Znvertebr. Taxon., 1993, 7, 1221-302
Poecilosclerida (Porifera :Demospongiae)
from the New Caledonia Lagoon
John N. A . ~ o o ~ e and
r * Claude ~ 6 v i ~
Queensland Museum, P.O. Box 3300, South Brisbane, Qld 4101, Australia.
Laboratoire de Biologie des InvertBbrts marins et Malacologie,
Museum national d'Histoire naturelle, 57, rue Cuvier, 75005 Paris, France.
A
Abstract
Twenty species of shallow-water poecilosclerid demosponges are described from the lagoon and reefs
of New Caledonia, including thirteen new species and six new locality records, belonging to five
families and eight genera {Myxillidae (1, Acarnus), Desmacellidae (1, Neofibularia), Crellidae (1, Crella),
Microcionidae 113, Clathria (Clathriopsamma), C. (Clathria), C. (Thalysias), Echinochalina (Echinochalina), E. (Protophlitaspongia)] and Raspailiidae [4 species, Raspailia (Raspailia), Ceratopsion,
Aulospongus]}. The shallow-water fauna contained a mixture of endemic species (65%) and those with
affinities to both northern and southern Australia, in contrast to the deeper-water fauna which was
previously found to have high endemism (70%) and ancestral affinities to northern New Zealand.
The non-endemic shallow-water fauna represents the easternmost extent of species' distributions in
the Indo-west Pacific, and the endemic component was also usually immediately recognisable as
(transformed) sister-species from Australian tropical and temperate provinces.
Introduction
Only about 170 species of Porifera have been described from the New Caledonian region,
although many hundreds of other species have already been collected by ORSTOM, and the
fauna is thought to consist of about 600 species. Furthermore, most of the published studies
have concentrated on the deeper-water fauna (Demospongiae, including the polyphyletic
Lithistida; LCvi and LCvi 1983a, 1983b, 1988; Uvi 1991), or have dealt exclusively with
Hexactinellida (LCvi and L h i 1982), hypercalcified 'sclerosponges' (Vacelet 1981), or Calcarea
(Vacelet 1981). Prior to the present study, 76 species of deeper-water (> 150 m depth)
Demospongiae had been described from the New Caledonian region, including the Norfolk
Rise, whereas only 68 species were known from shallow-waters (LCvi 1967, 1983; LCvi and
LCvi 1978; Vacelet 1981; Desqueyroux-Faundez 1984, 1987; Hooper and Bergquist 1992).
The present study concerns only the shallow coastal demosponge fauna of New Caledonia,
from lagoon and coral reef habitats surrounding the island. This first paper describes a
number of species of the order Poecilosclerida; a second paper will deal with the order
Axinellida (Hooper and LCvi 1993); the biogeographical relationships of the New Caledonian
shallow-water microcionid and raspailiid fauna, with respect to other provinces within the
Indo-west Pacific, will be discussed in a separate contribution. These papers represent
the results of a collaborative investigation amongst several 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.
1222
J. N. A. Hooper and C. LCvi
Methods
Methods used to prepare and examine sponges for light microscopy and electron microscopy are
described in Hooper (1991b). Spicule measurements are based on examination of 25 random samples
of each spicule category for each specimen, and 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, Natural History Museum, London; CSIRO, Commonwealth
Scientific and Industrial Scientific Organisation, Division of Fisheries, Hobart; DITAC, Commonwealth
of Australia Department of Industry, Technology and Commerce, Canberra; LFM, Merseyside County
Museums, Liverpool; MNHN, Museum National d'Histoire Naturelle, Paris; NCI, US National Cancer
Institute shallow-water collection contract (Australian Institute of Marine Science, Townsville; primary
colle&ions in National Museum of Natural History, Washington, duplicates and fragments in NTM
and QM); NMB, Naturhistorisches Museums zu Basel; NMV, Museum of Victoria, Melbourne; N.S.W.,
New South Wales; N.T., Northern Territory; NTM, Northern Territory Museum of Arts and Sciences,
Darwin; ORSTOM, Institut Francais de Recherche Scientifique pour le Developpement en Cooperation,
Centre de Noumea; QFS, Queensland Fisheries Service; Qld, Queensland; QM, Queensland Museum,
Brisbane; S.A., South Australia; SMF, Natur-Museum und Forschungsinstitut Senckenberg, Frankfurt;
Tas., Tasmania; USNM, United States National Museum, Smithosonian Institution, Washington; Vic.,
Victoria; W.A., Western Australia; ZMA, Institut voor Taxonomische Zoologic, Zoologisch Museum,
Universiteit van Amsterdam; ZMB, Zoologisches Museum fiir Naturkunde an der Humboldt-Universitat
zu Berlin, Berlin; ZMH, Zoologisches Institut und Zoologisches Museum der Universitat Hamburg,
Hamburg.
Systematics
Order POECILOSCLERIDA Topsent
Family MYXILLIDAE Topsent
Definition (emended)
Poecilosclerida with a renieroid, regularly reticulate o r (plumo-) reticulate choanosomal
skeleton, composed of tracts of smooth or acanthose monactinal megascleres (styles, smooth
or spined, rarely oxeote or strongylote), with or without echinating spicules; ectosomal
tangential skeleton composed of diactinal megascleres, typically being smooth tylotes with
microspined bases. Microscleres commonly arcuate isochelae, but may also include anchorate,
unguiferous, birotulate or palmate isochelae, anisochelae, sigmas, bipocilli, forceps, and
toxas (modified from van Soest 1984~).
Remarks
Only two other species of Myxillidae have so far been described for the New Caledonian
region, both from deeper waters (410-525 m depth)-Lissodendoryx bifacialis LCvi and
LCvi, 1983b and L. stylophora LCvi & LCvi, 1983b.
Genus Acarnus Gray
Acarnus Gray, 1867: 544 [type species Acarnus innominatus Gray, 1867: 5441 [full synonymy and
discussion given by Hooper (1987: 72), Hiemstra and Hooper (1991: 433) and van Soest et al.
(1991: 49)].
Diagnosis
Myxillidae with ectosomal tylotes, choanosomal megascleres include styles or subtylostyles,
completely smooth or with basal spines, echinating cladotylotes of 1 o r 2 sizes, with o r
without echinating acanthostyles of 1 o r 2 sizes; microscleres include palmate isochelae and
toxas, the latter with up to 3 categories including a thick, evenly curved form with recurved
ends ('oxhorns') (from Hiemstra and Hooper 1991).
Remarks
All species of Acarnus share the apomorphic feature of cladotylotes echinating fibres,
and some species also have echinating acanthostyles, a symplesiomorphic character for the
Poecilosclerida (also found in the Microcionidae, Raspailiidae, Crellidae and Anchinoidae).
The presence of acanthostyles was previously used t o differentiate the nominal genus
Acanthacarnus LCvi, 1952, from species lacking acanthostyles (i.e. Acarnus s.s.), but the
New Caledonian Poecilosclerids
1223
character is considered to be valid only at the infrageneric level (Hooper 1987; van Soest
et al. 1991).
There is still some debate about the correct family placement of Acarnus, and this stems
mainly from the different emphasis given by various authors to the importance of chelae
geometry at the family (or even generic) level. Acarnus is included in the Myxillidae,
following the scheme of de Laubenfels (1936) and repeated by Bakus (1966), van Soest
(1984a) and Hooper (1987), because it has a specialised tangential layer of ectosomal tylote
megascleres bearing spined bases, the primary apomorphy for the family, as well as a
more-or-less renieroid reticulate skeleton shared by many (but not all) members of the
family. The alternative point of view is that the genus is not a 'typical myxillid' because
it has palmate isochelae and toxa microscleres, and might better be included in the
Microcionidae (e.g. Burton 1959; LCvi 1973; Vacelet et al. 1976). Certainly, many of the
nominal myxillid genera have arcuate isochelae, but this feature cannot be considered to be
apomorphic for the family (being also found in many other families of PoeciloscleridaCrellidae, Anchinoidae, Coelosphaeridae, Microcionidae, Hymedesmiidae), nor is it found
in all genera of Myxillidae: other myxillids have only anchorate isochelae (e.g. Ectyomyxilla
Lundbeck, Plocamissa Burton), birotulate isochelae (e.g. Iotrochota Ridley, Iotrochopsamma
de Laubenfels, Sigmarotula Bergquist & Fromont), palmate anisochelae (e.g. Acanthorhabdus
Burton), unguiferous isochelae (some Myxilla Schmidt), or bidentate sigmoid isochelae
(Plumocolumetta de Laubenfels), and some genera have various combinations of these
chelae spicules, such as arcuate and unguiferous isochelae (e.g. Anomomyxilla Burton),
anchorate and arcuate isochelae (e.g. Forcepia Carter), anchorate and unguiferous isochelae
(e.g. Fibulia Carter, Stelotrochota Bakus), palmate anisochelae and birotulate isochelae
(e.g. Iophon Gray), or unguiferous and birotulate isochelae (e.g. Iotaota de Laubenfels).
It should also be noted that a similar range of chelae diversity is also found in the
Microcionidae, and the relative importance of these microscleres, at different levels of
classification, has been discussed by Hooper (1991a). Apart from Acarnus, only one other
genus (Allocia Hallmann), presently included in Myxillidae, has palmate isochelae, and both
these genera have also been previously included in the Microcionidae [by Hallmann (1920)
and Burton (1959) respectively], on the basis of their possession of palmate isochelae and
toxa microscleres. However, in the senior author's opinion, this interpretation ignores
the importance of the tangential ectosomal skeleton of tylote megascleres as the primary
apomorphy of the Myxillidae, and it also over-emphasises the importance of chelae
microsclere geometry as a family character. Hooper (1991a) suggested that, from a phylogenetic perspective, the significant difference between the Microcionidae and other
poecilosclerid families, including Myxillidae, concerned the geometry, derivation and
distribution of megascleres within the choanosomal and ectosomal skeletons, whereas the
presence or absence of particular skeletal structures or particular forms of microscleres was
of lesser importance at the family level. It was also demonstrated that extraordinary levels
of homoplasy occurred within the classification when both these sets of features (i.e. the
geometry, origin and distibution of megascleres and chelae microsclere geometry) were used
in combination and given equal importance in the classification. It was also suggested that
because the conjunctive use of these characters produced high levels of homoplasy, then a
parsimonious classification based on rnicrosclere geometry (e.g. de Laubenfels 1936) was
probably mutually exclusive from one based on structural megascleres, as proposed in part
by van Soest (1984a) and developed further by Hooper (1991a). The implication of accepting
this latter scheme is that modifications to the geometry of chelae microscleres, which is
interpreted by the senior author as being one of grade and not one of clade, must have
occurred independently in several poecilosclerid families. This debate is not yet resolved,
and for now Acarnus is retained in the Myxillidae following van Soest (1984) and Hooper
(1987).
Acarnus caledoniensis, sp. nov.
(Figs 1-2, Table 1)
Material Examined
Holotype. QM G300724 (ORSTOM R806): stn 124, Il6t Maitre, 22°20~1'S.,166025~1'E.,
23 m
depth, 15.vii.1976, coll. P. Laboute, SCUBA.
1224
J . N. A. Hooper and C. Ltvi
Paratype. QM G300749 (ORSTOM R806): same locality.
Other New Caledonian material. ORSTOM R499: stn 124, Ilbt Maitre, 22°20.1'S.,166025.l'E.,
25 m depth, 1.x.1991, coll. G. Bargibant, SCUBA; ORSTOM R522, R528: stn 147, ENE. Ilbt N'Due,
22°12~0'S.,166016~(YE.,
22 m depth, 8.ix.1976, coll. P. Laboute, SCUBA; ORSTOM R617: stn 160,
W. reef flat, Ilbt Maitre, 22°20.0'S.,166023l'E., 5 m depth, 13.x.1976, coll. A. Intts, SCUBA.
Description
Colour. Dark red 'fistules' (Munsell 5R 4/10), red-orange body (5R 6/6) alive, siltcovered in situ, light brown in ethanol.
Shape. Small spherical, subspherical or massive sponges, up to 90 mm diameter and
45 mm high, not usually firmly attached to substrate but able to roll around freely over
sand and rubble.
Surface. Covered with silt and evenly distributed microconules, up to 3 mm high, only
seen in live state. Several prominent oscules on upper surface, each 8-15 mm diameter,
8-18 mm high, each surrounded by a prominent membraneous lip ('fistule'); 'fistules' and
microconules usually collapse upon preservation; surface simply optically smooth with
partially translucent ectosomal membrane when preserved. 'Under-side' of sponge usually
retains embedded detritus (but this may not be a permanent feature as the sponge appears
to be able to roll over the substrate).
Ectosome. Skeleton at surface microscopically hispid, with paratangential layer of
tylotes concentrated in peripheral region, many protruding through surface. Isochelae
abundant and collagenous spongin moderately heavy in peripheral skeleton; collagen granular
and brown pigmented.
Choanosome. Main skeleton plumose, slightly reticulate, vaguely halichondroid in
arrangement due to preponderance of echinating spicules interconnecting adjacent skeletal
columns. Skeletal tracts composed of basally spined subtylostyles, containing few (5-10)
spicules abreast, 30-80 pm wide; adjacent tracts run more-or-less parallel to each other,
ascending to surface, with numerous plumose styles protruding through tracts at acute
angles, but only few cross-connections between them. Two size classes of both echinating
acanthostyles and cladotylotes present, all relatively abundant, embedded at acute angles to
skeletal tracts, together producing misleading impression that skeleton is reticulate. Spicules
not visibly enclosed within spongin fibres, although collagenous spongin abundant around
skeletal tract. Numerous microscleres dispersed between spicule tracts, with radial columns
of toxas particularly abundant. Choanocyte chambers oval, 40-70 pm diameter; chambers
frequently vaguely outlined by plumose tracts of both toxas and isochelae.
Megascleres. Choanosomal principal subtylostyles long, slender, slightly curved, tapering
to sharp points, with swollen tylote bases, usually microspined; some longer, more slender,
sinuous subtylostyles also present usually with smooth tylote bases: length holotype 321(354.3)-423 prn [paratype 266-(343.1)-394 pm), width 4-(7.5)-10 pm (4-(7.4)-11 pm].
Ectosomal tylotes long, slender, straight or very slightly curved asymmetrically, with
prominently swollen, invariably microspined, tylote bases: length holotype 304-(330.1)365 pm [paratype 299-(337.8)-383 pn), width 3-(5.0)-6 pm (3-(5.2)-7 pm] .
Cladotylotes of 2 sizes: larger forms (type I) relatively long, slender, slightly curved, with
recurved spines concentrated in middle of shaft, and shaft thicker at base than at apex; main
cladome at apex with 4 sharply pointed long clads, minor cladome at base with 4 small
clads: length holotype 122-(165.7)-192 pm [paratype 166-(184.9)-206 pm], width 1 .5(2.3)-3 pm 12.543 ~5)-4 pm], width at base 2.5-(3.9-5 pm [4-(4.4-6 pm]. Smaller
cladotylotes (type 11) with similar morphology, more slender, usually straight, with granular
spines over shaft: length holotype 68-(I 11.2)-152 pm [paratype 82-(111. @-I40 pm],
width 1.5-(1.3-2 pm [I-5-(2.6)-3 pm], width at base 2-(2.4)-3 a5 pm [2-5-(3-4)-4 pm].
Several examples of 'intermediate'-sized cladotylotes, 150-170 pm long, also occur, and the
recognition of the two size categories defined above is based on a bimodal distribution of
measured spicules.
Acanthostyles of 2 sizes, divided on the basis of a bimodal distribution of spicule
New Caledonian Poecilosclerids
1225
measurements; both size categories cylindrical, straight, fusiform, with slightly swollen
base, with fewer spines in centre of shaft than at ends; larger acanthostyles relatively rare:
length holotype 95-(111.7)-129 pm, [paratype 89-(113.3)-138 pm], width 2-(3.3)-4 pm,
[2-(3 8)-4 pm]. Smaller acanthostyles very abundant: length holotype 69-(76.3)-84 pm
[paratype 69-(77.2)-82 pm], width 2-(3.1)-4 pm [l 5-(2.8)-4 pm].
Microscleres. Only 2 varieties of toxas present. Oxhorn (type I) toxas absent. Smaller
centrally curved (type 11) toxas very thin, with angular central curvature and recurved arms:
length holotype 52-(93 -2)-141 pm [paratype 48-(78.0)-117 pm], width 0.5-(0.9)-1.2 pm
[O-5-(0.9)-1.4 pm]. Larger accolada (111) toxas very long, slender, mostly straight or very
slightly reflexed arms, with angular symmetrical or asymmetrical central curvature: length
holotype 185-(269.6)-386 pm [paratype 119-(243.4-416 pm], width 0.4-(1.1)-1.8 pm
[0.5-(1 3-3 pm]. Some intermediate examples between the type I1 and I11 toxa geometries
were also observed, but generally these were smaller, probably developmental forms of one
or the other type of toxon, and were not included in measurements given above.
Palmate isochelae large, homogenous: length holotype 14-(18.4)-22 pm [paratype 17(19.5)-22 pm].
Fig. 1. Acarnus caledoniensis, sp. nov.: A , holotype QM G300724, choanosomal
principal subtylostyles (enlarged spined base); B, ectosomal tylote; C, cladotylotes (type I);
D, cladotylotes (type 11); E, acanthostyles; F, larger accolada (type 111) toxas; G, smaller
centrally curved (type 11) toxas; H, palmate isochelae; I, fibre characteristics; J, section
through peripheral skeleton.
1226
J. N. A. Hooper and C . Ldvi
Distribution
Known so far only from the south-west coast of the Noumea lagoon, in Halimeda beds,
on sand and coral rubble substrate, rolling across the sea bed or only loosely attached to
benthos, 5-25 m depth.
Remarks
Acarnus caledoniensis, sp. nov. is the 22nd member of the genus so far described, and
ninth member belonging to the 'souriei' species-complex. It is also the most easterly record
of this group in the Indo-Pacific (cf. van Soest et al. 1991: fig. 8).
The species is most closely related to A. bicladotylota and A. tener, which together
comprise a subgroup within the 'souriei' species-complex distributed on the western rim of
the Pacific Ocean, from Japan to the Indo-Malay archipelago and western Micronesia.
It also shows similarities to A. peruanus from the eastern Pacific. Other members of
the 'souriei' species-complex include: A. levii (Vacelet, 1960) (from the Mediterranean),
A. primigenius Hiemstra & Hooper, 1991 (from Indonesia), A . nicoleae van Soest, Hooper
& Heimstra, 1991 (from the Caribbean), A. radovani (Boury-Esnault, 1973) (from Brazil),
and A. souriei (LCvi, 1952) (restricted to populations living in west Africa-Senegal, Cape
Verde and Gulf of Guinea).
