0'S.,131029.0'E., 20 m, 14.xi.1990, coll. J. N. A. Hooper and NCI, SCUBA. Northwest Shelf,
W.A.: NTM 23423: 150 m W. of Enderbry I., Dampier Archipelago, Dampier, 20°35.8'S.,1 16'28 .OIE.,
14 m depth, 31.viii.1988, coll. Low Choy, D. and NCI, SCUBA; NCI Q66C-1447-0 (fragment NTM
23332): S. reef 500 m from Direction I. Natl Pk, 21°33~1'S.,1150071'E., 6 m, 24.viii.1988, coll.
Low Choy, D. and NCI, SCUBA. Sahul Shelf, W.A.: QM G301141: Hibernia Reef, entrance to
lagoon, NE. side reef, 11°57.8'S.,123022.3'E., 23 m depth, 10.v.1992, coll. J. N. A. Hooper, SCUBA;
QM G301104: Cartier I., outer reef slope, N. side of reef, Australia, 12"31.4'S.,l23"33.3'E., 22 m
depth, 7.v.1992, coll. J. N. A. Hooper, SCUBA.
Description
Colour. Light orange-brown to bright orange exterior alive (Munsell 7.5YR 7/10-5YR
6/10), often with silt covered 'dusty' surface, orange-brown exterior and beige interior in
ethanol.
Shape. Massive, subcylindrical, vaguely elongate, rounded, bulbous growth form,
partially burrowing soft sediments or excavating hard sediments, up to 350 mm long,
200 mm wide, 160 mm thick (although on the Sahul Shelf specimens several metres in
diameter were observed).
Surface. Pineapple-like, convoluted, crustose surface, with large conules or rounded or
polygonal plates, 18-35 mm diameter, slightly raised above surface and separated by shallow
but disctinct grooves; apex of sponge with irregularly meandering or discrete series of
relatively deep, excavated channels (sieve-plates or porocalyces), up to 60 mm deep,
containing large oscules (up to 50 mm diameter) especially near apex of sponge, each oscule
with a raised membraneous lip. Exterior surface invariably silt covered, interior soft,
mango-like. Texture harsh, firm, spiculose.
Ectosome. Distinct, thick, detachable, paratangential crust of smaller (ectosomal)
oxeas, 400-850 pm wide, with innermost layer nearly horizontal and outermost layer nearly
perpendicular to the surface, together forming a continuous palisade of spicules. Ectosomal
crust supported by long, pillar-like tracts of large oxeas, usually widely spaced (450780 pm long, 750-900 pm apart), producing an excavated subdermal region containing
large cavities, about 650 pm diameter, with sparse collagen, collagenous fibrils, bundles of
raphides and sparsely scattered smaller oxeas.
1450
J. N. A. Hooper and C . LCvi
Choanosome. Cavernous, reticulate skeletal structure, with differentiated primary and
secondary spongin fibres and spicule tracts; primary, ascending, multispicular fibres (70250 pm diameter) more-or-less regularly spaced, 300-650 pm apart, interconnected by
secondary, transverse or oblique, pauci-, uni- or aspicular fibres (30-80 pm diameter).
Fibres lightly or heavily invested with spongin, depending on their thickness, and cored only
by larger (choanosomal) oxeas. Fibre meshes evenly rectangular, triangular or irregularly
oval, 90-260 pm diameter, containing abundant collagen, collagenous fibrils, many scattered
smaller oxeas, and bundles of raphides. Choanocyte chambers small, oval, 40-70 pm.
Megascleres. Two categories of megascleres of similar morphology, clearly distinguished
only by size and distribution in skeleton; both entirely smooth, relatively large, straight or
slightly curved at centre, rarely asymmetrical, tapering to sharp fusiform points. Smaller
(ectosomal) oxeas: length 319-(535.3)-708, width 4-(9.2)-12. Larger (choanosomal) oxeas:
length 644-(688.1)-782, width 13-(17.3)-22.
Fig. 35. Myrmekioderma granulata (Esper), specimen Q M G300022: A, larger
(choanosomal) oxeas; B, smaller (ectosomal) oxeas; C, bundles of raphides
(trichodragmata); D, section through cavernous peripheral skeleton.
New Caledonian Axinellids
1451
Microscleres. Raphides rarely seen individually, but more commonly occurring as
bundles of hair-like raphides (trichodragmata). Dimensions of bundles up to 140 x 15 pm.
Distribution
Widely distributed throughout the Indo-west Pacific: Madagascar, Aldabra (LCvi 1956,
1961), Seychelles (Dendy 1922; Hooper, unpublished data); Gulf of Manaar (Dendy 1905;
Burton 1938); Indonesia (Esper 1830; Ehlers 1870; van Soest et al. 1990); north-west
Australia (Hooper, unpublished data); central western Pacific: Ponape, Truk, Ebon Atoll
(de Laubenfels 1954), Palau, Ifaluk (Bergquist 1965); Chesterfield Islands and New Caledonia
(present study). Common habitats range from heavily sedimented fringing coral platforms
and coral pools, in sand, silt, beach rock and dead coral rubble substrates, to pristine coral
reef slopes, often in spurs and grooves; sublittoral depths to approximately 20 m.
Fig. 36. Myrmekioderma granulata (Esper): A , specimen in situ (ORSTOM R1347)
(photo J.-L. Menou); B, SEM fibre junction (specimen QM G300022) (magnified
100~)C
; , SEM skeletal structure (magnified 1 0 0 ~ ) D,
; peripheral skeleton
(scale = 500 pm).