It has been shown elsewhere that these 'souriei'-like species have small but consistent
differences from the west African A. souriei (s.s.). In general these morphological differences
correlate with regional species distributions, supporting both the decision to recognise them
as valid species, and the notion that they may have evolved from a previously widespread
'souriei'-like ancestor (van Soest et al. 1991). Therefore, as would be expected by its
transitional distribution, Acarnus caledoniensis has several character states of intermediate
condition between the western (A. bicladotylota, A. tener) and eastern (A. peruanus)
Pacific subgroups.
Fig. 2. Acarnus caledoniensis, sp. nov.: A , holotype QM G300724 (ORSTOM R806) in situ (photo
P . Laboute); B, SEM of cladotylotes (magnified 600 x); C, SEM of skeletal structure (magnified 60x);
D, SEM of echinating acanthostyles (magnified 600 x).
New Caledonian Poecilosclerids
Table 1. Comparison in spicule dimensions between Acarnus
The new species is compared t o other related species in the 'sourieri' species complex. Measurements
are given in micrornetres; comparative data from van Soest et al. (1991)
A. caledoniensis A. souriei
A. bicladotylota A. tener
A. peruanus
A. levii
Massive
Massive
Encrusting
260-340
4-10
244-371
9-12
315-480
6-9
Smooth
Spined
Spined
180-330
2.5-5
21 1-282
3-4
260-460
3-4.5
121-141
4-5
180-210
4.5-6
66-75
3
80-140
2-4
Few
Few
Thick
Thick
82-97
4
55-115
3-4.5
Absent
Absent
Absent
20-140
Subspherical
Encrusting
Shape
Encrusting
266-423
4-11
160-350
3-10
195-460
5-12
Spined
Rugose
Spined
299-383
3-7
210-400
3-5
143-404
3-6
80-180
4-6
80-2 10
4-7
Absent
38-95
1-6
Styles
Base of style
Tylotes
Cladotylotes I
130-190
6
Cladotylotes I1
Absent
Spines on shaft of cladotylotes
Few
Many
Thick
Thick
Thick
100-165
4-6
70-120
1-5
63-90
2-3
Absent
50-80
48-75
Many
Many
Basal part of shaft of cladotylotes
Thin
Acanthostyles I
70-130
4-6
Acanthostyles I1
Absent
Toxas I ('oxhorn')
Absent
70-1 10
Toxas I1 (deeply curved)
50-150
30-212
160-200
130-250
11-15
12-20
35-40
Toxas I11 ('accolada')
Absent
188-21 1
Absent
12-18
13-17
13-16
Isochelae
Acarnus caledoniensis is dark orange, and has a massive, subspherical growth form with
large raised oscules on the upper surface (Fig. 24). It has a plumose, slightly reticulate
skeleton, lacks oxhorn (type I) toxas, has much larger type I11 toxas with an upper range
of 416 pm long, has two sizes of cladotylotes, two sizes of acanthostyles, and has generally
1228
J. N. A. Hooper and C. Ltvi
larger styles and tylotes than its sibling species (refer to Table 1). By comparison, Acarnus
souriei s.s. is red, invariably encrusting, has a plumose skeleton, has all three forms of
toxas, type I11 toxas have an upper range of only 200 pm long, has only one size class of
cladotylotes but two sizes of acanthostyles, and styles and tylotes are generally smaller.
Acarnus peruanus is a thickly encrusting mat, has a plumoreticulate skeleton, also lacks
oxhorn (type I) toxas, type I11 toxas have an upper size range of only 211 pm long,
cladotylotes are smaller and consist of only one size class, there is only one size class of
acanthostyles, tylotes are shorter and thinner, and acanthostyles are relatively rare as
compared to those of A. caledoniensis. Acarnus bicladotylota is red-orange to blood red,
encrusting with embedded detritus, without raised oscules, has a plumose skeleton, has all
three forms of toxas, and type I11 toxas have an upper range of only 250 pm long, it also
has two size classes of cladotylotes, of similar size to those in A. caledoniensis, but it has
only one size of acanthostyles. Acarnus tener is red, massive, with a plumose skeleton, has
all three forms of toxas, type I11 toxas have an upper range of only 230 pm long, and has
only one category of cladotylotes and acanthostyles, similar in size to those found in
A. caledoniensis. The new species should also be contrasted with A. levii-both which show
most similarities in their cladotylote geometry and spination, but A. Ievii is bright red to
maroon-red, encrusting, has a plumose skeleton, lacks accolada (type 111) toxa, has only
one size of acanthostyles, although it also has two size classes of cladotylotes.
In the present contribution, it is not appropriate to represent an emended worldwide
phylogenetic classification of Acarnus (van Soest et al. 1991) to include the New Caledonian
species, but it is possible to hypothesise on where A. caledoniensis fits into the proposed
phylogenetic scheme depicted by those authors (van Soest et al. 1991: fig. 6 ) . As already
noted, A. caledoniensis is most closely related to the 'souriei' group of species. It belongs
to the subgroup of 'souriei' containing A. radovani, A. souriei, A. tener and A. nicoleae,
based on similarities in skeletal structure (being plumose, an apomorphy for the 'souriei'
group), and cladotylote morphology [with none or only minor thickening of the shaft below
the clad; and the presence of hook-shaped spines along the shaft on both size categories
of cladotylotes (both pleisomorphic conditions for the 'souriei' group)]. The new species
branches from (and is more primitive than) this subgroup, in having microspined bases on
styles (which is the ancestral condition; character I l a of van Soest et al. 1991), but it
stands out from those species by its growth form (being subspherical and functioning like
a 'tumbleweed', a new apomorphy for the genus), and the independent loss of type I
'oxhorn' toxas, which is the fourth occurrence of this (homoplastic) synapomorphy in the
genus (also found in A. peruanus, A. deweerdtae, and A. tenuis, all belonging to separate
species-gr oups).
Etymology
Named for the type locality.
Family DESMACELLIDAE Ridley and Dendy
Definition
Sponges with choanosomal axial skeleton ranging from axially condensed, radial,
reticulate, plumo-reticulate or halichondroid, produced by variably compressed reticulation
of spicules enclosed in spongin fibres; extra-axial skeleton plumose or halichondroid.
Megascleres typically monactinal styles or tylostyles, sometimes modified to oxeas or
strongyles, occasionally with flexuous interstitial strongyles. Microscleres sigmas, toxas,
microxeas, raphides (usually in trichodragmata), spheres or commas in all combinations
(after Hooper 1984).
Remarks
There is only one other published record of Desmacellidae from the New Caledonian
region, the hypercalcified sponge Merlia normani Kirkpatrick, 1908, published as
M. deficiens Vacelet, 1980 (Vacelet 1981).
New Caledonian Poecilosclerids
1229
Genus Neofibularia Hechtel
Neofibularia Hechtel, 1965: 22 [type species Amphimedon nolitangere Duchassaing and Michelotti,
1864: 821 [full synonymy and discussion given by Hartman (1967: 3), van Soest (1984a: 77) and
Hooper (1984: 33)].
Diagnosis
Desmacellidae with an irregular isodictyal reticulate choanosomal skeleton, well-defined
spongin fibres, membraneous or tangential ectosomal skeleton, typically smooth surface,
megascleres mostly diactinal (strongyles, oxeas), sometimes styles; microscleres sigmas,
raphides and microxeas (commas in one species); usually toxic (from Hooper et al. 1991).
Remarks
This definition is based on knowledge of only three species prior to the present work
[i.e. N. nolitangere (Duchassaing & Michelotti, 1864), N. irata Wilkinson, 1978, and
N. mordens Hartman, 19671, and from which the genus is supposedly clearly distinguished
from the allied Biemna. On paper, Neofibularia has a reticulate skeleton (cf. plumoreticulate in most Biemna), diactinal (cf. monactinal) megascleres, and all species are toxic
(v. no toxicity) (Hartman 1967: 6). It also apparently differs in having a well-developed
spongin fibre skeleton (cf. poorly developed) and species may produce copious amounts of
clear mucus (Bergquist and Fromont 1988: 30). Although one species of Biemna is also
known to be toxic and have a renieroid reticulate skeleton, B. saucia Hooper et al., 1991,
this species has probably retained ancestral characters no longer present in most other
Biemna species (i.e. a primitive Biemna).
Neofibularia hartmani, sp . nov.
(Figs 3-4, Table 2)
Material Examined
Holotype. QM G300281 (ORSTOM R234; fragment NTM 23891): stn 106, N. entrance, RCcif des
Cinq Milles, SW. New Caledonia lagoon, 22'29.3'S., 166"44.4'E., 8 m depth, 30.iv.1976, coll.
P. Laboute, SCUBA.
Fig. 3. Neofibularia hartrnani, sp. nov.: A , holotype QM G300281, choanosomal strongyles;
B, smaller microoxeas; C, larger microoxeas; D, raphides; E, large sigmas; F, small sigmas;
G, section through peripheral skeleton.
1230
J. N. A. Hooper and C. LCvi
Paratype. NTM 23892 (ORSTOM R1300): stn 192, E. coast IlBt Maitre, 22"20.01S.,16624.OrE.,
1 m depth, 22.ix.1977, coll. P. Laboute, by hand.
Other New Caledonian material. ORSTOM R97: stn 172, SE. Passe de Upe, RCcif de ]'Adventure,
Ile des Pins, 22'31. 1'S.,167°30.0'E., 15 m depth, 7.i.1977, coll. A. Intes, SCUBA; ORSTOM R1244:
stn 268, sand isles, RCcif Croissant, 22°20.2'S.,166022.5'E., 12 m depth, 25.ii.1980, coll. P. Laboute,
SCUBA.
Description
Colour. Pale yellow-brown or khaki-brown alive (Munsell 2.5Y 8/10), often partially
silt-covered, dark brown in ethanol.
Shape. Thickly lobate, massive, bulbous-encrusting sponge with lobes/bulbs fused to
adjacent bulbs, 20-55 mm diameter, up to about 40 mm high, spreading about 120 mm
across dead coral substrate.
Surface. Well developed sculpturing, consisting of close-set, interconnected microconules,
forming web-like striations on surface. Large oscules situated on apex of bulbs, 2-8 mm
diameter, with slightly raised rims, contracting to smaller diameter when preserved.
Ectosome. Membraneous, lightly arenaceous, with abundant heavily pigmented spongin,
but without specialised spicule skeleton. Irregular plumose tufts of choanosomal strongyles
protrude through surface, paratangential or erect on surface, more-or-less corresponding to
position of surface conules; ectosome rarely intact in preserved material.
Choanosome. Skeletal architecture reticulate, with differentiated primary (multispicular)
and secondary (uni- and paucispicular) spongin fibres. Primary fibres usually fully cored
with choanosomal strongyles, 50-90 pm diameter; secondary fibres with fewer spicules
Fig. 4. Neofibularia hartmani, sp. nov.: A, holotyue OM G300281 (ORSTOM R234) in situ
(photo P. Laboute); B, SEM of fibre node (magnified 4 0 0 ~ ) ;C, SEM of skeletal structure
(magnified 200 x); D, SEM of sigma microscleres (magnified 1000 x ).
New Caledonian Poecilosclerids
1231
packed abreast, 15-40 pm diameter. Fibre reticulation produces elongate, oblong or oval
meshes, 35-140 pm diameter, with moderate quantities of collagen and foreign particles,
and abundant microscleres. Choanocyte chambers not observed, obscured by foreign debris,
microscleres and loose megascleres.
Megascleres (refer to Table 2 for dimensions). Choanosomal strongyles long, thick or
thin, slightly curved, symmetrical or asymmetrical.
Table 2. Comparison in spicule dimensions between species of Neofibularia
Measurements are given in micrometres. Spicule numbering convention follows Wilkinson (1978).
Comparative material of N. irata consists of previously unpublished specimens from the Great Barrier
Reef (NTM 21113-14)
N,hartmani,
sp. nov.
N. irata
Wilkinson
N.nolitangere
(Hartman)
N. mordens
Hartman
Source of measurements
Present study
Present study
218-(257.3)-280
3-(5.3-9
(strongyles)
128-(163.4)-179
1.5-(2.1)-3.0
(strongyles)
28-(38.0)-51
0.5-(0.9)-1.2
24-(31.2)-37
0.4-(0.5)-0.8
69-(79.1)-94
0.8-(1.6)-2.0
54-(58.3-67
0.8-(1.1)-1.5
Absent
Absent
79-(86.5)-115
0.2-(0.6)-0.8
60-(67.4)-84
0.2-(0.3)-0.3
13-(22.4)-36
0.8-(1.1)-1.5
14-(31.8)-41
0.5-(0.9)-1.5
50-(67.3-85
1.8-(2.4)-3.5
Absent
Hartman (1967)
Hartman (1967)
Choanosomal megascleres
167-(326)-433
3.1-(9.9)-24.4
(strongyles and
styles)
235-(314-357
4.4-(10.2)-13.4
(styles)
Absent
Absent
Microxeas I
Microxeas I1
Absent
Microxeas 111
98-(122)-150
1.8-(2.9)-5.2
Raphides
82-(105)-126
0.6-(1.1)-2.0
Absent
Sigmas I
14-(19)-24
0.9-(1 .+Is6
Sigmas I1
Absent
Sigmas I11
Absent
Absent
Absent
Absent
Absent
Commas
5-(7.9)-12
Absent
J. N. A. Hooper and C. L6vi
1232
Microscleres. Microxeas clearly divided into two sizes, both relatively long, stout, widest
at midsection.
Raphides abundant, long, straight, hair-like.
Sigmas incompletely divided into two sizes, both with c- and s-shaped examples, relatively
thick, evenly curved at centre.
Distribution
Known only from the southern New Caledonian lagoon, from the reef flat and shallow
subtidal, on dead coral substrate, 1-15 m depth.
Remarks
This species is most similar to N. irata in the diversity and geometry of its spicules, but
otherwise the two species differ substantially in most features: spicule dimensions (Table 2),
growth form (Fig. 4A; N. irata is a flat spreading sponge with erect, leaf-like, folded
lamellae), surface features (N. irata has a smooth unornamented surface), and skeletal
structure (N. irata has a fragile, flattened, honeycomb-like fibre reticulation, cored by
whispy, hair-like tracts of thin strongyles). It is also suspected that N. hartmani produces
a toxic reaction in humans upon contact with skin (Dr C. Debitus, personal communication),
as is typical for the genus, but this has yet to be substantiated experimentally.
Etymology
This species is named for Professor Willard Hartman, Peabody Museum of Natural
History, Yale University, for his pioneering work on Neofibularia, and his other eminent
contributions to the biology of the Porifera.
Family CRELLIDAE Hentschel
Definition
Poecilosclerida with a thick ectosomal skeleton, usually a crust of tangentially placed,
densely packed acanthoxea or acanthostyles, a regularly reticulate choanosomal skeleton of
smooth diactinal megascleres grouped into vague bundles, sometimes reduced to a basal layer
of acanthostyles erect on substrate in encrusting forms, and spined megascleres may also be
distributed within choanosome, between tracts of smooth diactines. Microscleres may include
arcuate isochelae, sigmas, less often arcuate anisochelae (modified from van Soest 1984~).
Remarks
There is only a single previous record of the family Crellidae from the New Caledonian
region, Yvesia acanthosclera LCvi & LCvi, 1983b, from deeper waters (425-430 m depth).
Genus Crella Schmidt
Crella Gray, 1867: 521 [type species Cribrella elegans Schmidt, 1862: 701 [full synonymy and
discussion of the literature given by Bergquist and Fromont (1988: 76)].
Diagnosis
Crellidae with an ectosomal skeleton composed of erect or tangential acanthoxeas in
densely packed tracts; primary choanosomal skeleton with plumose, plumo-reticulate or
reticulate spongin fibres cored by diactinal megascleres (oxeas or tornotes), with or without
echinating acanthostyles; secondary choanosomal skeleton a more-or-less even renieroid
reticulation overlaying primary skeleton; in encrusting forms skeleton reduced to a basal
layer of fibres lying on substrate with erect acanthostyles embedded in spongin; microscleres
usually include arcuate isochelae, sometimes arcuate anisochelae, and sigmas.
Remarks
The genus Crella has not been substantially revised in the contemporary literature,
although descriptions of several species have been published since the 1980s (e.g. van Soest
New Caledonian Poecilosclerids
1233
1984a; Bergquist and Fromont 1988). These works (and Hallmann 1912) should be consulted
for a more detailed overview of the genus.
Crella spinulata (Hentschel)
(Figs 5-6, Table 3)
Grayella spinulata Hentschel, 1911: 340, text-fig. 29.
Crella spinu1ata.-Burton, 1934: 553; Hooper et al., 1992: 263.
Material Examined
Holotype.
ZMH: presently unknown.
New Caledonian material. QM G300014 (ORSTOM R1228): stn 261, SW. Il6t Nda, Lagon Sud,
21°52.5'S,166"51~2'E., 33 m depth, 4.xii.1979, coll. P. Laboute, SCUBA; NTM 23871 (fragment
QM G300686): Baie des Citrons, Noumea lagoon, 22°18.0'S.,166025.2'E., 3 m depth, 25.ix.1990,
coll. M. Kelly-Borges, snorkel.
Comparative material. Houtman-Abrolhos Islands, W.A.: NTM 22916, 2917 (fragment QM
300194): N. of North Wallabi I., 28°17.9'S.,113047.8'E., 39 m depth, ll.vii.1987, coll. J. N. A.
Hooper, USSR RV 'Akademic Oparin', beam trawl; QM G300221 (fragment NTM 23216): N. edge
of Pelsart I., 28°47.2'S.,113058.5'E., 22 m depth, 10.vii.1987, coll. J. N. A. Hooper, USSR RV
'Akademik Oparin', SCUBA; NCI Q66C-4228-M (fragment QM G300687): NW. side of Goss Passage,
Wallabi Group, 28929'S.,113°37'E., 18 m depth, 13.ix.1990, coll. NCI, SCUBA; NCI Q66C-4629-W
(fragment QM G300502): top of Assail Bank, between North I. and Wallaby Group of Is, 28'211S.,
113"42'E., 15 m depth, 16.ix.1990, coll. NCI, SCUBA. Northwest Shelf, W.A.: NTM 2645: W.
of Eighty Mile Beach, 19°33.1'S.,119005.1'E., 35 m depth, 04.v.1982, coll. CSIRO R.V. 'Sprightly',
dredge; NTM 2674: N. of Port Hedland, 19"16'S.,118"50'E., 70 m depth, 04.v.1982, coll. CSIRO
R.V. 'Sprightly', dredge; NTM 2746: N. of Eighty Mile Beach, 19°16'S.,118050'E., 75 m depth,
04.v.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 2744: N. of Adele I., Collier Bay, 15'58.3'S.,
122"39.7'E., 59 m depth, 21.iv.1982, coll. CSIRO R.V. 'Sprightly', dredge; NTM 21771: W. of
Port Hedland, 19°03.3'S.,l18049.9'E., 82 m depth, 29.viii. 1983, coll. Trevor Ward, R.V. 'Soela',
trawl; NTM 22295: NW. of Lacepede Is, 16°31'S.,121028'E., 38-40 m depth, 17.iv.1985, coll. Barry
Russell, pair trawl; NTM 23038: N. of Amphinome Shoals, 19°19.7'S.,119008.8'E., 50 m depth,
19.vii.1987, coll. J. N. A. Hooper, USSR R.V. 'Akademik Oparin', beam trawl. Darwin Harbour,
N.T.: NTM 22695: East Point Aquatic Life Reserve, 12°24.5'S.,130048~O'E.,9-12 m depth, 03.iv.