J. N. A. Hooper and C. L h i
1452
Remarks
There is no doubt that M. granulata, and probably all of its sibling species, fits poorly
with the present concept of the axinellids, and possibly the hypothesised relationship with
the halichondrids, as proposed by van Soest et al. (1990), may be more appropriate.
However, this relationship between the desmoxyids, axinellids and halichondrids is still very
poorly understood, and indeed none of these taxa have yet been revised or well investigated.
The conservative choice of leaving Myrmekioderma in Desmoxyidae is presently the best
alternative and also supported by chemotaxonomic data (Hooper et a[. 1992).
The New Caledonian specimen (described above) differs from typical morphotypes of
the species in lacking any spination on the smaller (ectosomal) oxeas, and in lacking any
modification to the oxea morphology; in lacking acanthose oxeas the species is not clearly
a desmoxyid. However, in most other comparative material examined (listed above), the
smaller oxeas are clearly acanthose (and also sometimes modified to strongyles, anisoxeas
or styles). Bergquist (1965) provides a comprehensive comparison in spicule dimensions
between all nominal species and regional populations of M. granulata, and a detailed
discussion of the species' morphometric variability and relationships.
Genus Higginsia Higgin
Higginsia Higgin, 1877: 291.-Hallmann,
1916b: 655; Dendy, 1922: 126; Topsent, 1928: 39;
de Laubenfels, 1936: 132; Burton, 1959: 255; Wiedenmayer, 1977: 156 [type species Higginsia
coralloides Higgin, 1877: 291 (Hallmann 1916b: 656), possible junior synonym of Spongia
strigilata Lamarck, 1814: 4501.
Dendropsis Ridley & Dendy, 1886: 483.-Ridley and Dendy, 1887: 191; Hallmann, 1916b: 693;
de Laubenfels, 1936: 132; LCvi, 1973: 606 [type species Dendropsis bidentifera Ridley and Dendy,
1886: 4831.
Desmoxya Hallmann, 1916b: 649.-de Laubenfels, 1936: 132; van Soest et al., 1990: 18 [type species
Higginsia lunata Carter, 1885: 3581.
Diagnosis
Growth forms erect, lamellate, massive, vasiform or lobate; surface conulose, papillose,
often silt covered or membraneous. Skeletal structure ranges from halichondroid with a
partially compressed, reticulate axis, and an irregularly plumo-reticulate extra-axial region
(Higginsia), a compressed axis and a radial, non-plumose extra-axial region (Dendropsis),
to a lax plumose or plumo-reticulate axial and extra-axial region, without axial compression
or regional differentiation of the skeleton (Desmoxya); spongin fibres usually poorly
developed although heavy collagen forms mesohyl, usually with numerous megascleres and
microscleres scattered between main skeletal tracts; all skeletal tracts cored by monactinal
and/or diactinal megascleres. Ectosome without specialised spiculation, but with extra-axial
spicule tracts (1 or 2 categories of megascleres) protruding through surface. Megascleres
oxeas, strongyles and/or styles of 1-3 sizes. Microscleres include spined, centrangulate
curved or straight microxeas, and sometimes also raphides occurring singly or in bundles
(trichodragmata) (modified from Wiedenmayer 1977).
Remarks
The nominal genera Higginsia, Dendropsis and Desmoxya differ essentially in skeletal
construction (reticulate or plumo-reticulate; with a compressed axis and radial extra-axis;
and plumose-halichondroid, with meandering, occasionally reticulate skeletal tracts respectively).
Most species lack definite axial compression of the skeleton (except Dendropsis bidentifera),
having instead a halichondroid, vaguely reticulate axis, and in most species there is often
some differentiation of axial and extra-axial skeletons (except Desmoxya lunata), suggesting
some sort of affinity with the concept of Axinellida. All three nominal genera share the
apomorphy of spined microxeas, but we are not completely convinced that the major
differences in their skeletal structures can be ignored, particularly in the case of Dendropsis
which has a nearly 'classical axinellid' architecture, in recognising these taxa merely as
synonyms of Higginsia (e.g. Hallmann 1916b; van Soest et al. 1990).
New Caledonian Axinellids
1453
It is possible that resurrection of Desmoxya is required to accommodate Higginsialike species that lack any evidence of axial compression (as seen in both H. lunata and
H. anfractuosa, sp. nov.), having instead a simply halichondroid, meandering Rhaphoxyalike reticulation of choanosomal tracts. These species also differ from Higginsia sensu strict0
only in having one size category of megascleres, having raphides in addition to spined
microxeas, as well as the more lax skeletal architecture. However, intermediate forms of
architecture between Higginsia and Desmoxya are present in some species of Higginsia
(e.g. H. massalis), and this condition is interpreted here as being merely a reduced form of
the skeleton that is typical for Higginsia (van Soest et al. 1990).
Hallmann (1916b: 655-9) provides a comprehensive review of the genus and many of the
species it contained at that time, but many more species have since been included in the
Fig. 37. Higginsia spp.: A , Higginsia lunata Carter (holotype BMNH 1886.12.15.138); B, peripheral
skeleton from BMNH slide (scale = 500 pm); C, Dendropsis bidentlfera Ridley and Dendy (holotype
BMNH 1887.5.2.59); D , skeleton (scale=500 pm); E, Higginsia coralloides var. rnassalis Carter
(holotype of variety, BMNH 1886.12.15.122); F, skeleton (scale=500 pm); G, Higginsia coralloides
(sensu Carter 1885) ('representative specimen' BMNH 1886.12.15.79); H, skeleton (scale=500 pm).