1986, coll. J. N. A. Hooper, A.-M. Mussig, C. Hood, SCUBA. Arafura Sea: QM G301206: Flattop
Bank, NE. Joseph Bonaparte Gulf, 12°16'S.,129015'E., 17.v.1992, 32 m depth, coll. J. N. A. Hooper,
dredge. Great Barrier Reef, Qld: NTM 21513: E. of Murdock I., Howick Group, 14°36'S.,145003'E.,
14 m depth, 18.ix.1979, coll. Qld Fish. Service; NTM 21575: E. of Coquet I., Howick Group,
l4"32.2'S., 145"07.5'E., 22 m depth, date unknown, coll. Qld Fish. Service; NTM 21568: Stanley
Reef, outer slope and lagoon, 19°16'S.,148006'E, 13 m depth, 15.x.1980, coll. Alice Kay; NTM 23514:
Dingo Reef, NE. edge of reef front, Whitsunday Is Group, 19°07.2'S,148028.7'E., 12 m depth,
20.xi.1988, coll. NCI, SCUBA.
Description
Colour. Red-orange alive (Munsell 2.5R 5/10), orange-brown in ethanol.
Shape. (QM G300014): Groups of solitary, conical or cylindrical digits, up to 60 mm
high, 20 mm basal diameter, 9 mm diameter at apex, erect on substrate, with common
basal attachment on or below substrate. Digits evenly cylindrical or slightly flattened.
(NTM 23871): Bulbous encrusting in sand sediment.
Surface. Slightly conulose, granular surface, with visible ectosomal membrane. Oscules
prominent in live material, evenly dispersed over sides and apex of digits, about 1-2 mm
diameter, slightly raised above surface 2-3 mm high.
Ectosome. Thick crust of acanthoxeas lie tangential to surface, 100-350 pm thick, with
plumose brushes of smooth oxeas, arising from major choanosomal spicule tracts, standing
perpendicular to surface crust and protruding slightly through it, but more-or-less obscured
by tangential surface crust. Moderately light granular collagen permeates surface crust.
1234
J. N. A. Hooper and C . Lhi
Choanosome. Skeleton composed of 2 distinct structures. Main skeleton a meandering
reticulation of thick, well-formed spongin fibres fully cored by multispicular tracts of smooth
oxeas, 50-85 pm diameter, eventually culminating in erect plumose brushes of smooth oxeas
at surface. Fibre meshes relatively cavernous, 140-220 pm diameter, oval to eliptical, with
light collagen visible around eliptical choanocyte chambers, 60-130 long, 30-50 pm wide.
Secondary skeleton renieroid, more-or-less regularly reticulate, overlaying primary reticulate
fibre skeleton, each element of renieroid reticulation composed of 2 or more acanthoxeas
joined at nodes by collagenous spongin. Secondary skeleton ceases about 200-250 pm from
surface crust. Most megascleres confined to one or the other skeletal structures, few spicules
scattered within mesohyl.
Megascleres (refer to Table 3 for dimensions). Choanosomal fibres cored by smooth
oxeas, long, thin, slightly curved, usually symmetrical, occasionally modified to styloids,
usually with hastate points.
Ectosomal and choanosomal acanthoxeas identical in morphology and size, long, thin,
symmetrically curved, with small granular spines evenly dispersed over spicule and with
sharp fusiform points.
Microscleres. Isochelae arcuate, moderately large, uncommon; thinner, poorly silicified
examples also seen, with rudimentary development of alae, presumably juvenile forms of
large chelae.
Distribution
Tropical Australasia: Shark Bay, W.A. (Hentschel 1911); Houtman-Abrolhos Islands,
W.A.; Northwest Shelf, W.A.; Timor Sea, N.T.; Darwin Harbour, N.T.; Penguin Channel,
Snake Reef, Stanley Reef, Howick Islands and Whitsunday Islands, Great Barrier Reef, Qld
(6-55 m depth) (Burton 1934; present study); and New Caledonia (3-33m depth; present
study). Partially burrowing into soft sediments (sand, gravel, shell-grit, coral rubble), or
bioeroding dead coral substrates, rarely found on live coral substrate.
Fig. 5 . Crella spinulata (Hentschel): A, specimen QM G300014, choanosomal oxeas; B, ectosomal
and choanosomal acanthoxeas; C,arcuate isochela; D, section through peripheral skeleton.
New Caledonian Poecilosclerids
1235
Remarks
The holotype of this species is presently unknown, possibly extant in the ZMH. Nevertheless, the species is perfectly recognisable from Hentschel's (1911) description. In addition,
abundant fresh material has been recollected from the south-, mid- and north-west coast of
Western Australia (including from areas close to the type locality in Shark Bay), and also
Fig. 6. Crella spinulata (Hentschel): A ,
specimen QM G300014 (ORSTOM R1228)
in situ (photo P. Laboute); B, specimen
ORSTOM R177 in situ (photo P. Laboute);
C, specimen QM G300014 (ORSTOM
R1228), SEM of peripheral skeleton
(magnified 150x); D, SEM of acanthoxeas
(magnified 1000 x ).
Table 3. Comparison in spicule dimensions between specimens of CreNa spinulata (Hentschel, 1911)
Measurements are given in micrometres; n, number of specimens examined
-
-
-
-
New Caledonia
Holotype
Shark Bay, W.A.
Present study
(n = 2)
Hentschel 1911
-
West coast
Australia, W.A.
East coast
Australia, Qld
Present study
(n = 14)
Present study
(n = 4)
Source
L
W
256-(244.0)-33 1
1.8-(2.9)-4.0
Choanosornal smooth oxeas
223-(236.8)-271
264-328
3.0-(5.8)-7.0
3-5
L
W
148-(152.4-157
1.5-(1.8)-3.0
120-152
4-5
228-(288.5)-355
2.5-(4.9-7.0
Acanthoxeas
95-(119.3)-147
1 .O-(4.3)-7.0
122-(143 .O)-162
2.0-(4.1)-6.0
12-(19.0)-28
16-(21.2)-25
Arcuate isochelae
L
18-(20.2)-22
16-20
1236
J. N. A. Hooper and C . Lkvi
from the north-east coast of Queensland. Some of this material has already been analysed
in an earlier chemotaxonomic study (Hooper et al. 1992). There is little doubt, on paper at
least, that all this material is conspecific with Hentschel's species, but it is also possible that
presently undetectable sibling species make up C. spinulata, given the great variation in
growth forms possible. A digitate branching morphology is typical, but encrusting and
massive specimens are also known.
The two representatives of the Noumea population, described above, represent new
locality records for the species and differ from most other specimens collected from
Australian waters in having only acanthoxeas, i.e. in many specimens of this species, and
indeed in many species of Crella, acanthoxeas frequently co-exist in equal proportions with
acanthostyles, or sometimes only acanthostyles are present (e.g. Bergquist and Fromont
1988). This feature may eventually be found to correlate with other differences (e.g. growth
forms), and hence be important in differentiating possible sibling species, but for the present
it not possible to clearly differentiate New Caledonian and Australian specimens on the
basis of morphometric features.
Family MICROCIONIDAE Carter
Definition (emended)
Poecilosclerida with ectosomal skeleton composed of styles or anisoxeas (exceptionally
oxeas), in erect bundles, forming continuous crust, lying tangential or sparsely dispersed
on surface. Choanosomal (or axial) skeleton with well-developed spongin fibres forming
hymedesmoid, microcionid, plumose, plumo-reticulate, reticulate or axially condensed tracts.
Subectosomal (or extra-axial) skeleton rarely well developed (cf. Raspailiidae), although
subectosomal (auxiliary) spicules may be dispersed between fibres, sometimes forming
ectosomal structures, and occasionally producing distinct extra-axial tracts (Axociella).
Spongin fibres cored by smooth or partially spined large styles, occasionally modified to
quasidiactinal forms (Echinochalina) and sometimes replaced partly or entirely by detritus
(Clathriopsamma), and with wholly or partially spined small styles (acanthostyles), or
modified forms such as acanthoxeas, acanthostrongyles or smooth styles embedded perpendicular to (i.e. echinating) fibres. Microscleres include diverse forms of isochelae, toxas
and occasionally raphides or microxeas.
Remarks
The Microcionidae is a vast and diverse group of the Porifera. Worldwide there are
522 nominal species, of which probably 432 species are valid, and in the Australian fauna
(i.e. including coastal, shelf, oceanic island and subantarctic island species) there are
140 valid species (or about 32% of the world's microcionids).
In the past, reliable species identifications of Microcionidae have amounted to little more
than educated guesswork, considering that the literature is vast, scattered, and many species
are neither adequately illustrated nor well recognisable from their published descriptions.
Furthermore, species seem to exhibit virtually every combination of characters, and virtually
every character has dozens of character states, making the clear delineation of even the
genera difficult (e.g. Hooper 1991~).Towards a solution, a major revision of species and
higher taxa within the family is currently in progress, which will make the Australian fauna,
at least, more accessible.
Prior to the present work, only two published records of Microcionidae existed for the
New Caledonian region, Antho novizelanica (Ridley, 1881) (LCvi and LCvi 1983b) from
deeper water (250-350 m depth; also found in New Zealand), and a shallow-water species
originally assigned to Clathria australiensis (Carter) from southern Australia (LCvi 1967),
but which is described here as a new species (Clathria rugosa, sp. nov.). The present study
adds another 12 species to the shallow lagoon and coral reef fauna, bringing the total
number of microcionid species known for the region to 14, of which 10 are suspected
endemics.
New Caledonian Poecilosclerids
Genus Clathria Schmidt
Restricted synonymy
Clathria Schmidt, 1862: 57 [type species Clathria compressa Schmidt, 1862: 581.
Thalysias Duchassaing and Michelotti, 1864: 82 [Spongiajuniperina Lamarck, 1814: 4441.
Clathriopsamma Lendenfeld, 1888: 227 [type species Clathriopsamma reticulata Lendenfeld, 1888:
2271.
Axociella Hallmann, 1920: 779 [Esperiopsis cylindrica Ridley and Dendy, 1886: 3401.
Dendrocia Hallmann, 1920: 767 [Clathriapyramida Lendenfeld, 1888: 2221.
Isociella Hallmann, 1920: 784 [Phakellia jacksoniana Dendy, 1897: 2361.
Diagnosis (emended)
Microcionidae with 1 (subgenus Clathria) or 2 (subgenus Thalysias) categories of auxiliary
styles or subtylostyles, sometimes absent entirely, forming peripheral skeleton; choanosomal
spicules usually enclosed within spongin fibres, sometimes simply with nodal spongin,
consisting of principal styles or subtylostyles usually well differentiated from auxiliary styles;
sometimes fibres cored by undifferentiated auxiliary spicules (subgenus Dendrocia) or simply
cored by detritus with or without megascleres (subgenus Clathriopsamma); echinating
acanthostyles usually present, sometimes absent (subgenera Axociella, Isociella), occasionally
with vestigial spination or completely smooth; choanosomal skeleton varies from leptoclathriid, microcionid, plumose, plumo-reticulate, renieroid (subgenus Isociella), to reticulate,
with (subgenus Axociella) or without structural differences between axial and extra-axial
skeletons; microscleres present or absent, typically consisting of palmate isochelae but
sometimes including other diverse forms of isochelae, and smooth or spiny toxas.
Remarks
Within Clathria (with five subgenera recognised here) there are 383 described species
worldwide, of which 328 appear to be valid; there are also another 9 recognisable but as
yet unnamed species in the literature, as well as many other unrecognisable species.
As noted above, only a single record of the genus has previously been published for the
New Caledonia region, Clathriopsamma australiensis (Carter) (LCvi 1967), although this
population is now considered to represent a new sibling species of the Australian taxon.
The emended definition of Clathria presented here allows the inclusion of species with
isochelae other than the usual palmate form. Various nominal genera have been created
for species otherwise similar to Clathria, but with modified chelae, such as cleistochelae
[e.g. Wetmoreus de Laubenfels (no material seen)], arcuate isochelae [e.g. Quizciona
de Laubenfels (BMNH 1901.12.26.13), Paradoryx Hallmann (AM G3046)], anchorate
isochelae [e.g. Cionanchora de Laubenfels (MNHN DT939), Folitispa de Laubenfels
(BMNH 1921.11.7.69)], and bidentate sigmoid isochelae [e.g. Damoseni de Laubenfels
(SMF969)l. Under some interpretations (e.g. Burton 1959), these genera should be assigned
to different families, whereas Hooper (1991a) has suggested that their importance at the
suprageneric level is questionable (see also Remarks above for Myxillidae and Acarnus).
The use of the subgeneric classification, rather than the retention of these taxa at the
generic level, is based on a phylogenetic analysis of the Microcionidae (Hooper 1991a).
The development of this system, to apply to a geographically wider fauna, is presently in
development by the senior author.
Clathria (Clathriopsamma) rugosa, sp. nov.
(Figs 7-8, Tables 4-5)
Clathria australiensis. - Ldvi, 1967: 22, fig. 6, pl. 2, fig. D.
Not Wilsonella australiensis Carter, 1885: 366.
Material Examined
Holotype. QM G300278 (fragment NTM 23880): Ile aux Canards, Noumea lagoon, New Caledonia,
22°19~0'S.,166026~O'E.,
22 m depth, 26.ix.1990, coll. J. N. A. Hooper, SCUBA, in sand, rubble,
sea grass beds.
1238
J. N. A. Hooper and C. LCvi
Paratype. QM G300696 (fragment NTM 23889): Baie des Citrons, Noumea, New Caledonia,
22"18'S.,l66"25'E., 3 m depth, 15.ix.1990, coll. M. Kelly-Borges, snorkel, fringing coral reef.
Other New Caledonian material. Nominotypical variety (flabellate-digitate, without astrorhizae):
ORSTOM R235: stn 182, NE. IlBt KouarC, 22°46.1'S.,1660471'E., 24 m depth, 7.vi.1977, coll.
P. Laboute, SCUBA; ORSTOM R194: stn 185, Pt Eacho, Lifou (ChCpCnCe), 20°47.0'S.,167007.1'E.,
12 m depth, 19.vi.1977, coll. P. Laboute, SCUBA; ORSTOM R547: stn 150, S. of entrance, Baie
St Vincent, 22"02.2'S.,165"59.5'E., 18 m depth, 9 . k 1976, coll. P. Laboute, SCUBA; ORSTOM R276:
stn 193, E. coast, Rtcif Tabu, 22°29.1'S.,166026.1'E., 30 m depth, 13.x.1977, coll. G. Bargibant,
SCUBA; ORSTOM R558: stn 148, NE. pt, IlBt Mboa, 22°08.3'S.,166009.3'E., 13 m depth, 9.ix.1976,
coll. P. Laboute, SCUBA; ORSTOM R771: stn 206, Banc de la Torche, Ile des Pins, 22051.11S.,
167040.01E.,36 m depth, 8.iii.1978, coll. P. Laboute, SCUBA; ORSTOM R1288: stn 302, ENE. lagon
Belep, Lagon Nord de la Grande terre, 19°39~0'S.,161"33.OIE., 38 m depth, 22.vi.1981, coll. J.-L.
Menou, SCUBA; ORSTOM R620: stn 124, IlBt Maitre, 22°20.1'S.,166025.1'E., 20 m depth, 15.vii.
1976, coll. P. Laboute, SCUBA; ORSTOM R466: stn 109, Baie des Citrons, Noumea, 22018.01S.,
166"25.11E., 10 m depth, 31.v.1976, coll. P. Laboute, SCUBA. Variety with astrorhizae, flabellatedigitate: QM G301265 (ORSTOM R157) ('representative specimen'): stn 124, I16t Maitre, 22"20,11S.,
166"25.11E., 20 m depth, 30.xi.1977, coll. G. Bargibant, SCUBA; ORSTOM R586: stn 147, midchannel between Ton Du (Tiart) and IlBt Ie, 22°121'S.,166016~5'E., 24 m depth, 8.ix.1976, coll.
P. Laboute, SCUBA: ORSTOM R260: stn 114, NE. IlBt TarCti, Banc Gail, 22°221'S.,166039~0'E.,
35 m depth, 29.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R263: stn 110, SE. IlBt RCdika,
22'31. 1'S.,166"36.01E., 19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R428: stn 124,
IlBt Maitre, 22°20.1'S.,1660251'E., 20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM
R404: stn 114, NE. Ilat Tartti, Banc Gail, 22°22~1'S.,166039~0'E.,35 m depth, 29.vi.1976, coll.
P. Laboute, SCUBA; ORSTOM R498: stn 124, IlBt Maitre, 22°20.1'S.,166025~1'E., 20 m depth,
15.vii.1976, coll. P. Laboute, SCUBA; ORSTOM R626: stn 162, W. (lee) of Ilat KiC (Havannha),
22°221'S.,167004~WE.,15 m depth, 26.x.1976, coll. A. Intis, SCUBA; ORSTOM R1069: stn 249,
NW. I16t Vua, Lagon Sud, 2lo33.4'S. 166"43.5'E., 35 m depth, 10.xii.1978, coll. P. Laboute,
SCUBA; ORSTOM R1304: stn 315, NE. IlBt Dii-Ami, Ile des Pins, 21°35.5'S.,167017.4'E., 33 m
depth, 30.vii.1981, coll. P. Laboute, SCUBA. Variety with fistules, massive: QM G301261 (ORSTOM
R1418) ('representative specimen'): stn 453, Nokanhoui, Ile des Pins, 22"43.3'S.,167O35.2'E., 40 m
depth, 27.xi.1983, coll. J.-L. Menou, SCUBA; QM G300870 (fragment NTM 23881, ORSTOM
cfR53): stn 158, Baie de Saint-Vincent, between Passe de St Vincent and Hugon, 22°03.0'S.,165059.0'E.,
10 m depth, 16.ix.1976, coll. P. Laboute, SCUBA.
Description
Colour. Bright red-orange alive (Munsell 5R 5/10), rarely dark orange (10R 6/10);
light grey-brown in ethanol.
Shape. Flabellate, palmate-digitate or simply digitate sponges, occasionally massive,
subspherical, varying considerably in size (70-340 mm long, 45-140 mm maximum width,
10-20 mm thickness), and varying in development of digits, typically consisting of several
short or long cylindrical digits (15-140 mm long), arising from margins of a fan, or at
least with an enlarged point of attachment for digits above basal stalk; usually with short
cylindrical stalk (30-50 mm long) often attached to coral rubble or dead molluscs.