1454
J. N. A. Hooper and C. Ltvi
genus, and all are In need of detailed revision. Few type specimens have yet been located
and seen, so it is not possible to undertake a review of these species at the present time.
Species thought to be valid are: H. bidentifera (Ridley & Dendy) from the Cape of Good
Hope, South Africa (holotype BMNH 1887.5.2.59; Fig. 37C-D); H. coralloides Higgin from
the Caribbean (holotype BMNH [not seen]), with varieties H. c. liberiensis Higgin, 1877,
and H. c. arcuata Higgin, 1877 [possible synonym of H. strigilata (Lamarck); Wiedenmayer
19771; H. higgini Dendy, 1922, from the western Indian Ocean (Okharnandal, Diego Garcia,
Providence, Egmont Reef, and south Arabian coast; Dendy 1922; Burton 1959) (holotype
BMNH [not seen]); H. lunata Carter from Port Phillip Heads (Dendy 1897a; Hallmann
1916b) (holotype BMNH 1886.12.15.138, schizotype MNHN LBIM DCL283; Fig. 37A-B);
H. massalis Carter, 1885, from Port Phillip Heads, Vic., and Ambon, Indonesia (Dendy
1897a; Topsent 1897; Hallmann 1916b) (holotype BMNH 1886.12.15.122; Fig. 37E-F), with
Fig. 38. Higginsia spp.: A , Higginsia mixta Hentschel ('representative specimen'
NTM 22236); B, skeleton (scale = 500 pm); C, Higginsia scabra Whitelegge (specimen
on deck NCI Q66C-0523-5) (photo NCI); D, skeleton of specimen NTM 22801
(scale = 500 pm); E, Spongia strigilata Lamarck (holotype MNHN DT637); F,
skeleton (scale = 500 pm).
New Caledonian Axinellids
1455
synonym 'H. coralloides7o f Carter (1885)from Port Phillip Heads ('representativespecimen'
BMNH 1886.12.15.79; Fig. 37G-H), and 'H. strigilata' o f Desqueyroux-Faundez (1981);
H. mixta Hentschel, 1912, from southern Indonesia (holotype SMF 968 [not seen]) and
Palau (Bergquist 1965), also known from north-west and north-east Australia (Hooper,
unpublished data; 'representative specimen' NTM 22236; Fig. 38A-B); H. natalensis
Carter, 1885, from the Cape o f Good Hope, South Africa (holotype BMNH [not seen]);
H. papillosa Thiele, 1905, from Calbuco, Chile (holotype probably ZMB [not seen]);
H. petrosioides Dendy, 1922, from the Seychelles and Indonesia (holotype BMNH [not
seen]); H. pumila (Keller, 1889) from the Red Sea (holotype ZMB 442 [not seen]);
H. robusta Burton, 1959, from the Gulf o f Aden (holotype BMNH 1936.3.4.342 [not
seen]);H. scabra Whitelegge, 1907, from Port Jackson, N.S.W. (Hallmann 1916b), and
north-west and north-east Australia (Hooper, unpublished data) (holotype AM 2480;
'representative specimen NCI Q66C-0523-J; Fig. 38C-D); H. strigilata (Lamarck) from an
uncertain locality (Turgot collection), possibly originating from the West Indies (holotype
MNHN DT637; Fig. 38E-F) (Topsent 1932; Wiedenmayer 1977); and H. thielei Topsent,
1904, from the Azores (holotype possibly Monaco).
O f these species, only five have been recorded from the Indo-west Pacific: H. lunata,
H. massalis, H. mixta, H. petrosioides and H. scabra.
Higginsia anfractuosa, sp. nov.
(Figs 39-40, Table 8 )
Material Examined
Holotype. QM G300723 (ORSTOM 'cfR806'): stn 181, E. reef flat, Il6t Maitre, New Caledonia
lagoon, 22°20.1'S.,166025.0'E., 1.5 m depth, 2.vi.1977, coll. G. Bargibant, by hand.
Description
Colour. Pale orange alive (Munsell 7.5YR 8/8), olive-brown in ethanol.
Shape. Erect, globular, cylindrical digit, tapering towards base and apex, 62 mm long,
24 mm diameter at base, 32 mm widest diameter, attached directly to substrate (with
embedded detritus in basal end), without stalk or other processes.
Surface. Evenly distributed, rounded Cliona-like papillae, up to 2.5 mm diameter, only
raised slightly above surface, each with a terminal apical oscule (now closed), 1.0-1.5 mm
Table 8. Comparison in spicule dimensions between Higginsia anfractuosa, sp. nov., and H. lunata
All measurements given in micrometres, and expressed as minimum-(mean)-maximum range of
measurement. N = 2 5 for each specimen
Spicules
Species
H. anfractuosa, sp. nov.
(holotype QM G300723)
Megascleres
Microscles
Acanthoxeas
Raphides I
Raphides I1
L
W
L
W
L
W
L
W
H. lunata Carter
(holotype
BMNH1886.12.15.138)
238-(298.2)-318
2.5-(3.3)-4.5
(Vestigial, predominantly
strongyles)
(Robust, predominantly
styles, fewer strongyles)
106-(129.3)-173
2.5-(2.9)-3.5
(Straight, evenly spined)
176-(252.9)-286
1.5-(2.1)-2.5
91-(99.8)-112
0.5-(1 .I)-2.0
30-(38.1)-48
2.0-(2.8)-4.0
(Curved, terminally spined)
448-(540.8)-620
0.5-(1.3)-2.0
60-(132.3)-210
0.5-(1.3)-2
1456
J . N. A. Hooper and C. L h i
diameter. Surface with distinct (non-detachable) dermis, more darkly pigmented than
choanosomal region, and has overall goose-flesh appearance. Texture rubbery, compressible.