Surface. Nominotypical variety. Optically smooth, even, membraneous surface when
alive, with few microconules on margins of digits, none on lateral surfaces of fans;
ectosomal membrane destroyed upon preservation, surface beneath membrane very porous,
with numerous ostia approximately 0.5-1 mm diameter. Oscules, 2-3 mm diameter, surrounded by symmetrical membraneous lips which usually collapse upon preservation, slightly
raised (up to 2 mm) from surface, without any obvious subectosomal canals (astrorhizae);
oscules located mainly on margins or less frequently scattered over lateral surfaces in live
material, but these are not easily visible after preservation.
Variety with astrorhizae.
Oscules raised above surface with prominent subectosomal canals (astrorhizae) converging
towards central pore, and surrounded by slightly raised membraneous lips, producing
Variety with fistules. Oscules on upper and lateral
prominent subectosomal sculpturing.
surfaces (of subspherical massive growth forms), raised above surface on relatively large
fistules, up to 12 mm long, 4 mm diameter, without subectosomal canals (astrorhizae)
or other surface sculpturing. However, fistules temporary, membraneous, collapsing upon
preservation.
New Caledonian Poecilosclerids
1239
Ectosome. Single category of auxiliary spicules (subectosomal subtylostyles) forms
irregular paratangential brushes on surface. Surface mineral skeleton not dense, usually
consisting of bundles, occasionally individual spicules embedded in or dispersed between
relatively close-set, horny spongin fibres; fibres marginally smaller but more compacted
(80-140 pm mesh size) in peripheral skeleton than in axis of choanosome; small quantities
of detritus (mostly sand grains) embedded in peripheral skeleton, but less abundant than in
axis; light brown collagen and numerous microscleres abundantly scattered in periphery.
Plumose tracts of choanosomal styles, in paucispicular tracts, terminate just below auxiliary
spicules in the peripheral skeleton, and do not protrude through the sponge surface.
Choanosome. Skeleton reticulate, spongin fibres heavy, relatively widely spaced (230400 pm mesh size); spongin fibres divisible into primary and secondary components; primary
fibres (80-115 pm diameter) sparsely cored by uni- or paucispicular tracts of choanosomal
principal styles, usually ascending towards peripheral region; primary fibres heavily echinated
by acanthostyles, dispersed relatively evenly over all fibres; many primary fibres also
contain sand grains, but these do not usually occupy entire fibre diameter; secondary fibres
(20-45 pm diameter) usually uncored, without detritus, but heavily echinated by acanthostyles; collagen moderately heavy, containing abundant microscleres; choanocyte chambers
oval, 50-80 pm diameter.
Megascleres (refer to Tables 4-5 for dimensions). Choanosomal principal spicules
short, straight styles with slightly subtylote bases, rounded or abruptly pointed tips, usually
Clathria (Clathriopsarnrna)
rugosa, sp. nov.: A , paratype QM
G300696, subectosomal auxiliary
subtylostyles (quasidiactinal); B,
choanosomal principal style and
acanthose style; C, echinating
acanthostyles; D, toxas; E, palmate
isochelae; F, section through peripheral
skeleton; 6, fibres characteristic; H,
fibre node.
Fig. 7.
1240
J. N. A. Hooper and C. L h i
completely smooth, occasionally with sparse spines over shaft and both ends; similar in
morphology to auxiliary spicules, differentiated mainly by being slightly shorter and thicker.
Subectosomal auxiliary spicules quasidiactinal modifications of styles, long, straight with
slightly subtylote microspined bases, rounded microspined points, completely smooth shaft.
Echinating acanthostyles short, cylindrical, slightly subtylote bases, rounded, slightly
swollen tips, evenly spined; spines small, granular.
Microscleres (refer to Tables 4-5 for dimensions). Palmate isochelae small.
Toxas with consistent morphology, variable in size, slightly curved at centre, recurved
and sharply pointed at tips.
Fig. 8. Clathria (Clathriopsamma) rugosa, sp. nov.: A , paratype QM G300696, SEM of skeletal
structure (magnified 1 0 0 ~ ) ;B, SEM of fibre characteristics (magnified 600x); C, nominotypical
variety, specimen in situ (ORSTOM R53) (photo P. Laboute); D, specimen in situ (ORSTOM R450)
(photo P. Laboute); E, variety with astrorhizae, specimen in situ (ORSTOM R626) (photo P. Laboute);
F, variety with fistules, specimen in situ (ORSTOM R771) (photo P. Laboute).
New Caledonian Poecilosclerids
1241
Table 4. Comparison in spicule dimensions between Clathria rugosa, C. australiensis and C. bulbosa
Clathria rugosa is compared with the allopatric populations of its sibling species C. australiensis
(Carter), as well as the sympatric C. bulbosa, sp. nov. Measurements are given in micrometres
C. rugosa, sp. nov. C. bulbosa, sp. nov. C. australiensis*
(New Caledonia)
(New Caledonia)
(Vic.)
134-159
4.5-8
Principal
124-180
2-4
Auxiliary
162-206
2.5-4
Spined
184-207
1.5-3
Smooth
55-74
2-4
32-92
0.8-2.5
14-19
A
Choanosomal styles
92-152
2.5-4
Principal
C. australiensisB
(N.S.W.)
C, australiensisC
(W-4.)
92-146
1.5-4
Principal
106-173
2.5-5
Principal
Subectosomal auxiliary styles
92-155
94-141
1.5-4
2-3.5
Spined
Spined
Echinating acanthostyles
45-77
2-4.5
54-78
2-4.5
Toxas
28-89
0.5-1
32-74
0.5-1
Isochelae
11-18
12-16
98-172
2-5
Spined
Lectotype BMNH 1886.12.15.43.
Holotype of Clathiopsamma lobosa Lendenfeld (a synonym of C. australiensis), AM G9053
Holotype of Thorects ramsayi Lendenfeld (a synonym of C. australiensis), AM G8820.
Table 5. Comparison in spicule dimensions between specimens (varieties) of Clathria rugosa
Measurements are given in micrometres
Holotype
nominotypical
(QM G300278)
Paratype
variety
(QM G300696)
'Fistulose'
variety
(QM G300870)
'Astrorhizae'
variety
(QM G301265)
Choanosomal principal styles
L
W
134-(147.8)-159
4.5-(5.8)-8
L
W
162-(188.2)-206
2-(2.8)-4
135-(140.0)-146
4.545.9-7
117-(132.6)-156
2.5-(4.4)-6
134-(145.6)-159
4-(4.9)-6
Subectosomal auxiliary styles
180-(184.8)-194
2.5-(3.3)-4.5
144-(172.0)- 192
2.5-(3.1)-4
173-(182.0)-204
2.5-(3.2)-4
Echinating acanthostyles
L
W
66-(74.2)-91
4.5-(4.9)-6
58-(67.2)-77
4-(5.5)-7
59-(65.8)-72
3-(4.2)-5
55-(65.9-75
3.5-(4.2)-5
L
W
24-(75.2)-122
0.8-(1.8)-2.5
Toxas
35-(68.3)-84
0.8-(1.9)-3
32-(58.2)-84
0.8-(1.9)-3.0
32-(63.2)-117
0.8-(1.5)-2.5
L
13-(15.1)-17
Isochelae
12-(14.0)-16
12-(14.2)-18
13-(14.3)-16
1242
J. N. A. Hooper and C. Ltvi
Distribution
Known so far only from the southern and south-west regions of the New Caledonian
lagoon, on dead coral, coral rubble, Halimeda beds and sand substrates; depth range
3-38 m.
Remarks
There are three distinct populations of this species in the New Caledonian lagoon, as
described above. As far as is known, these populations are sympatric, appear to occupy
similar ecological niches, and two are virtually identical in their external morphology,
coloration, skeletal structure, spicule geometry and spicule size (Table 9, whereas the third,
fistulose variety differs more substantially in being subspherical (Fig. 8F) rather than
flabellate (Fig. 8C, E) or digitate (Fig. 80). However, these varieties are apparently easily
differentiated in the field by their surface sculpturing: the nominotypical variety has oval,
symmetrical, slightly raised oscules; the variety with astrorhizae has prominent subectosomal
canals (astrorhizae) converging towards the centre of each oscule; the variety with fistules
has oscules raised relatively further above the surface, on short membraneous stalks.
SCUBA divers report that these features are distinctive in situ (J.-L. Menou, personal
communication), although they are virtually undetectable after preservation. These three
populations are not formally differentiated here (at the subspecies level) because no other
conventional morphological characters can be found to distinguish them. Nevertheless,
the described material of each variety is separated above to emphasise these apparently
prominent field characteristics, but only non-morphological evidence will eventually
determine whether or not their formal separation is warranted.
Clathria rugosa is a member of the subgenus Clathriopsamma in having its mineral
spicule skeleton supplemented by detritus, mostly sand grains (Fig. 8B). However, this
feature may have a fairly high level of homoplasy for the Demospongiae, probably
independently acquired in several other Poecilosclerida (e.g. Desmacididae) and other
demosponges (e.g. Dysideidae), and must be re-evaluated in a phylogenetic context.
Spongin fibres on the ectosome of C. rugosa are more closely compacted than fibres
within the choanosome, and this feature is reminiscent of the typical Callyspongia condition
(Haplosclerida), but it is undoubtedly one of convergence. Clathria rugosa is also similar to
a small group of other Clathria species- C. (Thalysias) major, C. (Dendrocia) pyramida
and C. (Clathriopsamma) australiensis-in having microspines on both the bases and tips
of auxiliary spicules (Fig. 7 A ) . These spicules at first appear to be diactinal megascleres, and
as such suggest affinities with the Myxillidae, but they are in fact modified styles (i.e. they
are always asymmetrical), and the feature is interpreted here as being one of convergence
with the Myxillidae.
Clathria rugosa and C. australiensis are obviously closely related, showing many
similarities in their skeletal architecture, spiculation and growth form, but there are several
significant differences between them that justify the creation of a new species for the New
Caledonian populations. Moreover, it is possible that the widely distributed, temperate
Australian C. australiensis itself consists of several heterogenous populations, as each
geographic morph differs slightly from the 'original' population from Port Phillip, Vic.
There is presently not enough contemporary material (i.e. in good condition, with accurate
locality data), nor any available non-morphological, corrobative evidence to divide these
populations into sibling species.
Clathria rugosa has field characteristics notably different from C. australiensis (s.s.):
the former with flabellate, flabellate-digitate, digitate or rarely subspherical growth forms,
bright red-orange live coloration, prominent oscules, with or without subectosomal canals
(astrorhizae) or small fistules; the latter predominantly lobate, club-shaped or bulbous, pale
orange alive, and oscules without ornamentation (as far as is known). Similarly, there are
other cryptic morphological differences seen in preserved material that distinguish the two
species. Clathria rugosa has auxiliary spicules with blunt or rounded points, and terminal
spines are perched only on the extremities of these 'points', whereas in C. australiensis there
is a prominent terminal spine/point, and spines occur at least part of the way along the
shaft of the spicule. In this feature C. rugosa resembles most closely C. major. Other
New Caledonian Poecilosclerids
1243
differences are that in C. australiensis acanthostyles are pointed and tapering, toxas are
very thin, strongly curved at the centre and not abundant, and spicule dimensions vary
slightly (Table 4). Clathria rugosa is also similar to another species, described below from
the Chesterfield Islands (approximately midway between the Great Barrier Reef and New
Caledonia), C. bulbosa, sp. nov. (assigned to the subgenus Clathria in lacking detritus
in the skeleton), in its skeletal structure and spicule geometry, although there are several
important but cryptic differences between the two species which indicate that the two are
not conspecific (described below and Table 4).
Etymology
The name rugosa refers to the roughened, rugose texture.
Clathria (Clathriopsamma)litos, sp. nov.
(Figs 9-10)
Material Examined
Holotype. Q M G301269 (ORSTOM R190): stn 109, Baie des Citrons, Noumea, New Caledonia,
22"18.3'S.,l66O25.8'E., 15 m depth, 17.v.1977, coll. P. Laboute, SCUBA.
Description
Colour. Pale orange-brown alive (Munsell 10R 7/8), beige in ethanol.
Shape. Clump of branching, cylindrical digits (clump 240 mm wide, 115 mm maximum
height, 90 mm maximum thickness; individual digits 25-65 mm long, 8-16 mm diameter),
each at least partially fused to adjacent digits, attached to a common base, forming
branching, non-anastomosing growth form. Digits more-or-less regularly cylindrical, usually
bifurcating at least once, and often expanded at apical end, with terminal oscule.
Surface. Highly porous, both when alive and when preserved, with wide fibre meshes
interconnected on surface by ectosomal membrane stretched across adjacent fibres, usually
detached in preserved state, leaving an insubstantial, wide-meshed, honeycombed-reticulate
appearance. Large oscules on apex of each digit, 6-13 mm diameter, surrounded by
membraneous lip. Each oscule with single, prominent (but non-asterose) subectosomal canals
(astrorhizae) running longitudinally from oscule to fused portion of digit. In life subectosomal canals covered by membraneous dermis, collapsing after preservation, leaving large
longitudinal furrow on lateral side of each digit, up to 4 mm wide, 5 rnm deep. Surface
shaggy, microconulose, with fibre endings protruding through ectosomal membrane. Texture
very soft, fragile, insubstantial, easily torn, slightly arenaceous.
Ectosome. Membraneous, with tips of primary spongin fibres protruding for up to
300 pm through ectosomal membrane, 300-700 pm apart. No spicules associated with
ectosomal skeleton, but moderate quantities of detritus, mostly sand grains, scattered on
both exterior and interior of peripheral spongin fibres.
Choanosome. Skeletal structure irregularly reticulate. Spicule skeleton absent (except
for microscleres), but spongin fibre system well developed, differentiated into primary and
secondary fibres. Primary fibres predominantly ascending, 50-70 pm diameter, interconnected by thinner secondary, mostly transverse fibres, 15-35 pm diameter. All fibres
lightly laminated but without central pith, and sparse detritus also incorporated into some
primary fibres (secondaries completely clear). Fibre meshes cavernous, oval or elongate,
variable in size, 130-550 pm diameter. Heavy collagen dispersed within fibre meshes,
pigmented light brown, granular, containing sparse microscleres and abundant detritus (sand
and foreign spicules). Choanocyte chambers oval, 20-40 pm diameter.
Megascleres. Absent.
Microscleres. Arcuate isochelae, large, thick, with well developed alae. Length 16(19.9)-24 pm.
Toxas uncommon, long, thick, slender, with generous central curvature, long straight
(unreflexed arms). Length 114-(164.7)-196 pm, width 1.5-(2.1)-2.5 pm.
1244
J. N. A. Hooper and C. Ltvi
Distribution
Known only from the Noumea region of the lagoon, New Caledonia, coral rubble
substrate, 15 m depth.
Remarks
This species is included with Clathria (Clathriopsamma) on a speculative basis, since it
lacks structural megascleres that would otherwise confirm its affinities. It is assigned to the
Microcionidae in having toxa and chelae microscleres [and resemblance in its digitate growth
form, surface sculpturing and reticulate spongin fibre skeleton to other microcionids, such
as Echinochalina, Clathria (Zsociella)],and it is presently included in the species-group
Clathriopsamma in having detritus incorporated into spongin fibres (Fig. 10D-E). The
'keratose'4ke nature of this species makes its inclusion with the Microcionidae obscure;
that prompted a more thorough than usual investigation of whether or not the recorded
isochelae and toxa microscleres were native to the sponge or acquired. Examination of many
fragments, taken from several different places on the sponge, consistently turned up these
microscleres, albeit they were never more than 'uncommon', supporting the present placement
with Poecilosclerida.
Fig. 9. Clathria (Clathriopsamma) litos, sp. nov.: A , holotype QM
G301269, toxas; B, arcuate isochelae; C , section through peripheral
skeleton.
New Caledonian Poecilosclerids
1245
Thus, the affinities of Clathria litos must be inferred by the geometry of these isochelae
microscleres, having arcuate isochelae instead of the usual palmate sort found in the
majority of microcionids. In fact, several species-groups within the Microcionidae (see
below), and throughout Poecilosclerida in general (e.g. Ectyodoryx in the Myxillidae), have
been created predominantly or exclusively on this basis.
Nominal microcionid genera, defined by possession of arcuate isochelae, include (with the
type material of the type species indicated, seen by the senior author): Anaata de Laubenfels
[type species Aaata spongigartina de Laubenfels (USNM 21428)], Anthoarcuata Bakus [type
species A. graceae Bakus (USNM 36284)], Paradoryx Hallmann [type species Clathria dura
Whitelegge (AM G3046) from N.S.W. and Tas.], Qasimella Thomas [no material seen],
Quizciona de Laubenfels [type species Microciona heterospiculata Brondsted (BMNH
19Ol.l2.26.13)], and Wetmoreus de Laubenfels [type species Microciona novaezealandica
Brondsted (no material seen)]. Many other Australian microcionid species are also positively
known to have arcuate isochelae, including two species of Dendrocia Hallmann [D. myxilloides (Dendy) (NMV G2376), and D. pyramida (Lendenfeld) (AM GgON), both from
south-east Australia], and a number of species of Clathria [C. scabida (Carter) (BMNH
1887.7.11.9) and C. echinonematissima (Carter) (no extant type material), both from Vic.;
Fig. 10. Clathria (Clathriopsamrna) litos, sp. nov.: A, holotype QM G301269 (ORSTOM R190);
B, holotype in situ (photo P. Laboute); C, fibre structure (scale=200 pm); D, SEM of fibres
and debris incorporated (magnified 200x); E, SEM of skeletal structure (magnified 6 0 ~ ) .
1246
J. N. A. Hooper and C. L6vi
C. grisea (Hentschel) (ZMB 4435) from the Houtman-Abrolhos Islands, W.A.; C. curvichela
(Hallmann) (AM Z59) from the Gulf St Vincent, S.A., C. elegantula Ridley and Dendy
(BMNH 1887.5.2.91) from south-east and south-west Australia, C. oxyphila (Hallmann)
(AM Z51) from Tas. and S.A.]. By comparison, other obvious sister-species have palmate
isochelae but do not differ substantially in their other morphological and/or skeletal
characters, indicating that they also clearly belong to these genera [e.g. an undescribed
species of Dendrocia from the Houtman-Abrolhos Islands, W.A. (NTM Z3218), and most
species assigned to the genus Clathria]. This material cited above has been comprehensively
re-examined and it is confirmed that all nominal taxa are undeniably microcionids. Consequently, the importance of microsclere geometry at supraspecific levels of classification
must be questioned. Indeed, Hooper (1991a) indicated that the existing generic classification,
which was partially based on chelae geometry (after de Laubenfels 1936), and reiterated most
recently by Bergquist and Fromont (1988), requires substantial revision because of high levels
of homoplasy due to these and other features (the characters reappearing in many places,
inferring that they have evolved independently many times).