Ectosome. Heavy collagenous ectosomal layer, 100-250 pm wide, darkly pigmented,
containing scattered megascleres and microscleres, predominantly paratangential to the
surface. Around oscules spicules ordered into diverging rays, presumably supporting surface
papillae and providing support for oscule contractile mechanism (Fig. 40C).
Choanosome. Skeleton plumose-halichondroid, with predominantly ascending skeletal
tracts, meandering and rejoining throughout skeleton; fibre meshes irregular, elliptical-oval,
more cavernous in interior than peripheral skeleton; no axial compression and no differentiation between axial and extra-axial skeletons; spongin fibres only lightly invested with
spongin, 30-90 pm diameter, with poorly differentiated primary and secondary elements;
fibres fully cored with megascleres, dispersed in plumose-diverging tracts. Mesohyl contains
heavy collagen and abundant loose megascleres and rnicroscleres. Choanocyte chambers
20-85 pm diameter.
Megascleres (refer to Table 8 for dimensions). Vestigial megascleres, predominantly
strongyles, occasionally styloid or strongyloxeas, straight, slender, with symmetrical, rounded
ends, or slightly tapering hastate points.
Microscleres (refer to Table 8 for dimensions). Acanthoxeas long, slender usually straight,
occasionally asymmetrical, evenly spined, granular spination, with tapering, sharply pointed
ends.
Two size categories of raphides present, both straight, thin, tapering to sharp fusiform
points, the larger category nearly the same length as megascleres, differentiated only by
Fig. 39. A-D, Higginsia anfractuosa, sp. nov., holotype QM G300723: A , vestigial
strongyles, strongyloxeas and styloids; B, two sizes of raphide microscleres; C,
acanthoxeas; D, section through peripheral skeleton. E, Higginsia lunata Carter,
holotype B M N H 1886.12.15.138, curved acanthoxeas.
New Caledonian Axinellids
Fig. 40. Higginsia anfractuosa, sp. nov.: A , holotype (QM G300723); B, peripheral skeleton
(scale = 500 am); C, oscule and supporting spicules radiating around oscule (scale = 200 pm);
D, SEM fibre characteristics (scale = 200 pm).
their terminations and relative thickness. Raphides dispersed individually in skeleton, no
trichodragmata observed.
Distribution
Known only from the type locality, coral rubble.
Remarks
This species is a closely related, sibling species of Higginsia lunata Carter (Carter 1885;
Dendy 1897a) from Port Phillip, Vic., meticulously redescribed by Hallmann (1916b: 650).
The two species differ substantially only in the size of megascleres and microscleres
(Table 8), and geometry of acanthoxeas (curved, terminally spined, slightly centrangulate in
H. lunata; straight, evenly spined, not swollen at centre in the present species; Fig. 40C, E ) .
Growth form, papillose surface features and skeletal structure are comparable between
the two species, and both are atypical of all other Higginsia in having a lax, plumosehalichondroid, meandering skeleton. Nevertheless, these differences between the New
Caledonian and Port Phillip populations are greater than normally attributed to interspecific
variability, substantially more than cryptic differences, and the New Caledonian population
is considered to represent a separate species.
Etymology
Named for the winding, meandering skeletal columns; from anfractuosus (L.), very
winding, sinuous.
Higginsia tanekea, sp. nov.
(Figs 41-42, Table 9)
Material Examined
Holotype. QM G300024 (ORSTOM R1298): stn 305, N. Ile Paaba, New Caledonia lagoon,
19"55.3'S.,161°37.3'E., 27 m depth, 24.vi.1981, coll. G . Bargibant, SCUBA.
1458
J. N. A. Hooper and C. Lkvi
Description
Colour. Pale orange alive (Munsell 7.5YR 7/10), beige in ethanol.
Shape. Massive, irregularly bulbous, subspherical, subcylindrical sponge, 210 mm long,
80 mm wide, 55 mm thick, without stalk or other processes, loosely attached directly to
substrate, with embedded detritus on 'ventral surface', or unattached and rolling free on
substrate ('tumbleweed' sponge).
Surface. Slightly bulbous surface, with low, rounded ridges, distinct skin-like, detachable
dermis and irregularly dispersed microconules, up to 2 mm diameter, conical or elongate and
irregular in shape, not raised more than 2 mm from surface, interconnected by shallow
canals and grooves. Surface smooth, not hispid. Texture soft, compressible, relatively fragile,
easily torn. Internal consistency porous, cavernous, Echinoclathria-like. Oscules not observed.
Ectosome. Collagenous detachable dermis, 100-350 bm wide, containing darkly pigmented collagen, sparsely dispersed thinner ectosomal oxeas forming paratangential tracts,
interdispersed with crust of acanthoxeas mostly erect on surface, mostly confined to
peripheral skeleton. Choanosomal megascleres not protruding beyond surface. Subdermal
region cavernous, directly below skin-like collagenous layer, with sparse tracts of choanosoma1 megascleres supporting dermal layer.
Choanosome. Skeleton halichondroid-reticulate, with vaguely ascending spongin fibres
and skeletal tracts forming wide-meshed reticulation. Spongin fibres light, more-or-less
Fig. 41. Higginsia tanekea, sp. nov., holotype QM G300024: A , choanosomal oxeas and styloids;
B, ectosomal oxeas; C, acanthoxeas; D, section through peripheral skeleton.
New Caledonian Axinellids
1459
divided into primary, ascending, multispicular fibres, 60-160 pm diameter, and shorter,
interconnecting, predominantly transverse, uni- or paucispicular, secondary fibres, 30-75 pm
diameter; fibre reticulation forming cavernous, oval or elongate meshes, 350-1200 pm
diameter, wider in peripheral skeleton than in deeper regions of choanosome; all fibres
cored by larger oxeas, forming irregular, slightly confused, sometimes meandering tracts,
with spicules usually occupying most of fibre diameter; numerous larger choanosomal and
Fig. 42. Higginsia tanekea, sp. nov.: A , holotype [QM G3000241, peripheral
skeleton (scale=500 pm); B, specimen in situ (ORSTOM 'cfR1298') (photo
P. Laboute); C, holotype in situ (ORSTOM R1298, on right) (photo P. Laboute);
D, SEM skeletal structure (scale =200 ~ m ) .
1460
J. N. A. Hooper and C. LCvi
smaller ectosomal oxeas also dispersed between fibre skeleton, but few acanthoxeas seen in
choanosome. Fibre meshes contain abundant collagen, more lightly pigmented than in
peripheral skeleton, and small oval or eliptical choanocyte chambers, 25-70 ym diameter in
peripheral region.
Megascleres (refer to Table 9 for dimensions). Choanosomal megascleres predominantly
oxeas, rarely styloid, usually long, slender, symmetrically curved, with fusiform, sharply
pointed ends.
Ectosomal oxeas with same morphology, slightly shorter and substantially thinner.
Microscleres (refer to Table 9 for dimensions). Acanthoxeas variable in length, mostly
relatively long, slender, usually with slight, angular, central curvature, occasionally straight
or with acute bend, sharply pointed tips, with evenly dispersed, large spines.
Distribution
Known only from the New Caledonia lagoon, in Halimeda beds in the inter-reef region,
27 m depth.
Remarks
The relationship of Higginsia tanekea to other members of the genus is uncertain.
It is not obviously closely related to any other species in the combination of its morphological characters (skeletal structure, spicule size or spicule geometry), although individually
most of these features can be found in at least one other species of the genus. In fact, the
only apomorphic character in H. tanekea appears to be the possession of unusually large,
perpendicular spines on acanthoxeas; a plesiomorphic feature is the lack of long, extra-axial
styles which are present in most other species. Higginsia tanekea shows some similarities,
in some of its features, with several other species, but these can be differentiated as follows:
H, scabra has long slender extra-axial styles, lacks smaller, slender ectosomal oxeas, and
has an axinellid skeleton with distinctive plumo-reticulate, diverging architecture, and
differentiated axial and extra-axial regions (depicted by Hallmann 1916b: pl. 41, figs 1-3);
H. massalis has a similar skeletal architecture and has similarities in the geometry of some
of its spicules, but it has shorter, more robust choanosomal oxeas (often asymmetrical and
with telescoped points), long extra-axial styles, and shorter acanthoxeas; H. mixta has much
thicker choanosomal oxeas (some styloid, some with telescoped points), also with long,
slender extra-axial styles, longer ectosomal oxeas, and has an axinellid skeletal structure;
H. robusta has much thicker choanosomal spicules (varying from styles to oxeas), long,
slender extra-axial styles are present, acanthoxeas substantially shorter, and it too has an
axinellid skeleton; H. pumila is poorly known but, from Keller's (1891) description, it
differs in having choanosomal spicules consisting of thick styles, and the extra-axial spicules
are long, slender styles. Spicule dimensions of these species are compared in Table 9.
Etymology
Named for the long, slender, sharp spines on acanthoxea microscleres; tanekes (Gk),
long-pointed.
Higginsia massalis Carter
(Figs 43-44, Table 9)
Higginsia coralloides.-Carter, 1885: 357.
Not Higginsia coralloides Higgin, 1877: 291, pl. 14, figs 1-5.
Higginsia coralloides Higgin var. massalis Carter, 1885: 357.-Dendy, 1896: 243; Hallmann, 1916b:
656, 659-65, pl. 29, fig. 6, pl. 38, figs 6-7, pl. 39, figs 1-2, pl. 40, figs 1-4.
Material Examined
Holotype. BMNH 1886.12.15.122: Port Phillip Heads, Vic., coll. J. B. Wilson.
New Caledonian material. QM G300023 (ORSTOM R1222): stn 124, Ili3t Maitre, 22"20.11S.,
166"25.11E.,20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM unregistered: Fosse aux
New Caledonian Axinellids
1461
Canards, 22"19.2'S.,l66"26'E., 20-25 m depth, date of collection unknown, coll. P. Laboute, SCUBA;
QM G301321: Croisant-Larkgritre, IlBt Maitre, off Noumea, 22°20.2'S.,166022 S'E., 20 m depth,
13.x.1992, coll. J. N. A. Hooper, SCUBA.