The present interpretation is that the distinction between palmate versus arcuate isochelae
is certainly not one of priority at the family level of classification (e.g. Microcionidae versus
Myxillidae; Burton 1959); and, furthermore, the differentiation of genera on the basis of
this feature must also be questioned [encompassing many of de Laubenfels' (1936) genera].
These questions, and the high levels of homoplasy in the classification resulting from their
continuous use, have already been posed by Hooper (1991a) and this topic must now be
investigated more seriously (see also comments noted above for Myxillidae and Acarnus).
In any case, apart from the possession of arcuate isochelae, Clathria litos shows no close
similarities to any of the above mentioned species. In external morphology it resembles
to some extent the tropical Australian Clathria (Isociella) eccentrica (Burton), and also
C. (C.) menoui, sp. nov. (see below), but this similarity probably merely reflects the
porous, reticulate nature of the surface skeleton. The precise affinities of this species
are uncertain.
Etymology
From the Greek litos (plain, simple), referring to the depauperate, virtually vestigial
nature of the spicule skeleton in this species.
Clathria (Thalysias) vulpina (Lamarck)
(Figs 11-12, Table 6)
Restricted synonymy
Spongia vulpina Lamarck, 1814: 449.
Halichondria frondifera Bowerbank, 1875: 288-89.
Clathria diche1a.-Vacelet et al., 1976: 71, pl. 3, fig. b.
Clathria corallitincta Dendy, 1889: 85, pl. 4, fig. 8.
Rhaphidophlus seriatus Thiele, 1899: 14, pl. 1, fig. 6, pl. 5, fig. 7.
Clathria reinwardti var. palmata. -Ridley, 1884: 447.
Material Examined
Holotype. MNHN DT639: ? Australia, Peron and Lesueur collection (dry).
New Caledonian material. QM G300279 (fragment NTM 23882): Baie des Citrons, off Noumea,
22°18~0'S.,166025.0'E., 3 m depth, 25.ix.1990, coll. M. Kelly-Borges, snorkel, fringing coral reef;
QM G300692 (fragment NTM 23878): stn 106, N. entrance, RBcif des Cinq Milles, SW. New Caledonia
lagoon, 22°293'S.,166044.4'E., 8 m depth, 30.iv.1976, coll. P. Laboute, SCUBA; ORSTOM R183:
stn 124, Ilbt Maitre, 22°20.1'S.,166025.1'E., 20 m depth, 15.vii.1976, coll. P. Laboute, SCUBA;
ORSTOM R264, R471: stn 110, SE. Ilbt Redika, 22O3l. 11S.,166"36.0'E., 19 m depth, 3.vi.1976,
coll. P. Laboute, SCUBA; ORSTOM R588: stn 147, mid-channel between Ton Du (Tiari) and Ilbt Ie,
22°121'S.,166016~5'E., 24 m depth, 8.ix.1976, coll. P. Laboute, SCUBA; ORSTOM R275: stn 113,
L'Cpave du 'Bonhomme', Grand Rtcif de Mbhre, 22'21 ~O'S.,166°14.0'E., 25 m depth, 21.vi.1976, coll.
P. Laboute, SCUBA.
New Caledonian Poecilosclerids
1247
Comparative material. Sri Lanka: BMNH 1889.1.21.17: Gulf of Manaar, Ceylon, 80501N.,
79"401E., coll. E. Thurston. Micronesia: USNM 23149: Agana Bay, NW. coast of Guam, 14"N.,
154OE., 2 m depth, 22.ix.1949, coll. unknown; ZMA POR769: Siboga stn 257. Philippines: USNM
21256, 21257: precise locality unknown. Indonesia: BMNH 1877.5.21.1351-2: Gaspar Straits, off
Belitung I., 3°10'S.,107015'E., coll, unknown; SMF 1699 (fragment MNHN LBIM DCL2352): Straits
of Dobo, Aru I., 6°S.,134050'E., 40 m depth, 20.iii.1908, coll. H. Merton, dredge; SMF 1673 (fragment MNHN LBIM DCL2230): same locality, 16 m depth, 20.iii.1908; NMB 16, 17 (fragments
BMNH 1908.9.24.165-166, ZMB 2897): Kema, off Minahassa, Celebes (Sulawesi), 2°50'S.,123030'E.,
30 m depth, 1895, coll. P. and F. Sarasin, dredge. Darwin region, N.T.: NTM 2974, 978: Dudley
Pt, East P t Marine Reserve, 12'24.5-25'S.,13O048-48+4'E., 10-12 m depth, 13.ix.1982, coll. J. N. A.
Hooper, SCUBA; NTM 21092, 1101: same locality, 6-7 m depth, 22.xii.1982; NTM 22691: same
locality, 9-12 m depth, 3.iv.1986; NTM 23194: same locality, 9 m depth, 16.ix.1987; NTM 21976,
1981: Weed Reef, 12°29.2'S.,130037.6-47. l1E., 8 m depth, 1l.v.1984, coll. J. N. A. Hooper, SCUBA;
NTM 22097: same locality, 23.viii.1984; NTM 22188: same locality, 16.xi.1984. NTM 2437: Lee Pt
Reef, 12"19~2'S.,130"53~1'E.,intertidal, 13.xii.1981, coll. J. N. A. Hooper; NTM 2782: Angler's
Reef, Lee Pt, 12°18.7'S.,130052.0'E., 5-6 m depth, 19.vii. 1982, coll. P. N. Alderslade, SCUBA;
NTM 22101, 2103: Fish Reef, Bynoe Harbour, 12"26~2'S.,130°26.2'E., 9-10 m depth, 5.ix.1984,
coll. J. N. A. Hooper, SCUBA. Cobourg Peninsula region N.T.: NTM 252, 54, 65: Coral Bay,
6 m depth,
Port Essington, Cobourg Pen. Marine Natl Pk, 11'09.4-11~5'S.,132"02~0-04~0'E.,
17.x.1981, coll. J. N. A. Hooper, SCUBA; NTM 288: same locality, 4 m depth, 18.x.1981; NTM
294: same locality, 5 m depth, 19.x.1981; NTM 2442: same locality, 4-5 m depth, 18.x.1981;
NTM 21354, 1375: same locality, 4-5 m depth, 16.v.1983; NTM 22515: same locality, 2-3 m depth,
l 5 . i ~1985;
.
NTM 22520: Orontes Reef, mouth of Port Essington, 11°03 .6'S.,l32"05.4'E., 18-20 m
depth, 16.ix.1985, coll. J. N. A. Hooper, SCUBA; NTM 2626: Unnamed Shoal, N. of Melville I.,
Arafura Sea, 11°36.2'S.,129052.0'E., 47 m depth, 22.v.1982, coll. R. Lockyer, SCUBA; NTM 2923:
Cootamundra Shoals, N. of Melville I., Arafura Sea, 10°50.2'S.,129"13 .2'E., 36 m depth, 7.v.1982,
coll. R. Lockyer, SCUBA. Northwest shelf, W.A.: NTM 21222: W. of Port Hedland, 19"28.5'S.,
118"55.3'E., 40 m depth, 26.iv.1983, coll. J. N. A. Hooper, CSIRO RV 'Soela', beam trawl; NTMl810:
same locality, 19°26~9'S.,l18054.2'E., 50 m depth, 30.viii.1983, coll. T. Ward, CSIRO RV 'Soela',
beam trawl; NTM 21421: NE. of Port Hedland, 19°1.0'S.,119025.0'E., 80 m depth, 19.iv.1983, coll.
R. S. Williams, pair trawl; NCI Q66C-1557-5 (fragment NTM 23401): 0.5 km E. of Hermite I.,
Monte Bello Is, 20°27.6'S.,115"32.98'E.,3 m depth, 29.viii.1988, coll. D. Low Choy and NCI,
50 m depth,
SCUBA; NTM 23043: N. of Amphinome Shoals, 19°19~7-23.3'S.,119008.8-12.2'E.,
19.vii.1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', beam trawl. Mid-west coast,
W.A.: NTM 22968: W. of Carnarvon, 24°55.6-56.5'S.,112050.8-53.5'E.,80-85 m depth, 14.vii.
1987, coll. J. N. A. Hooper, USSR RV 'Akademik Oparin', beam trawl; NTM 23373: 4 km N. of
Barrow I., 20°38.8'S.,115028.8'E., 22 m depth, 26.viii.1988, coll. D. Low Choy and NCI, SCUBA;
NTM 23397: 3 km W. of Hermite I., Small Is group, Monte Bello Is, 20'27. lfS.,115"34.2'E., 6 m
depth, 29.viii.1988; NCI Q66C-1557-5 (fragment NTM 23401): 1 km E. of Hermite I., Small Is
group, Monte Bello Is, 20°27.6'S.,115"32.8'E., 10 m depth, 29.viii.1988; NTM 23446: 2 km N. of
Bessieres I., Anchor Is, Exmouth Gulf, 21°30~6'S.,114045.4'E., 17 m depth, 23.viii.1988. Queensland:
BMNH 1881.10.21.288: Thursday I., Torres Strait, 10°35'S.,142013'E., 8-10 m depth, coll. HMS
'Alert'; Bird I., Torres Strait, Qld, 11°42'S.,143"05'E., coll. HMS 'Alert'; QM GL708: Moody Reef,
Home Is, Cape York, 12°05.6'S.,143014.5'E., 29.7 m depth, 25.ix.1979, coll. QFS, trawl; QM GL863,
1986, 2755: Green I., GBR, 16°46'S.,146058'E,, Feb. 1979-Oct. 1980, coll. L. Cannon and B. Goeden,
trawl; QM GL705: Osterland Reef, SE. of Cooktown, 15°36.2'S.,145026.8'E., 24 m depth, 10.ix.1979,
coll. QFS, trawl; QM GL710: Noggin Passage, Frankland Is, SE. of Cairns, 17"13~5'S.,146"17~0'E.,
37.8 m depth, lO.x.1979, coll. QFS, trawl; NTM 22859: Low Isles, N. of Cairns, 16°23'S.,145034'E.,
0.3 m depth, 18.x.1986, coll. A. Ayling.
Description
Colour. Vivid red-orange alive (Munsell 5R 4/10), beige in ethanol.
Shape. Numerous folded and rejoined lamellae together producing elongate-tubular,
tubulo-digitate, flabellate or sometimes simply subspherical growth forms, always with
cavernous reticulate construction; size varies from small (70 mm diameter) to massive lobes
(up to 300 mm diameter).
Surface. Many rounded conules, small digits, or spiky projections on ridges and margins
of surface, with area between conules highly excavated to produce cavernous interior; large
oscules (up to 6 mm diameter) on apex of surface projections or on margins of branches,
1248
J. N. A. Hooper and C. LCvi
usually raised above surface and with a membraneous lip. Surface fleshy alive, harder when
preserved, not noticeably porous.
Ectosome. Variably developed ectosomal skeleton, ranging from typical Thalysias
condition consisting of erect brushes of both ectosomal and subectosomal auxiliary styles
(mainly on surface conules), to Clathria condition where paratangential or tangential
bundles lie above plumose subectosomal skeleton (mainly between surface conules). Subectosomal region usually cavernous, with tracts of choanosomal styles becoming plumose
towards the peripheral skeleton, but usually not protruding through surface.
Choanosome. Skeletal architecture regularly reticulate, with well-developed spongin fibres
forming more-or-less square meshes; fibres differentiated into primary, multispicular,
predominantly ascending fibres, where spicules occupy about half of fibre diameter, and
secondary, uni- or paucispicular, mostly transverse connecting fibres; fibres cored by both
choanosomal principal styles and long, slender subectosomal styles, although the former are
most common; fibres relatively heavily echinated by acanthostyles. Mesohyl with light
collagen, relatively few free megascleres, and only rare microscleres.
Fig. 11. Clathria (Thalysias)vulpina (Lamarck):A, specimen OM G300692,
choanosomal principal styles; B, subectosomal auxiliary subtylostyles; C,
ectosomal auxiliary styles; D, echinating acanthostyles; E, larger toxas;
F, smaller toxas; G , section through peripheral skeleton.
New Caledonian Poecilosclerids
1249
Megascleres (refer to Table 6 for dimensions). Choanosomal principal styles, entirely
smooth, straight or slightly curved near base, with rounded or only faintly subtylote bases,
tapering to sharp points; similar in morphology to subectosomal auxiliary styles, although
thicker and shorter; moderately common.
Subectosomal auxiliary styles long, thin, straight, rounded smooth bases, rarely subtylote,
rarely microspined, tapering to sharp points; abundant.
Ectosomal auxiliary styles short, thin, straight, rounded smooth bases or slightly subtylote,
rarely microspined, tapering to sharp points; less common.
Echinating acanthostyles short, thin, slightly subtylote bases, sharp points, spined mostly
on basal and distal ends, with a more-or-less spine-free area below basal swelling; abundant.
Fig. 12. Clathria (Thalysias) vulpina
(Lamarck): A, specimen QM G300692
in situ (photo P . Laboute); B, SEM of
peripheral skeleton (magnified 80 x); C,
SEM of fibre characteristics (magnified
600x).
Table 6. Comparison in spicule dimensions between Clathria vulpina from New Caledonia and
populations from northern Australia
Measurements are given in micrometres. Refer to text for sources of information
Spicule type
Dimension
New Caledonia
Northern Australia
Choanosomal principal styles
L
W
L
140-(156.8)-195
5-(6.2)-8
142-(205.8)-244
2-(3.1)-5
79-(97.7)-128
2-(2.6)-4
37-(51.1)-59
2 5-(3.5)-6
18-(79.3)-194
0.5-(0.6)-0.8
Absent
Absent
113-(184.4-253
4-(11.2)-19
124-(221.4)-3 11
1-(4.7)-10
64-(99.1)-161
1-(3.3)-6.9
52-(73.1)-95
1-(6.6)-12
7-(97.9)-200
0.5-(1.2)-3.2
7-(12.4)-18
1-(3.8)-7
Subectosomal auxiliary styles
W
Ectosomal auxiliary styles
Echinating acanthostyles
Toxas
Isochelae
L
W
L
W
L
W
LI
LII
1250
J. N. A. Hooper and C. L h i
Microscleres. Isochelae absent.
Toxas always thin, hair-like, divided into 2 forms, the smaller with central curvature and
slightly reflexed arms, the larger with small central bend and straight arms; both forms
uncommon.
Distribution
Widely distributed throughout tropical and subtropical waters of the Indo-Pacific, from
Mozambique, Madagascar, Amirante Is, Seychelle Is, Aldabra Is, Red Sea, west coast of
India, Gulf of Manaar, Sri Lanka, Mergui Archipelago, Andaman Sea, Straits of Malacca,
throughout the Indo-Malay archipelago, Philippines, Vietnam, Japan, Guam, Papua New
Guinea, west, north, and eastern Australia and New Caledonia; intertidal to 80 m depth,
coral reef, coral rubble, Halimeda beds, beach rock (Hooper, unpublished data).
Remarks
This species is now known from collections made throughout the Indo-Pacific region:
Madagascar, Sri Lanka, Vietnam, Philippines, Guam, Papua New Guinea, Indonesia,
Western Australia (Shark Bay to Exmouth Gulf), northern Australia (Darwin region to
the Wessel Islands), and eastern Australia (Cape York to Cairns). The New Caledonia
population described here is a new locality record and also the most easterly record for the
species so far recorded. Not surprisingly, therefore, specimens were found to vary in a
number of features from the Indian Ocean-Australasian populations. However, these New
Caledonian specimens are still easily recognisable as belonging to C. vulpina, and this
recorded variability is not considered to warrant the creation of a new taxon.
The New Caledonia population is consistently brighter red in life than other populations
(which are typically mauve-red in shallow water, or drab brown-beige in deeper water
material). The bases of subectosomal and ectosomal auxiliary styles are predominantly
smooth (v. predominantly spined), isochelae appear to be entirely absent (v. isochelae of
two size categories present, and in only 3% of material were they rare or absent), toxas are
invariably thin, hair-like, never more than 0.8 pm diameter [v. toxa thickness varying from
very thin to relatively thick (0.5-3.2 pm diameter)]; ectosomal skeleton is thin, paratangential, relatively poorly developed (v. more substantial development of ectosome,
although specimens range from the bushy, erect condition typical of Thalysias, to paratangential as described above), and all categories of spicules are marginally smaller in the
New Caledonian specimens (Table 6 ) .
Detailed analysis of most morphological characters from large samples of the species
(Hooper, unpublished data), in an attempt to document its intra-specific variability, failed
to find any substantial, cryptic differences between widely dispersed populations, suggesting
that C. vulpina is truly a circum-Indo-Pacific species.
Clathria (Thalysias) flabellifera, sp. nov.
(Figs 13-14, Table 7)
Material Examined
Holotype. QM G300693 (ORSTOM R1416; fragment NTM 23884): stn 106, N. entrance, RCcif
des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 3O.i~.1976, coll.
P . Laboute, SCUBA.
Other New Caledonian material. QM G301271 (ORSTOM 'cfR1416'): stn 456, Lagoon pool, E.
coast Iles des Pins, 22°35.1'S.,167032.0'E., 38 m depth, 28.xi.1987, coll. J.-L. Menou, SCUBA;
ORSTOM 'cfRl416': stn 457, E. coast Ile des Pins, 22"36.0'S.,167"33.O'E., 15 m depth, 28.xi.1987,
coll. G. Bargibant, SCUBA.
Description
Colour. Orange-red alive (Munsell 5R 5/10), pale cream in ethanol.
Shape. Flabelliform, flattened in 1 plane, produced by fusion of adjacent, flattened
branches, but branch fusion incomplete producing large gaps on surface of fan; fan 170 mm
high, 230 mm wide, up to 8 mm maximum breadth, with uneven, digitate margins.
New Caledonian Poecilosclerids
1251
Surface. Mostly even, slightly folded, especially towards digitate margins; without
conules or other processes; prominent oscules visible in live specimen, approximately 2 mm
diameter, each surrounded by white subectosomal canals (astrorhizae) converging on oscules
in all directions, but oscules not visible optically in preserved material; surface microscopically porous, with 'dusty' coloured surface in preserved state.
Ectosome. Variable development of ectosomal skeleton, ranging from specialised
Thalysias condition with erect brushes of intermingled ectosomal and subectosomal auxiliary
spicules forming continuous palisade, to Clathria condition with sparse tangential or
paratangential tracts of auxiliary spicules; subectosomal region slightly cavernous, with
relatively large mesh sizes (200-350 pm maximum diameter); tips of choanosomal principal
megascleres may protrude through ectosome, but usually masked by 'true' ectosomal
skeleton.