Comparative material. BMNH 1886.12.15.79: Port Phillip Head, Vic., 22 m depth, coll. J. B.
Wilson.
Description
Colour. Pale orange alive (Munsell 5YR 8/6), grey-brown in ethanol.
Shape. Massive, elongate, irregularly subspherical, 73 mm long, 46 mm diameter,
without stalk or other processes, attached directly but loosely to the substrate with embedded
detritus in 'ventral surface', or rolling freely on the substrate ('tumbleweed' sponge).
Table 9. Comparison in spicule dimensions between Higginsia tanekea, sp. nov., and related
species
All measurements given in micrometres, and expressed as range of measurement
Spicule
Species (locality; source of data)
Choanosomal
Extra-axial
Ectosomal
megascleres
megascleres
megascleres
Higginsia tanekea, sp. nov. (New Caledonia; holotype QM G300024)
L
628-(824.3)-993
Absent
392-(512.9)-622
W
4-(10.2)-14
3-(4.3-7
(Slender oxeas,
(Oxeas)
rarely styloid)
H. mixta (Hentschel) (Aru I.; Hentschel 1912)
775-1175
L
624-744
2240
4-5
W
28-3 1
20-3 1
(Robust oxeas)
(Long styles)
(Oxeas)
H. mixta (Hentschel) (Palau; Bergquist 1965)
650-912
L
1025-1150
1900-3125
5-7
W
16-21
14-18
H. mixta (Hentschel) (NW. Australia; unpublished data, NTM collection)
L
570-(871.3)-1250
850-(1260.0)-2015
Absent
W
25-(38.6)-50
4-(9.8)-16
H. scabra (Port Jackson, N.S.W.; Hallmann 1912)
L
550-770
950-1 100
Absent
W
8-35
15-25
(Robust oxeas)
(Long styles)
H. scabra Whitelegge (NW. Australia; unpublished data, NTM collection)
L
550-(608.3)-675
222-(327.3)-417
Absent
W
18-(33.4)-42
5-(7.3)-11
H. massalis Carter (Port Phillip, Vic.; holotype BMNH 1886.12.15.122)
321-(568.4)-704
492-(1129.8)-1750
L
535-(599.8)-706
2-(6.4)-11
W
10-(13.4)-17
5-(9.0)-14
(Robust oxeas,
(Long styles,
(Oxeas, occasionless often
rarely strongyles
ally bidentate)
styloid)
or oxeas)
H. massalis Carter (Port Phillip, Vic.; specimen BMNH 1886.12.15.79)
265-(468.8)-669
L
464-(610.1)-692
412-(867.0)-1221
4-(6.4)-9
7-(10.6)-15
W
8-(12.8)-16
H. massalis Carter (Port Phillip, Vic. ; Hallmann 1916b)
Up to 900
200-400
L
560-700
4-5
9
W
14-18
H. massalis Carter (New Caledonia; specimen QM G300023)
512-(716.8)-843
L
841-(898.8)-936
632-(1484'3)-2121
2-(5. 1)-8
6-(8.2)-10
W
12-(14.6)-18
(Robust oxeas,
(Long styles,
(Oxeas)
never styloid)
never oxeas)
Acanthoxea
microscleres
71-(111 .@-I43
1.5-(2.9)-4.5
88-152
3-5
62-200
2-5
70-(108.3)-175
3-(6.1)-10
60-130
Up to 5
72-(110.0)-154
1-(4.1)-6
72-(97'8)-124
2-(3.7)-5
71-(84.9-96
2.5-(3.8)-5
40- 130
4-5
74-(96.4-137
2-(3.1)-4.5
J. N. A. Hooper and C. Livi
1462
Surface. Uneven, irregular, lumpy surface, with a distinct skin-like, detachable dermis,
covered with evenly dispersed, irregularly shaped microconules, up to 3 mm diameter, raised
no more than 2 mm from surface, forming meandering, crenellated ridges and valleys.
Oscules, 3-6 mm diameter, slightly raised above surface, with membraneous lip; smaller
ostia, less than 1 mm diameter, situated between ridges. Texture soft, compressible, easily
torn; internal consistency compact, only slightly cavernous, spiculose, friable, with low
spongin fibre component.
Ectosome. Minutely hispid surface produced by sparsely dispersed, long extra-axial
styles, protruding through a highly collagenous, darkly pigmented dermal layer, 70-220 pm
wide. Ectosomal layer contains sparse, paratangential tracts of smaller, thinner oxeas
and a thick, paratangential crust of acanthoxeas; acanthoxeas mainly confined to dermal
skeleton. Subdermal region slightly cavernous, with elongate canals, 350-900 pm diameter.
Fig. 43. Higginsia rnassalis Carter, specimen Q M G300023: A , choanosomal oxeas;
B, extra-axial styles; C, 'ectosomal' oxeas; D, acanthoxeas; E, section through peripheral
skeleton.