Choanosome. Skeletal architecture regularly reticulate, composed of long, primary,
ascending spongin fibres (70-150 pm diameter), and short, thinner, secondary, transverse
fibres (40-100 pm), together producing elongate-rectangular or square meshes near periphery,
more compact oval meshes towards axis (150-200 pm diameter); axial fibres cored by
multispicular tracts of choanosomal principal spicules occupying most of fibre diameter;
secondary interconnecting fibres paucispicular; all fibres in peripheral skeleton cored by
uni- or paucispicular tracts of spicules, forming plumose brushes at surface; echinating
acanthostyles abundant throughout skeleton; collagen light, choanocyte chambers oval
(40-90 pm diameter), usually lined by abundant isochelae and toxas.
Megascleres (refer to Table 7 for dimensions). Choanosomal principal styles relatively
short, ranging from narrow to robust, usually straight, slightly subtylote or with rounded,
smooth bases, tapering to fusiform points.
Subectosomal auxiliary styles long, slender, straight, with rounded or slightly subtylote
smooth bases, tapering to sharp points.
Fig. 13. Clathria (Thalysias)jlabeNifera,sp. nov.: A, holotype QM G300693, choanosomal principal
styles; B, subectosomal auxiliary styles; C, ectosomal auxiliary styles; D, echinating acanthostyles;
E, smaller toxas; F, larger toxas; G, palmate isochelae; H, section through peripheral skeleton.
J. N. A. Hooper and C. L6vi
1252
Ectosomal auxiliary styles short, slender, straight, with rounded or slightly subtylote
bases, usually smooth, sometimes microspined, tapering to sharp points.
Echinating acanthostyles relatively robust, subtylote, sharply pointed, with small spines
located mainly on bases and points of spicules.
Fig. 14. Clathria (Thalysias)
Jabellfera, sp. nov.: A , holotype
QM G300693 (ORSTOM R1416) in situ
(photo J.-L. Menou); B, SEM of axial
skeleton (magnified 1 0 0 ~ ) C,
; SEM of
fibre characteristics (magnified 500 x ).
Table 7. Comparison in spicule dimensions between the New Caledonian Clathriaflabellifera, sp. nov.,
and the related Western Australian species Clathria cancellaria
Measurements are given in micrometres
Spicule type
-
-
Dimension
-
Choanosomal principal styles
Subectosomal auxiliary styles
Ectosomal auxiliary styles
Echinating acanthostyles
Toxas I (short, thick, recurved)
Toxas I1 (long, thin, straight)
Isochelae I
Isochelae I1
A
C. Jabellifera
(New Caledonia)
-
C. cancellariaA
(Western Australia)
-
139-(151.4-163
6-(7.6)-9
209-(253.8)-293
3-(4.4-6
98-(131.7)-179
2-(2.3-4
49-(66.7)-84
3-(5.1)-7
13-(33.4-48
1.O-(I.7)-2.5
49-(124.8)-204
O.5-(0.8)-1 .O
11-(13.1)-15
Absent
161-(185.5)-218
6-(12.6)-16
135-(218.8)-286
3-(6.5)-8.5
86-(97. 6)-142
2-(3.3)-5
62-(79.8)-92.5
3-(8.6)-11 .5
Absent
Holotype MNHN DT528 and specimen NTM 21249 from the Northwest Shelf, W.A.
New Caledonian Poecilosclerids
1253
Microscleres. Palmate isochelae simple shape.
Two categories of toxas present: (I) most common short, relatively thick, large central
curve, reflexed arms; (11) long, hair-like, with small central curve, straight arms.
Distribution
Known so far only from the south-west coast of the Noumea lagoon, 8-38 m depth,
growing on coral and coral rubble substrates.
Remarks
Clathria (Thalysias)j7abellifera is very closely related to (and possibly may even eventually
be merged with) C. (T.) cancellaria (Lamarck, 1814). Both species possess several similar
features, indicating that they are at least sibling species (i.e. similar skeletal architecture,
comparable spicule geometry and spicule size in most cases, and nearly identical growth
form, surface features and live coloration).
Clathria cancellaria was originally collected from an unknown locality thought to be on
the south-west coast of Australia-the type material of Lamarck's (1814) (MNHN DT528)
has recently been re-examined-and its Western Australian distribution was also recently
confirmed by the discovery of additional fresh material from the Houtman-Abrolhos Islands
and Northwest Shelf of Western Australia (Hooper, unpublished data). There are no known
populations of either species living in between these widely separated New Caledonian
and Western Australian populations, a fact that has been established with a reasonable
degree of confidence from recent widespread collecting efforts in tropical Australasia.
Consideration that these two disjunct populations might be recent isolates of a single species
are rejected, given that there are several major, albeit relatively cryptic differences between
the two groups. These differences are considered to justify the erection of a new taxon for
the New Caledonian population, being relatively more substantial than allowing us to
recognise it simply as a geographical 'morph' of C. cancellaria (as done above for
C. vulpina, for example). Clathria flabellifera has two different sorts of toxa microscleres
(neither of which resemble the single toxon category found in C . cancellaria), one size of
isochelae microsclere (the latter species has two), acanthostyle spines are small and found
mainly towards the extremities of the spicule (whereas those on the latter species are much
larger, the spicules more robust, and spines more evenly dispersed over the spicule), with
the exception of a few smaller ectosomal megascleres most auxiliary spicules are completely
smooth (whereas C. cancellaria has basal spines on most of the smaller and larger auxiliary
spicules), and most spicules are substantially thinner in the New Caledonian population
(Table 7).
Of course, it is possible that these differences may be trivialised in future re-evaluations
of the Microcionidae, but on the basis of morphological criteria currently accepted for this
particular group (e.g. Hooper 1991a), and for the Poecilosclerida in general (e.g. van Soest
1984a; Bergquist and Fromont 1988), and corroborated by chemotaxonomic evidence (e.g.
Hooper et al. 1990; Hooper et al. 1992), there is certainly sufficient morphological
divergence to recognise these two sibling species as distinct.
Etymology
Named for its fan-shaped growth form.
Clathria (Thalysias) corneolia, sp. nov.
(Figs 15-16, Table 8)
Material Examined
Holotype. QM G300691 (ORSTOM R527; fragment NTM 23877): stn 106, N. entrance, RCcif
des Cinq Milles, SW. New Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 30.iv.1976, coll.
P. Laboute, SCUBA.
Other New Caledonian material. ORSTOM R340: stn 114, NE. IlBt TarCti, Banc Gail, 22022.11S.,
166"39.0rE., 35 m depth, 29.vi.1976, coll. P. Laboute, SCUBA; ORSTOM R526: stn 147, mid24 m depth, 8.ix.1976, coll.
channel between Ton Du (TiarC) and Ilat Ie, 22°12~1'S.,166016~5'E.,
1254
J . N. A. Hooper and C. LCvi
P. Laboute, SCUBA; ORSTOM R475: stn 110, SE. Ilbt RCdika, 22°31~1'S.,166036~0'E.,
19 m depth,
3.vi.1976, coll. P. Laboute, SCUBA.
Description
Colour. Pale red-pink alive (Munsell 2.5R 8/4), pale pink-brown in ethanol.
Shape. Clumped, branching growth form, with bulbous subcylindrical branches or thin
arborescent branches, attached to a common base, entirely free or partially fused with
adjacent branches, and with one or more points of attachment to substrate. Growth forms
range from 70 to 190 mm high; branches bulbous-cylindrical alive up to 45 mm diameter,
contracted and slightly flattened when preserved, 5-12 mm diameter, usually bifurcated
and reticulated, tapering to rounded tips.
Fig. 15. Clathria (Thalysias) corneolia, sp. nov.: A , holotype QM G300691,
principal styles (echinating fibres); B, subectosomal auxiliary styles; C,ectosomal
auxiliary styles; D, echinating acanthostyles; E, toxas; F, two sizes palmate
isochelae; G , section through peripheral skeleton.
New Caledonian Poecilosclerids
1255
Surface. Optically even, without any noticeable protrusions; surface with membraneous
ectosome stretched between branches, most clearly seen in live specimens, partially collapsing
upon preservation, but still retaining prominent subectosomal canals (astrorhizae) running
along lateral margins of branches. Large oscules, up to 25 mm diameter, on apex of
branches, surrounded by prominent membraneous lip.
Ectosome. Specialised ectosomal skeleton present, consisting of well-developed, discrete
plumose bundles of ectosomal styles, perched on peripheral spongin fibres; ectosomal spicule
bundles not always associated with ascending fibres or spicule tracts that arise from sub-
Fig. 16. Clathria (Thalysias) corneolia,
sp. nov.: A , specimen ORSTOM R340
in situ (photo P. Laboute); B, holotype
QM (3300691 (ORSTOM R527); C , SEM
of fibre characteristics (magnified 300 x);
D, SEM of axial skeleton (magnified 8 0 ~ ) .
Table 8. Comparison in spicule dimensions between the New Caledonian Clathria corneolia, sp. nov.,
and the related species Clathria cervicornis (Thiele) from Oceania and the Indo-Malay archipelago
Measurements are given in micrometres
Spicule type
Dimension
Choanosomal principal styles
L
Subectosomal auxiliary styles
W
L
W
Ectosomal auxiliary styles
Echinating acanthostyles
Toxas
Isochelae I
Isochelae I1
L
W
L
W
L
W
L
L
C . corneolia
(New Caledonia)
C. cervicornis
(holotype SMF 679)
Undifferentiated from
subectosomal styles
180-(258.9)-304
4-(5.8)-9
112-(128.3)-155
2-(3.3)-5
48-(62.0)-74
3-(5.9-7
12-(35.3)-55
O.5-(0.9)-1.5
10-(12.4)-15
3-(6.9)-9
1256
J. N. A. Hooper and C. Ltvi
ectosomal skeleton. Peripheral spongin fibre skeleton well developed, lying directly below
ectosomal membrane. Fibres usually terminate as swollen nodes on surface.
Choanosome. Subectosomal organic skeleton relatively cavernous, with large, elongate
fibre meshes, 160-350 pm diameter; subectosomal mineral skeleton more-or-less plumose,
with tracts of subectosomal auxiliary styles, in paucispicular bundles, coring fibres and many
also protruding from fibres in plumose array. Choanosomal skeleton irregularly reticulate,
with well-developed spongin fibre system divided into primary and secondary components;
fibre reticulation in choanosomal core produces elongate meshes, 230-400 pm maximum
diameter; primary fibres paucispicular, 100-180 pm diameter, cored by subectosomal
auxiliary styles; secondary fibres 50-90 pm diameter, aspicular or cored by uni- or
paucispicular tracts of subectosomal styles. Both acanthostyles and principal styles echinate
fibres, but these are sparse. Choanocyte chambers oval, 50-85 pm diameter, often paired.
Mesohyl reinforced by light collagen, with abundant microscleres and many auxiliary spicules
also dispersed between fibres.
Megascleres (refer to Table 8 for dimensions). Principal styles (echinating fibres) scarce;
straight, sharply pointed or with stepped points (telescoped ends); bases evenly rounded
or tapering, hastate, invariably smooth. Intermediate forms between principal styles and
acanthostyles also occur.
Subectosomal auxiliary styles long, usually straight, sharply pointed, with slightly
subtylote, smooth bases.
Ectosomal auxiliary styles short, thin, straight, sharply pointed, with slightly subtylote
bases.
Acanthostyles (echinating fibres) scarce; short, stout, sharply pointed or stepped tips,
with evenly rounded or slightly subtylote bases; spines concentrated on bases and mid-way
along shaft, sometimes in more-or-less verticillate rows.
Microscleres (refer to Table 8 for dimensions). Palmate isochelae scarce, two size
categories, both small, the larger with some twisted forms.
Toxas common, moderately thin, short or long, usually with large central curvature,
with or without straight arms.
Distribution
Known so far only from the south-west coast of the New Caledonian lagoon, sand,
coral rubble, Halimeda beds.
Remarks
This species is very similar to C. (T.) cervicornis (Thiele, 1903), which is widespread
throughout the western Pacific and Indo-Malay archipelago-Marshall Is, Palau Is, Caroline
Is, Moluccas, Arafura Sea (Thiele 1903; Brondsted 1934; de Laubenfels 1954; Bergquist
1965). However, there are several cryptic but important morphological differences indicating
that two species should be distinguished. In C. cervicornis, principal styles are virtually
undifferentiated from subectosomal spicules; most spongin fibres are cored by multi- or
paucispicular columns of subectosomal styles, and there is no differentiation of primary
or secondary fibre elements; most categories of spicules are substantially longer and more
robust (Table 8); and spination of acanthostyles is also quite different.
Etymology
Named for the bulbous, horny spongin fibres predominant in the surface and choanosoma1 skeletons (L. cornu, horn).
Clathria (Thalysias) araiosa, sp. nov.
(Figs 17-18, Table 9)
Material Examined
Holotype. QM G300694 (ORSTOM R1370; fragment NTM 23886): stn 106, N. entrance, Rtcif
des Cinq Milles, SW. New Caledonia lagoon, 22"29.3'S.,166"44.4'E., 8 m depth, 30.iv.1976, coll.
G. Bargibant, SCUBA.
New Caledonian Poecilosclerids
Description
Colour. Pale red alive (Munsell 5R 6/10), beige in ethanol.
Shape. Bulbous, lobate, thickly encrusting, up to 15 mm thick.
Surface. Uneven surface composed of numerous fused bulbs, each with apical osculum,
up to 4 mm diameter, raised on membraneous lip and with white or pale red subectosomal
canals (astrorhizae). Surface also excavated by parasitic barnacles.
Ectosome. Dense ectosomal skeleton, composed of a single layer of ectosomal auxiliary
styles forming compact brushes, perched on the ends of subectosomal spicule brushes.
Choanosome. Hymedesmoid skeletal architecture, with thin layer of spongin lying on
coralline substrate; bases of choanosomal principal styles and echinating acanthostyles
embedded in basal spongin layer, standing perpendicular to substrate; multispicular brushes
of subectosomal auxiliary styles arise from ends of erect choanosomal styles, in 2-3 discrete
layers, producing straight or sinuous tracts within mesohyl, and with peripheral layer of
subectosomal spicules intermingled with ectosomal spicules to produce the ectosomal
skeleton. Choanosome slightly cavernous near substrate, with oval choanocyte chambers,
40-130 pm diameter, and granular collagen permeating mesohyl.
Megascleres (refer to Table 9 for dimensions). Choanosomal principal styles short,
slender, straight or slightly curved towards base, sharply pointed, with smooth evenly
rounded or very slightly subtylote bases.
Subectosomal auxiliary styles long, slender or relatively thick, straight, sharply pointed,
with smooth, rounded or slightly subtylote bases.
Fig. 17. Clathria (Thalysias) araiosa, sp. nov.: A, holotype QM G300694,
choanosomal principal styles; B, subectosomal auxiliary styles; C, ectosomal
auxiliary styles; D, echinating acanthostyles; E, toxas, F, palmate isochela;
G, section through peripheral skeleton.
1258
J. N. A. Hooper and C. Ldvi
Ectosomal auxiliary styles only slightly differentiated from choanosomal principal styles,
marginally thinner, straighter, with smooth, evenly rounded, non-tylote bases.
Echinating acanthostyles relatively long, slender, slightly subtylote bases, sharply pointed,
with small spines; most spines on basal end and midway along spicule, with points and areas
below basal constriction free of spines.
Microscleres (refer to Table 9 for dimensions). Palmate isochelae rare, relatively large,
single size class.
Toxas small, thin, slightly curved at centre, with straight arms, rarely with reflexed tips.
Distribution
Known only from the New Caledonia lagoon, dead coral substrate, 8 m depth.
Remarks
Clathria araiosa is unremarkable in most of its characters, resembling many other
encrusting microcionid species [e.g. C. (C.) aceratoobtusa (Carter, 1887) (with representative
specimen seen by the authors NTM 23676 from the Gulf of Thailand, the holotype having
been destroyed), C. (C.) grisea (Hentschel, 1911) (fragment of the holotype ZMB 4435),
C. (T.) aruensis (Hentschel, 1912) (holotype SMF 955), C. (T.) toxifera (Hentschel, 1912)
(with representative specimen NTM 22136 from Darwin, N.T., the holotype not yet found
in the ZMH)]. Like these species, C.araiosa has a rnicrocionid skeletal structure (Fig. 17G),
red pigmentation, and a surface sculptured by subectosomal canals (astrorhizae) (Fig. 18A).
It is distinguished from these species only by a combination of characters such as the size,
Fig. 18. Clathria (Thalysias) araiosa, sp. nov.: A , holotype QM G300694 (ORSTOM R1370) in situ
(photo G. Bargibant); B, SEM of peripheral skeleton (magnified 80 x); C, SEM fibre characteristics
(magnified 500 x).
New Caledonian Poecilosclerids
1259
Table 9. Comparison in spicule dimensions between Clathria araiosa, sp. nov., and the related species
Clathria coralliophila (Thiele) from Indonesia, Papua New Guinea and the Great Barrier Reef
Measurements are given in micrometres
C. araoisa
(New Caledonia)
C. coralliophila
(lndonesiaA)
C. coralliophila
(Great Barrier
~ e eB,f
C. coralliophila
(Papua New
~uinea~)
L
W
126-(186.6)-267
6-(7.3-10
Choanosomal principal styles
285-(347.5)-408
145-(252.8)-406
12-(13.8)-15
7-(10.5)-13
224-(345.2)-452
10-(12.4)-15
L
W
278-(339.2)-365
2.5-(4.6)-6
Subectosomal auxiliary styles
312-(390.4)-488
321-(355.4)-390
5-(7.9)-10
5-(7.3)-11
223-(347.3)-430
5-(7.2)-9
L
W
121-(142.3)-183
2.5-(3.2)-4
Ectosomal auxiliary styles
118-(140.9)-174
104-(150.1)-208
2-(3.1)-4.5
2-(3.1)-4
109-(121.5)-138
2-(3.5)-5
L
W
50-(61.8)-72
2.5-(4.1)-5
48-(56.3)-7 1
4-(5.2)-6
84-(95.0)- 105
4-(6.3)-8
76-(85.8)-95
5-(6.6)-8
L
W
12-(25.3-42
0.5-(1.1)-2
Toxas
18-(76.6)-118
0.5-(0.9)-1.2
68-(140.3)-244
0.5-(1.4)-2
32-(125.5)-211
0.5-(1.2)-2
11-(13.5)-17
10-(12.1)-15
2-(4.4)-8
3-(5.1)-8
Echinating acanthostyles
Isochelae I
L
15-(16.6)-18
11-(13.1)-16
L
Absent
2-(4.6)-8
Isochelae I1
A
Lectotype SMF 1784 from Ternate, Moluccas.
Redescription of Burton's (1934) specimen from Low Is, BMNH 1930.8.13.107.
Specimen NCI Q66C-4518-A (fragment QM G300377) from Motupore I.
geometry and ornamentation of spicules. It is closest to C.(T.) coralliophila (Thiele, 1903)
from the Moluccas, Papua New Guinea and the northern Great Barrier Reef (lectotype
SMF 1784), although C. araiosa differs from that species in the dimensions of many
categories of spicules (Table 9), in lacking basal spines on either principal or auxiliary
spicules, the pattern of spination on acanthostyles, and in lacking a smaller category of
palmate isochelae.
Etymology
From the Greek araios, thin, porous.
Clathria (Thalysias) hirsuta, sp. nov.
(Figs 19-20, Table 10)
Material Examined
Holotype. QM GL2746 (fragment NTM 21551): Cairns region, Qld, 16"56'S.,146"00'E., depth
unknown, 1982, coll. QFS (stn 7, Cairns inter-reef survey, trawl).
Paratypes. QM GL2750 (fragment NTM 21555), QM GL2754 (fragment NTM 21560): Cairns
region, Qld, 16°56'S.,146000'E., depth and date of collection unknown, coll. QFS (stn 7, Cairns
inter-reef survey, trawl).
1260
J. N. A. Hooper and C. L h i
Other Queensland material. NCI Q66C-0813-X (fragments QM G300603, NTM 23494): N. end of
Deloraine I., Whitsunday Is, GBR, 20°09'S.,149004'E., 20 m depth, 15.x.1987, coll. NCI, SCUBA;
QM G300081: Shag Rock, N. Stradbroke I., vicinity of Moreton Bay, 27°25.0'S.,153031.4'E., 15 m
depth, 5.ii.1992, coll. J. N. A. Hooper and J. Wilkinson, SCUBA.
New Caledonian material. QM G301274 (ORSTOM R1548): 'Patate Proner', Ile des Pins, S.
lagoon, New Caledonia, 22°31.2'S.,167024.5'E., 30 m depth, 10.x.1991, coll. G. Bargibant, SCUBA.
Description of typical material (Queensland waters)
Colour. Bright red alive conules (Munsell 2.5R 5/10), paler between conules, dark
brown in ethanol.
Shape. Small, erect, tubular or flattened-lobate digits, reticulate-honeycombed in
construction, 98-165 mm long, 57-68 mm wide, 25-46 mm in diameter, with large digitate,
prominently conulose margins; with or without cylindrical basal stalk, up to 9 mm long,
6.5 mm diameter (Fig. 20E).
Surface. Highly clathrous, reticulate, Echinoclathria-like surface, consisting of numerous
small or large interconnected flattened or lobate branches, up to 15 mm long, with rounded
or pointed conulose edges (Fig. 20F). Surface membraneous between conules, and shaggy
on conules. Branches forming 'body' of sponge are completely fused although perforated by
large cavities, and adjacent branches are joined by a membraneous covering; porous meshes
between adjacent branches up to 4.5 mm diameter, excavating sponge entirely. Texture
firm, compressible, difficult to tear, slightly arenaceous. Large oscules, 3-5.3 mm diameter,
scattered between digitate surface projections; small pores, 0.5-0.8 mm diameter dispersed
over surface between conules.
Ectosome. Ectosomal skeleton lightly arenaceous, with a sparsely dispersed, thin,
tangential or paratangential, uni- to paucispicular skeleton composed of subectosomal
auxiliary subtylostyles lying on or just below surface; at tips of conules the surface skeleton
is raised to form irregular paratangential tracts which project through the surface.
Choanosomal principal styles also pierce surface in places on the ends of conules, occurring
individually or in light bundles, but these are relatively infrequent. Choanosomal fibres are
immediately subectosomal, with only a small subectosomal region, and the extra-fibre spicule
tracts composed of subectosomal auxiliary styles are confined to peripheral skeleton.
Choanosome. Choanosomal skeletal architecture irregularly reticulate, with thick,
flattened, relatively heavy spongin fibres divided into primary and secondary components.
Primary fibres, 58-95 pm diameter, contain multispicular tracts of intermixed choanosomal
principal styles and subectosomal auxiliary styles. Primary tracts relatively straight, ascending
to the surface (running longitudinally through sections), although near periphery they curve
towards surface and terminate in light plumose brushes; secondary fibres relatively short,
thick, 15-85 pm diameter, interconnecting primary fibres, with a uni- or paucispicular
core of intermingled principal and subectosomal spicules. Secondary fibres run mostly
perpendicular to surface, and predominant in deeper regions of choanosome. Echinating
acanthostyles moderately heavy on all fibres, particularly on those near periphery. Circular,
thick fibre nodes on all major fibre junctions, 60-98 pm diameter, contain a heavier core
of choanosomal mpgascleres than in other parts of fibres. Branching of spongin fibre
produces variable ovoid or elongate meshes, usually small, sometimes cavernous, 49-210 pm
diameter. Few spicules or detritus scattered between fibres. Abundant, unpigmented, granular
collagenous spongin in mesohyl, with oval choanocyte chambers, 42-57 pm diameter.
Megascleres (refer to Table 10 for dimensions). Choanosomal principal styles thin,
fusiform, straight, rounded or slightly subtylote, smooth bases; not well differentiated from
auxiliary subtylostyles, although shorter and thicker, with less well-developed basal swelling.
Subectosomal auxiliary subtylostyles long, slender, fusiform, straight, evenly cylindrical,
rounded or subtylote bases, usually microspined.
Acanthostyles straight, prominently subtylote, with light or vestigial spination, mostly
aspinose necks.
New Caledonian Poecilosclerids
1261
Microscleres (refer to Table 10 for dimensions). Palmate isochelae not common
(holotype and paratype GL2750), or common (other material), small, imperfectly divided into
two size categories.
Toxas very thin, raphidiform, abundant, very slight and gradually rounded central
curvature, with straight arms or very slightly reflexed points, rarely sinuous.
Description of Atypical Material (New Caledonian waters)
Colour. Tips of protruding surface conules red alive (Munsell 5R 5/10), honeycombed
portion of sponge pale mauve-red alive (2.5R 8/2), with slightly striated, paler surface
sculpturing (subectosomal canals (astrorhizae)) also visible in between surface conules; sponge
evenly beige in ethanol.
Fig. 19. Clathria (Thalysias) hirsuta, sp. nov.: A , New Caledonian specimen QM
G301274, choanosomal principal styles; B, subectosomal auxiliary styles; C, ectosomal
auxiliary styles; D, echinating acanthostyles; E, toxas; F, Queensland paratype QM
GL2754, choanosomal principal styles; G , subectosomal auxiliary subtylostyles; H,
echinating acanthostyle, I, toxas, J, palmate isochelae; K, specimen QM G301274,
section through peripheral skeleton.
1262
J. N. A. Hooper and C. Livi
Shape. Erect flattened digit, 65 mm long, 40 mm wide, 30 rnm thick, with honeycombed construction, excavated by pores and canals passing completely through sponge,
with highly conulose surface (Fig. 200, G).
Surface. Numerous close-set, pointed surface conules, 2-4 mm apart, 3-8 mm long,
2-3 mm diameter, usually bifurcate or trifurcate at apex, situated on ends of erect flat,
lamellate surface processes, 5-7 mm long, 5-9 mm wide, up to 2 mm thick, which encircle
edges of the excavated canals and rejoin to produce the honeycombed surface structure.
Texture firm, compressible, difficult to tear. Oscules, up to 5 mm diameter, scattered over
surface between microconules.
Ectosome. Ectosomal skeleton consists of sparse tangential or paratangential tracts
of intermixed ectosomal and subectosomal auxiliary styles, not producing a specialised
Thafysias-like skeleton. Collagen in ectosomal region only lightly developed.
Choanosome. Skeleton dominated by well-developed, horny spongin fibres, with mineral
skeleton greatly reduced in comparison with Queensland material. Spongin fibre skeleton
Fig. 20. Clathria (Thalysias) hirsuta, sp. nov.: A , holotype QM GL2746, SEM acanthostyles
; SEM skeletal structure
(magnified 1 0 0 0 ~ )B,
; SEM ectosomal spicule brushes (magnified 1 5 0 ~ ) C,
(magnified 1 0 0 ~ )D,
; specimen QM (3301274 (ORSTOM R1548) from New Caledonia; E, holotype
(QM GL2746) from Cairns region; F, surface of specimen QM G300603 from Great Barrier Reef (photo
NCI); G, specimen QM G301274 (ORSTOM R1548) from New Caledonia in situ (photo P. Laboute).
New Caledonian Poecilosclerids
1263
differentiated into primary, ascending fibres, 65-110 pm diameter, cored by multispicular
tracts of spicules; secondary, transverse fibres shorter and marginally thinner, 80-170 pm
long, 20-50 pm diameter, interconnecting ascending fibres, cored by pauci- or unispicular
tracts of megascleres. All fibres cored by abundant subectosomal auxiliary styles intermixed
with fewer principal choanosomal styles, and echinated by sparse echinating acanthostyles.
Coring spicules occupy only small proportion of fibre diameter, usually confined to centre
of fibre. Fibre reticulation forms triangular, oval or elongate meshes, more compact in
axial region (45-110 pm diameter) than in peripheral skeleton (90-180 pm diameter).
Mesohyl with abundant collagen, slightly granular, lightly pigmented, but with few interstitial
spicules. Choanocyte chambers large, oval, 25-50 pm diameter.
Megascleres (refer to Table 10 for dimensions). Choanosomal principal styles (exclusively
in fibres) scarce, very slender, straight, sharply pointed, with smooth evenly rounded bases.
Subectosomal auxiliary styles (both coring fibres and in the subectosomal skeleton) long,
very slender, straight or rarely slightly curved, with tapering sharp points, and bases nearly
anisoxeote, tapering to pointed or small rounded ends.
Ectosomal auxiliary styles short, straight, very slender, with evenly rounded bases.
Echinating acanthostyles scarce, short, slender, sharply pointed, with vestigial spines and
very slight subtylote bases.
Microscleres (refer to Table 10 for dimensions). Isochelae absent.
Toxas uncommon, raphidiform, with slightly angular or rounded central curvature and
straight arms.
Table 10. Comparison in spicule dimensions between Queensland and New Caledonian populations of
Clathria hirsuta, sp. nov.
Measurements are given in micrometres
Holotype
(QM GL2746)
Paratypes
(QM GL2750, 2754)
Other material
Queensland
New Caledonia
(n = 1 )
(n = 2)
Choanosomal principal styles
132-(158.9)-195
140-(158.8)-183
3.5-(6.2)-10
4-(6.0-9
Subectosomal auxiliary styles
183-(237.9-293
141-(202.2)-254
2-(3.6)-5
2-(3.5)-4.5
Ectosomal auxiliary styles
Absent
Absent
Absent
Echinating acanthostyles
43-(63.2)-79
3-(5.2)-8
52-(63.8)-75
3-(5.1)-8
Toxas
35-(100.7)-163
0.5-(0.7)-1.0
22-(104.6)-165
0.5-(0.8)-1 .O
Isochelae I
3-(4.3-6
3-(3.8)-5
Absent
9-(11.2)-14
Absent
Isochelae I1
9-(10.8)-14
1264
J. N. A. Hooper and C . L6vi
Distribution
Northern, central and southern inshore coastal waters of Queensland, and the New
Caledonia lagoon; coral reef, rock and coral rubble substrates, 20-30 m depth.
Remarks
Two independent descriptions of this species are given in order to highlight the differences
between Queensland and New Caledonian populations. Although these differences are not
considered here to be sufficient to recognise two distinct, sibling species, many characters
seen in New Caledonian material have been modified to a greater or lesser degree, or have
been lost completely, from those present in Australian populations. We consider that the
Queensland morph is typical of the species, whereas the loss or reduction of characters seen
in New Caledonian material, which is obviously secondary, is an apomorphy for the species.
Hence, the New Caledonian populations are interpreted as being derived sister-species of the
Queensland populations. Nevertheless, both populations are clearly conspecific, as seen by
their growth forms (spiky surface, vaguely digitate, reticulate-honeycombed construction),
live coloration (darker red conules, paler red or whitish between conules, with subectosomal
striations) and skeletal architecture (particularly the predominance of spongin fibres in the
skeleton, with well-differentiated primary multispicular and secondary paucispicular fibres),
whereas they are contrasted substantially in their spicule geometries (compare Fig. 19A-E,
F-J) and spicule sizes (Table 10) to the extent that comparison of these features alone
may not be enough to recognise conspecificity. Additional, non-morphological evidence is
required to determine the degree to which these two populations have diverged, but for the
present they are treated as conspecific.
In comparison with New Caledonian material, Queensland specimens have much more
robust spicules in all categories, and their relative abundance in the skeleton also differs
considerably: choanosomal principal styles are abundant inside fibres; there is only one size
class of auxiliary spicule (lacking specialised ectosomal megascleres) which is a large,
subtylote auxiliary style; acanthostyles are subtylote and relatively heavily spined; two size
classes of isochelae are present; and toxas are similar in thickness and central curvature,
but have slightly reflexed points.
Clathria hirsuta belongs to a species-group having vaguely similar skeletal characteristics,
but are generally morphologically depauperate in comparison with many other Clathria
species. To this group belong Clathria (ThaZysias) cactiformis (Lamarck) (lectotype MNHN
DT580), C. (T.) clathrata, sensu Topsent (not Schmidt nor Whitelegge) (no material seen),
C. (T.) cervicornis (Thiele) (holotype SMF 679), C. (T.) juniperina (Lamarck) (lectotype
MNHN DT570), C. (T.) paucispina (Lendenfeld) (lectotype AM G9121a), and the present
species. All these species, termed here the 'juniperina' species complex, and distributed
amongst the nominal genera Clathria and Thalysias, have choanosomal (coring) megascleres
that are barely differentiated from subectosomal auxiliary spicules (and in the present species
both are intermingled within the fibres), an irregular, heavy fibre skeleton, and raphidiform
toxas, but each species differs in one or more other significant features. In the case of the
present species, characteristics that distinguish it from other members of the 'juniperina'
species-group are its growth form, the possession of vestigially spined acanthostyles, very
thick, relatively flattened fibres, relatively thin megascleres of all categories, two sizes of
isochelae, and slightly curved, reflexed raphidiform toxas.
Hallmann (1912) contends that some of these species (i.e. those he suggested were related
to C. paucispina) are barely different in their gross morphology (i.e. more-or-less tubular,
digitate, with a spiky surface), and therefore should be combined, but resolution of this
species complex is not possible without much more comparative (preserved) material and
corroboration using non-skeletal characters. Whether or not the 'juniperina' species group
is a true monophyletic group has yet to be decided, requiring rigorous investigation of the
relative importance, and phylogenetic value, of the morphological characters concerned.
Etymology
The species name refers to the highly conulose, shaggy surface.
New Caledonian Poecilosclerids
Clathria (Clathria) kylista, sp. nov.
(Figs 21-22, Table 11)
Material Examined
Holotype. QM G300035: Inner Gneering Shoals, off Mooloolaba, SE. Qld, 26°38.5'S.,153009.5'E.,
10 m depth, 10.xii.1991, coll. J. N. A. Hooper and S. D. Cook, rock reef.
Paratype. QM G300690 (ORSTOM R1338; fragment NTM 23876): stn 106, N. entrance, Rkcif
des Cinq Milles, SW. New Caledonia lagoon, 22"29.3'S., 166"44.4'E., 8 m depth, 3O.i~.1976, coll.
G. Bargibant, SCUBA.
Description
Colour. Dark orange to pale orange alive (Munsell 10R 6/10-2.5YR 7/8), dark brown
to beige in ethanol.
Shape. Growth form ranges from erect, branching, bulbous, digitate tubes, with several
tubes attached to a common base, some bifurcating some single and each terminating with
a single osculum (paratype), to irregular more-or-less tubular lobes and low digits arising
from a bulbous, thickly encrusting base, with oscules scattered over the apical regions of
lobate digits (holotype). Specimens range from about 130 mm basal diameter up to
encrusting individuals exceeding 320 mm basal diameter, and lobate digits range from 20 mm
long, 25 mm diameter, to over 300 mm long, 70 mm diameter.
Surface. Prominently conulose, large, bulbous surface tubercles, producing regular
(paratype) or irregular (holotype) surface sculpturing. Oscules large, up to 35 mm diameter,
surrounded by a prominent membraneous lip.
Ectosome. Membraneous, with a sparse, tangential layer of irregularly dispersed subectosomal auxiliary styles and raphidiform spicules, presumably younger forms of these
spicules; ectosomal skeleton also contains erect brushes of auxiliary spicules scattered
Fig. 21. Clathria (Clathria) kylista, sp. nov.: A , paratype QM G300690, choanosomal
principal styles; B, subectosomal auxiliary subtylostyles; C, smooth echinating subtylostyles;
D, toxas; E, section through peripheral skeleton.
1266
J. N. A. Hooper and C. L6vi
throughout both peripheral and subectosomal regions, generally protruding towards
exterior. In peripheral region mesohyl contains moderately light collagen and few small
sand particles.
Choanosome. Skeletal structure plumoreticulate, with differentiated primary and
secondary skeletal lines. Primary fibres long, ascending, 50-70 pm diameter, paucispicular,
with never more than 4-5 spicules abreast; secondary fibres shorter, 15-30 pm diameter,
transverse, connecting primary fibres, uni- or aspicular. Primary fibres cored predominantly
by principal spicules but also with moderate quantities of longer, more slender auxiliary
megascleres interdispersed; spicules in primary fibres occupy only small proportion of fibre
diameter. Echinating styles moderately common on primary ascending fibres, sparse on
secondary connecting fibres, inserted at oblique angles to fibres; this uneven distribution
of both coring and echinating spicules gives skeleton an almost plumose appearance.
Choanosomal mesohyl contains sparse collagen, fibre meshes widely spaced, 160-350 pm
diameter, choanocyte chambers oval, 35-60 pm diameter, and small quantities of sand
grains scattered between fibre meshes.
Megascleres (refer to Table 1 1 for dimensions). Choanosomal principal styles slender,
straight, relatively short, with hastate points and smooth, slightly swollen, subtylote bases.
Subectosomal auxiliary styles long, slender, straight, with hastate points and smooth,
elongated, swollen subtylote bases.
Echinating styles entirely smooth, short, slender, sharply pointed, with prominent basal
constriction ('neck'), subtylote base and widest just below basal constriction.
Microscleres (refer to Table 11 for dimensions). Isochelae absent.
Toxas abundant, moderately long, ranging from slender to raphidiform, with straight
arms and prominent central curve.
Clathria (Clathria) kylista,
sp. nov.: A , paratype QM G300690
(ORSTOM R1338) in situ (photo
G. Bargibant); B, SEM of skeletal
structure (magnified 80 x); C , SEM
fibre characteristics (magnified 500 x).
Fig. 22.
New Caledonian Poecilosclerids
1267
Distribution
Although this species is known only from the Noumea lagoon and Sunshine coast region
o f Queensland, it is hypothesised that it also lives on the southern Great Barrier Reef region
and some other subtropical western Pacific islands; depth range 8-10 m .