New Caledonian Axinellids
1463
Choanosome. Skeleton plumo-reticulate, verging on disorganised-halichondroid, without
well developed axial compression of skeleton, with only poorly differentiated axial and
extra-axial regions, with extra-axial spicule tracts slightly more plumose than more reticulate
choanosomal spicule tracts, and containing long extra-axial styles in peripheral region,
usually perpendicular to surface; skeleton consists of differentiated primary, more-or-less
ascending, multispicular tracts, 70-110 pm diameter, interconnected by shorter, uni- or
paucispicular, secondary, mostly transverse tracts, 20-70 pm diameter, cored exclusively
by long choanosomal oxeas, with some thinner 'ectosomal' oxeas interdispersed; fibre
reticulation produces elongate meshes, 110-300 pm diameter, but often partially obscured
by choanosomal spicules dispersed between major skeletal tracts; spongin fibre system poorly
developed, spicules appear to be cemented together primarily by granular collagen; meshes
contain abundant collagen, with numerous choanosomal oxeas, only few acanthoxea
microscleres, and very small quantities of detritus (arenaceous); choanocyte chambers small,
oval to elongate, 25-45 pm diameter.
Fig. 44. Higginsia massalis Carter: A, specimen in situ [ORSTOM R1222 (specimen
QM G300023)l (photo P. Laboute); B, peripheral skeleton (scale=500 pm); C,
same specimen preserved; D, SEM peripheral skeleton (scale = 500 pm).
1464
J. N. A. Hooper and C . Ltvi
Megascleres (refer to Table 9 for dimensions). Choanosomal megascleres exclusively
oxeas, robust, relatively long, straight or slightly curved at centre, usually symmetrical, with
fusiform, tapering, sharp, rarely tel~scopedpoints.
Extra-axial styles variable in length, usually very long, slender, slightly curved, sometimes straight or sinuous, with evenly rounded bases, tapering to sharp or slightly
telescoped points.
'Ectosomal' oxeas very long, slender, usually slightly curved, sometimes greatly curved or
sinuous, sharply pointed; these spicules not confined exclusively to ectosomal region, but also
found in smaller quantities in choanosomal skeletal tracts.
Microscleres (refer to Table 9 for dimensions). Acanthoxeas relatively long, slender,
with slight angular curvature at centre, tapering to sharp points, evenly covered with small
spines; spines possibly larger at centre of spicule than on ends of spicule.
Distribution
Southern Australia and New Caledonia lagoon, inter-reef region on sand and coral rubble
substrate, 10-22 m depth.
Remarks
There are several differences between the New Caledonian specimen of H. massalis,
described above, and the holotype from southern Australia. Apart from minor differences
in spicule sizes (Table 9), the points of choanosomal oxeas are often telescoped in the
holotype, whereas they are rarely so in the present material, being more commonly sharply
pointed; ectosomal oxeas in the holotype often have slightly rounded or bidentate terminations, whereas those in the New Caledonian specimen appear to be exclusively sharply
pointed; extra-axial styles in the present material are always styloid, whereas those in the
holotype are occasionally oxeote; and skeletal architecture is a little more disorganised in the
New Caledonian specimen, whereas it is more regularly plumo-reticulate in the holotype.
According to Hallmann (1916b: 659), the skeletons of several non-typical specimens from
the Port Phillip region are composed of a series of thin lamellae, separated at the periphery
but united further towards the choanosome, producing a shaggy peripheral skeletal structure
(or 'skeleton-sponge' of Hallmann 1916b). This latter feature was not obvious in the New
Caledonian specimen, nor was it apparent in the holotype.
These apparently minor differences may eventually prove to be indicative of two cryptic
sibling species, possibly justifying the erection of a new taxon for the New Caledonian
population. However, examination of a second specimen from Port Phillip Heads, originally
assigned to the West Indies species H. coralloides by Carter (1885), showed it to be virtually
intermediate in most of these characters, making it nearly impossible to clearly delineate the
populations into separate species (without other, non-morphological characters to support
such a proposal).
Hallmann's (1916b) measurements of spicules in his material differ slightly from those
observed in both the holotype and Carter's (1885) specimen, especially the size of ectosomal
spicules, and the corrected measurements for this material are presented in Table 9. Species
related to H. massalis have already been discussed above and are also compared in Table 9.
Discussion
An earlier study of five families of poeciloscleridan sponges from the New Caledonian
region (Hooper and LCvi 1993) listed 28 species, 18 of which were apparently endemic
to this fauna. Although it was noted that there was a greater proportion of indigenous
poecilosclerid sponge species (71 %) living exclusively in deeper-waters of the shelf and slope
of New Caledonia, agreeing with the preliminary biogeographic model suggested by LCvi
(1979), the level of endemism for the shallow-water poecilosclerid fauna in the lagoon and
outer reefs was also found to be significantly higher (62%) than previously predicted by
LCvi. The present study described or redescribed 16 axinellid species from the shallow-waters
in the lagoon and outer reefs surrounding New Caledonia, bringing the total number of
New Caledonian Axinellids
1465
known axinellid species in this region to 25, of which 7 are new species and 7 are new
locality records.
The non-endemic, shallow-water axinellid fauna appears to be most closely related to the
north-east Australian (Solanderian) and Indo-Malay fauna, usually representing the easternmost extent of their known distributions in the Indo-west Pacific. The shallow-water fauna
also includes two authenticated 'widely distributed' species: Axinella carteri (Dendy) (distributed throughout coral reefs of the western and eastern Indian Ocean, the Indo-west Pacific
and the western Pacific rim), and Astrosclera willeyana Lister [found in the western Indian
Ocean (RCunion, Madagascar and Mozambique; Vacelet and Vasseur 1965, 1971; Vacelet
1967; Vacelet et al. 1976), eastern Indian Ocean (Christmas I.; Kirkpatrick 1910; Ashmore,
Cartier and Hibernia Reefs, Sahul Shelf; Hooper 1992; southern Indonesia; van Soest 1990),
central western Pacific Ocean (Guam; Hartman in Vacelet 1967), south-western Pacific
Ocean (Great Barrier Reef; Ayling 1982; Loyalty and Tuvalu Is; Lister 1900; New Caledonia;
Vacelet 1981), and central southern Pacific (French Polynesia; Vacelet 1977)], whereas other
previously suspected 'widely distributed' species were generally found to be separate, closely
related, cryptic sibling species. By comparison, the previous study on the poecilosclerid fauna
indicated that its affinities lay mainly with both the northern and southern Australian
shallow-water fauna, with surprisingly fewer similarities with the northern New Zealand
fauna, plus a relatively higher endemic element (Hooper and LCvi 1993). Similar to the
Poecilosclerida, however, many of the new axinellids described here were recognisable as
sister-species of known, mainly tropical Australasian axinellids. These observed cryptic
differences between many of the New Caledonian and Australasian sister-species are considered to be real, representing specific fixed differences in the genotype, rather than
phenotypic variability of single, widely distributed species. This point of view contrasts with
earlier studies in the western Pacific region (e.g. Burton 1934), and generally has empirical
support (biochemical, genetic and detailed morphometric analyses of both sympatric and
allopatric species; exhaustive comparison of relevant voucher material cited in these earlier
studies, with less reliance on the published literature; Hooper et al. 1990, 1992; van Soest
et al. 1991; van Soest and Hooper 1993).
Of the four families and 25 species of axinellids known to live in the vicinity of New
Caledonia, 12 have not been recorded elsewhere (48% endemism). Most of this endemism
can be accounted for by the deeper water fauna (three of five species), compared with only
nine of the 20 species being shallow-water species. The proportion of endemic species for
all four axinellid families is significantly lower than equivalent figures obtained for five
families of Poecilosclerida (71% deeper-water, 63% shallow-water; Hooper and LCvi 1993),
or for the entire described sponge fauna in this region (72% deeper-water, and 40% shallowwater). However, considering only the family Axinellidae so far collected and described for
the region, the proportion of endemic species is high for both the shallow- and deeper-water
species. Further comments on the biogeographic affinities of the New Caledonian Axinellidae
will be discussed in a separate contribution.
A summary of the New Caledonian axinellid fauna is presented below [S, predominantly
or exclusively shallow-water species (0-100 m depth); D, predominantly or exclusively deeperwater species (100-500 m depth); species marked with an asterisk are thought to be endemic
to the region].
Axinellidae (15 species have been described for this region, nine of which are apparently
endemic):
Cymbastela cantharella (LCvi, 1983) [S]*
C. concentrica (Lendenfeld, 1887) [S]
Reniochalina plumosa (LCvi & LCvi, 1983) [Dl*
R. condylia, sp. nov. [S]*
Axinella lifouensis LCvi & LCvi, 1983 [Dl*
A . carteri (Dendy, 1889) [S]
Phakellia columnata (Burton, 1928) [Dl
P. pulcherrima (Ridley & Dendy, 1886) [S]
P. stipitata (Carter, 1881) [S]
J. N. A. Hooper and C. Ltvi
Stylissa flabelliformis (Hentschel, 1912) [S]
Ptilocaulis fusiformis LCvi, 1967 [S]*
P. epakros, sp. nov. IS]*
P. papillatus, sp. nov. [S]*
Pseudaxinella debitusae, sp . nov. [S]*
Rhaphoxya systremma, sp. nov. IS]*
Desmoxyidae (five species known for the region, only two of which are possibly endemic):
Myrmekioderma granulata (Esper, 1830) [S]
Higginsia anfractuosa, sp. nov. [S]*
H. tanekea, sp. nov. [S]
Higginsia massalis Carter, 1885 [S]
Parahigginsia phakellioides Dendy, 1924 [Dl
Trachycladidae (one species known for the region, also recorded from south-eastern and
south-western Australia):
Trachycladus digitatus Lendenfeld, 1887 [S]
Agelasidae [now in the order Agelasida; two species known for the region, one apparently
endemic, and two hypercalcified 'sclerosponges' are also included here (following van Soest
1984 and others)]:
Agelas mauritiana (Carter, 1883) [S]
A . novaecaledoniae Levi & LCvi, 1983 [Dl*
Astrosclera willeyana Lister, 1900 [S]
Stromatospongia micronesica Hartman and Goreau, 1976 [S]
Acknowledgements
We are extremely grateful to Dr CCcile Debitus, ORSTOM, Noumea, for assisting with
the acquisition of specimens and in situ photographs of live material, which have greatly
facilitated the preparation of this publication. We also gratefully acknowledge funding
provided by both ORSTOM Noumea and DITAC Canberra, which enabled the authors
to participate in a series of workshops at ORSTOM Noumea on the taxonomy of New
Caledonian shallow-water sponges. This publication is one of several recent contributions
on the shallow-water fauna of the Noumea lagoon, as a prelude to the publication of a
forthcoming popular book on the subject, and we acknowledge the assistance and interaction
of our colleagues Chris Battershill, Patricia Bergquist, Jane Fromont, Michelle Kelly-Borges,
Jean Vacelet and Clive Wilkinson. For collection of specimens and photographs we thank
Pierre Laboute, Georges Bargibant, Jean-Louis Menou, and other members of ORSTOM
Noumea for the hospitality, helpful assistance and cooperation during this project.
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