Remarks
Under previous schemes for the Microcionidae (e.g. de Laubenfels 1936), this species
would probably have been referred to Echinoclathria because its echinating spicules are
entirely smooth. However, as noted elsewhere (Hooper 1991a), Echinoclathria differs from
Clathria not in the presence or absence o f spines on these spicules, but in the structure
(or derivation) o f spicules localised to particular parts o f the skeleton: in Echinoclathria,
spongin fibres are both cored and echinated by principal spicules (i.e. spicules with identical
geometry), whereas in Clathria there are distinct categories o f coring (principal) and
echinating megascleres.
Clathria kylista is remarkable in its growth form (Fig. 22A), skeletal architecture
(Fig. 22B-C) and spiculation (Fig. 21A-D). It superficially resembles a Callyspongia
(Haplosclerida) in its tubular digitate morphology, although it lacks the specialised
surface sculpturing characteristic o f that genus. Its skeletal structure o f sparsely cored
ascending primary fibres and uncored secondary connecting fibres is similar to that o f
C. (C.) angulifera Dendy (holotype NMV G2286), which has acanthose echinating spicules
and isochelae, and an undescribed species from South Australia (Clathria provisional species
number 503) which has acanthose echinators, isochelae and slightly curved toxas. The
possession o f entirely smooth echinating styles with basal constrictions, as well as long
slender toxas with prominent central curvature is reminiscent o f similar spicules found in
C. dubia (Kirkpatrick) (holotype BMNH 1898.12.20.37 from Christmas Island, re-examined).
This latter species has a specialised ectosomal skeleton (placing it with Thalysias), a
hymedesmoid skeletal structure, and anchorate isochelae. The unusual geometry o f the
echinating styles in C. kylista is the strongest apomorphy amongst other species o f Clathria.
Only two specimens are so far known for C. kylista, one from Queensland and one
from New Caledonia, and both o f these are very similar in their morphology and skeletal
characters. Several characters have been contrasted between this material in the description
above, and comparisons in spicule dimensions are given in Table 11.
Etymology
From the Greek kylistos, rolled, alluding to the tubular bulbous growth form peculiar
to this species.
Table 11. Comparison in spicule dimensions between New Caledonian and Australian specimens of
Clathria kylista, sp. nov.
Measurements are given in micrometres
Spicule type
Choanosomal principal styles
Subectosomal auxiliary styles
Ectosomal auxiliary styles
Echinating acanthostyles
Toxas
Isochelae
Dimension
New Caledonia
(paratype
QMG300690)
South-east Queensland
(holotype
QMG300035)
L
W
L
W
L
W
L
W
L
W
L
61-(81.6)-97
1.6-(2.3-3.6
138-(159.4)-183
0.4-(2.1)-3.5
Absent
64-(84.7)-112
2.5-(2.9)-4.0
143-(168.9)-181
1.0-(2.4)-4.1
Absent
32-(35.5)-39
1.1-(3.1)-5.0
65-(135.6)-266
0.2-(1 .I)-1 -6
Absent
26-(32.8)-40
1 .O-(2.6)-4.0
65-(129.6)-231.4
0.5-(I .3)-2.5
Absent
J. N. A. Hooper and C. Ltvi
1268
Clathria (Clathria) bulbosa, sp. nov.
(Figs 23-24, Table 4)
Material Examined
Holotype. QM G300021 (ORSTOM R1350): stn 423, E. rtcif Boody, Chesterfield Is (territory
of New Caledonia), 20°58.1'S.,158034.6'E., 42 m depth, 23.vii.1984, coll. G. Bargibant, SCUBA.
Description
Colour. Bright red alive (Munsell 5R 4/10), pale beige in ethanol.
Shape. Thickly encrusting on plate coral (Astreopora sp., Family Acroporidae), bulbous,
mushroom-shaped sponge (40 mm high, 60 mm diameter), with short, slightly constricted
basal attachment (10 mm long, 25 mm diameter), and two large oscular atria on apex of
sponge (10-18 mm diameter).
Surface. Few rounded irregular lobate bumps on lateral sides, whereas surface more even
on apex; entire surface microscopically porous. Oscular atria surrounded by symmetrical
membraneous lip. Large openings of excurrent canals (true oscules, 2-5 mm diameter) visible
inside central oscular atrium. Texture soft, compressible, difficult to tear.
Ectosome. Ectosomal skeleton with a special category of quasidiactinal auxiliary styles
erect or lying tangential to surface, usually not projecting from it, forming sparse bundles
or scattered individually on surface; these spicules confined to peripheral skeleton and
do not occur deeper within choanosomal region. Light collagenous spongin in mesohyl of
peripheral skeleton, with sparsely dispersed microscleres.
Fig. 23. Clathria (Clathria) bulbosa, sp. nov.: A, holotype QM G300021,
choanosomal auxiliary subtylostyles; B, subectosomal auxiliary quasidiactinal
styles; C, echinating acanthostyles; D, toxas; E, palmate isochelae; F, section
through peripheral skeleton; G, fibre characteristics.
New Caledonian Poecilosclerids
1269
Choanosome. Skeletal architecture reticulate, with a well developed organic spongin fibre
system predominant over inorganic spicule components. Fibres differentiated into primary
and secondary elements. Primary fibres (45-70 pm diameter) generally ascending, usually
cored by plumose paucispicular brushes of auxiliary styles (different from those on
ectosome). These spicule brushes not completely confined to axis of fibres, occasionally
bundles lie outside, adjacent to primary fibres. Secondary fibres (12-35 pm diameter) usually
transverse connecting, mostly uncored or rarely with single auxiliary styles lying in axis of
fibre. Both primary and secondary fibres heavily echinated by acanthostyles, heaviest at
fibre nodes. Fibre meshes oval to elongate, 70-350 pm diameter, more cavernous in
choanosome than in peripheral region. Mesohyl heavily permeated by collagenous spongin
containing abundant rnicroscleres. Choanocyte chambers oval (48-60 pm diameter), often
lined by microscleres.
Megascleres. 'True' principal megascleres absent; fibres cored by choanosomal auxiliary
styles, long, slender, straight, with smooth subtylote bases and hastate points: length
124-(167.6)-180 pm, width 2-(2.8)-4 pm.
Subectosomal auxiliary spicules quasidiactinal modifications of styles, long, slender,
straight, with smooth subtylote bases and smooth rounded or slightly swollen tips, often with
a terminal mucronate point: length 184-(196.8)-207 pm, width 1.5-(2.3)-3.0 pm.
Second category of specialised ectosomal spicules absent.
Echinating acanthostyles slender, straight, cylindrical, with slightly subtylote bases,
rounded or slightly swollen tips, often with terminal spines obvious; granular spines evenly
dispersed over entire spicule: length 55-(66.9)-74 pm, width 2-(2.9)-4 pm.
Microscleres. Palmate isochelae abundant, relatively large: length 14-(15.9)-19 pm.
Toxas abundant, slender, relatively short, usually with widely curved centres and slightly
reflexed arms, sometimes with straight arms: length 32-(56.3)-92 pm, width 0-8-(1.4)2 - 5 pm.
Fig. 24. Clathria (Clathria) bulbosa,
sp. nov.: A , holotype QMG300021
(ORSTOM R1350) in situ (photo
G. Bargibant); B, SEM of skeletal
structure (magnified 150 x ); C, SEM
of fibre characteristics (magnified 800 x).
1270
J. N. A . Hooper and C. L h i
Distribution
Known so far only from the Chesterfield Islands Group, coral reef habitat, 42 m depth.
Remarks
This species is obviously closely related to C. rugosa, although a number of characters
indicate that they are not conspecific but probably sibling species. Clathria rugosa is
characteristically flabellate-digitate (Fig. 8C-F), whereas C. bulbosa is bulbous encrusting
(Fig. 24A). Both species have quasidiactinal auxiliary spicules forming the inorganic part of
the ectosomal skeleton, but in C. bulbosa these auxiliary megascleres are completely smooth.
Echinating acanthostyles are also similar between the two species, with a tapering cylindrical
shape, rounded points and evenly distributed granular spines, and spicule sizes are similar
for several categories of spicules but different for others (Table 4).
Nevertheless, unlike C. rugosa, this species lacks principal spicules. Fibres are cored by
auxiliary styles: i.e. there are two categories of auxiliary styles, those coring the spongin
fibres differing only slightly in shape and dimension from the quasidiactinal forms located
in the ectosomal skeleton, but both forms appear to be derived from the auxiliary category
of spicules. Both species have reticulate skeletons and differentiated primary and secondary
spongin fibre systems, but the skeleton of C. rugosa contains abundant detritus, placing it
in the subgenus Clathriopsamma, whereas C. bulbosa lacks any foreign spicules or detritus
and is typical of most other Clathria (and included in the subgenus Clathria). These minor
differences between obvious sibling species, each of which on paper is referrable to a
different nominal genus, illustrates the possible artificiality of the genus Clathriopsamma,
as already suggested by Hooper (1991~). However, all these subgeneric groups (e.g.
Clathriopsamma, Clathria, Thalysias) do appear to contain homogeneous assemblages of
sister-species, with possible, but as yet undetermined, monophyletic origins. Interpretations
of these levels of divergence, and indeed the whole phylogenetic basis for the existing
systematics of Microcionidae, require investigation.
Etymology
From the Latin, bulbosus, swollen, referring to the bulbous encrusting growth form of
this species.
Clathria (Clathria) menoui, sp. nov.
(Figs 25-26)
Material Examined
Holotype. Q M G301267 (ORSTOM R1535): stn 490, Ilot N'Da, S. lagoon, New Caledonia,
22°41.5'S.,166058.VE.,47 m depth, 23.v.1991, coll. J.-L. Menou, SCUBA.
Description
Colour. Orange-brown alive (Munsell5YR 7/10), with bright yellow larvae (2.5Y 8/10);
light beige in ethanol.
Shape. Massive, encrusting on coral, up to 40 mm high, 60 mm wide, and extending
for 180 mm along substrate, composed of fused, bulbous (in life) or flattened flabellate
(when preserved), lobate-digitate projections, 15-35 mm high, 21-53 mm wide, 3-6 mm
thick (preserved material), attached to a common encrusting base. Smaller bulbous (live) or
flattened (preserved) conulose digits cover apex of larger digits, 4-5 mm diameter, up to
5 mm high.
Surface. Fleshy, porous, membraneous ectosome when alive, becoming detached in
most places when preserved, leaving highly porous surface. Margins of digits longitudinally
striated, uneven, and apex of most digits have single large oscule, 2-6 mm diameter, with
prominent membraneous lip. Each oscule surrounded by converging subectosomal canal
system (astrorhizae), with large canals covered by smooth membraneous surface (in contrast
to adjacent surface which is highly porous, excavated by large ostia, up to 0.8 mm
New Caledonian Poecilosclerids
1271
diameter); subectosomal canals collapse when preserved, leaving an excavated, highly porous
surface. Texture spongy, elastic, difficult to tear.
Ectosome. Membraneous, with a thin layer of subectosomal auxiliary subtylostyles
lying tangential to surface, through which protrude tips of spongin fibres from primary,
ascending spongin fibre system; terminal fibres tapering to sharp points, single or bifurcate
at apex, 50-100 pm apart, protruding approximately 50 pm through surface, and each with
a sparse paucispicular core of vestigial spicules which protrude through end of fibres.
Collagen in peripheral skeleton abundant but very lightly pigmented, with small quantities
of detritus scattered throughout (although not embedded in fibres).
Choanosome. Skeletal structure irregularly reticulate, cavernous, with spongin fibre
system dominant over spicule skeleton. Spongin fibres poorly differentiated into primary and
secondary categories; primary fibres more-or-less ascending, 40-60 pm diameter, cored
by uni- or paucispicular tracts of principal subtylostyles, sometimes uncored or cored by
vestigial spicules of indeterminant origin (merely resembling fibre pith), and coring spicules
occupy very small proportion of fibre diameter; secondary fibres 15-35 pm diameter,
interconnecting primary fibres at irregular intervals, more-or-less transverse, usually uncored
or rarely unispicular; fibre meshes irregular, generally large, oval or elongate, 80-250 pm
diameter; echinating spicules abundant, regularly dispersed over fibres. Collagen in axial
Fig. 25. Clathria (Clathria) rnenoui, sp. nov.: A, holotype QM G301267,
smaller choanosomal principal subtylostyles; B, larger subectosomal auxiliary
subtylostyles; C , smooth echinating subtylostyles;D, toxas; E, section through
peripheral skeleton.
1272
J. N. A. Hooper and C. Ltvi
region of skeleton abundant but very lightly pigmented, with numerous subectosomal
auxiliary subtylostyles scattered in fibre meshes; choanocyte chambers oval, 25-45 pm
diameter.
Megascleres. Choanosomal principal spicules, coring spongin fibres, nearly vestigial,
always with blackened axial canals, and barely different from auxiliary spicules, consisting
of short, straight, slender subtylostyles, with prominent tylote bases and sharply pointed tips.
Length 82-(91.2)-96 pm, width 2-(2.6)-3.5 pm.
Subectosomal auxiliary subtylostyles, in ectosomal skeleton and scattered between fibre
meshes, long, slender subtylostyles, straight or very slightly curved near tip, with prominent
subtylote bases, often tear-drop shaped, and long, tapering, sharply pointed tips. Length
154-(172.6)-185 pm, width 1 8-(2.5)-3 a0 pm.
Echinating spicules entirely smooth, straight subtylostyles, with well-marked constricted
'necks', evenly rounded bases and sharply pointed tips. Length 27-(30.1)-34 pm, width
1.5-(2.1)-3.0 pm.
Microscleres. Isochelae absent.
Toxas uncommon, exceedingly slender (raphide-like), with only slight angular curvature
at centre and straight arms (not reflexed). Length 62-(76.3)-85 pm, width 0.5-(0.7)0.8 pm.
Clathria (Clathria) menoui, sp. nov.: A , holotype QM G301267 (ORSTOM R1535),
peripheral spongin fibres (scale=200 pm); B, holotype; C, SEM of skeleton (magnified 80x);
D, holotype in situ (photo J.-L. Menou); E, SEM fibre characteristics (magnified 1OOOX).
Fig. 26.
New Caledonian Poecilosclerids
Distribution
Known so far only from the southern New Caledonian lagoon, coral reef habitat,
47 m depth.
Remarks
This species is greatly reduced in its inorganic skeletal characters. The species is distinctive
in its porous, ornamented surface features, live coloration, skeletal characteristics where the
organic fibre skeleton dominates over the coring spicule skeleton, the smooth echinating
spicules and the raphide-like toxas with only a slight angular centre and completely straight
arms.
In growth form and surface characteristics, C. menoui resembles to some extent the
common tropical Australian microcionid species C. (Isociella) eccentrica (Burton), but this
resemblance is purely superficial. In its fibre characteristics (well-developedhorny fibres, few
coring spicules; Fig. 26A, C, E), C. menoui also resembles the southern Australian species
Echinoclathria leporina (Lamarck), and in the geometry and ornamentation of its echinating
spicules (smooth, prominent bases, constricted 'necks', thickest at the centre of the shaft;
Fig. 25C) it shows some similarities to C.(T.) transiens Hallmann. However, none of these
species are closely related and the specific affinities of C. menoui are uncertain.
Etymology
Named for M. Jean-Louis Menou, ORSTOM, the original collector of the holotype, in
recognition of his substantial contributions to the documentation of the marine biology of
New Caledonia.
Genus Echinochalina Thiele
Echinochalina Thiele, 1903: 961 [type species Ophlitaspongia australiensis Ridley, 1884: 4421.
Protophlitaspongia Burton, 1934: 562 [Siphonochalina bispiculata Dendy, 1895: 2461.
Diagnosis (emended)
Microcionidae with monactinal, quasi-monactinal or thin diactinal auxiliary megascleres
coring irregularly reticulate or plumo-reticulate spiculo-spongin skeleton, with smooth or
acanthose principal spicules echinating fibres; ectosomal skeleton also with auxiliary spicules,
usually tangential or erect on surface; microscleres present or absent.
Remarks
Including the three New Caledonian species described here, there are now 20 nominal
species of Echinochalina worldwide, 18 of which appear to be valid, and 16 of which are
endemic to the Australasian region. The genus is divided into two subgenera: Echinochalina,
with diactinal, quasimonactinal or monactinal auxiliary spicules coring fibres, and monactinal
principal spicules echinating fibres; and Protophlitaspongia with diactinal or quasimonactinal
auxiliary spicules coring fibres and diactinal or quasimonactinal principal spicules echinating
fibres. This subdivision is based on an earlier phylogenetic analysis of the family (Hooper
1991~).
Echinochalina (Echinochalina) intermedia (Whitelegge)
(Figs 27-28, Table 12)
?Echinoclathria viminalis (in part). -Whitelegge, 1901: 87-8.
Not Thalassodendron viminalis Lendenfeld, 1888: 225.
Echinoclathria intermedia (in part). -Whitelegge, 1902: 214-15; Dendy, 1922: 7 1, pl. 2, fig. 8.
Echinochalina intermedia.-Hallmann, 1912: 294-95, text-fig. 69; Shaw, 1927: 427; Burton, 1934:
563.
Material Examined
Holotype. AM, Lendenfeld no. 365, presently missing.
J. N. A. Hooper and C. LBvi
1274
New Caledonian material. QM G300684 (ORSTOM R1299, fragment NTM 23868): stn 106, N.
entrance, RCcif des Cinq Milles, SW. New Caledonia lagoon, 22'29.3'S., 166"44.4'E., 8 m depth,
30.iv.1976, coll. P. Laboute, SCUBA; ORSTOM R433: stn 104, NE. RCcif des Cinq Milles, 22"27.5'S.,
166"43.5'E., 12 m depth, 29.iv.1976, coll. P. Bourret, SCUBA; ORSTOM R267, R268, R472: stn 110,
SE. IlBt RBdika, 22"31~1'S.,166°36~0'E.,19 m depth, 3.vi.1976, coll. P. Laboute, SCUBA; ORSTOM
R1278: stn 184, SE. of IlBt Ua, 22"42. 1'S.,166°49.0'E., 16 m depth, 8.vi.1977, coll. G. Bargibant,
SCUBA; ORSTOM R1070: stn 249, NW. I1Bt Vua, Lagon Sud, 21°33.4'S.,166043.5'E., 35 m depth,
10.xii.1978, coll. P. Laboute, SCUBA.
Comparative material. Northwest shelf, W.A.: NTM 21021: NW. of Darnpier Archipelago,
20°14'S.,117037'E., 29 m depth, 20.x.1982, coll. J. Blake, CSIRO RV 'Soela', trawl. Great Barrier
Reef, Qld: QM GL707 (fragment NTM 21529): Noggin Passage, E. of Frankland Is, 17'13.5'S.,
146"17'E., 42 m depth, date of collection unknown, coll. QFS, dredge; QM GL2748 (fragment NTM
21553): W. of Sudbury Reef, E. of Cairns, 17°03'S.,1460071'E., 34.5 m depth, 28.i.1981, coll.
QFS, dredge; QM GL2752 (fragment NTM 21558): Stanley Reef, outer slope and lagoon, 19O16'S.,
14S006'E., 13 m depth, 18.xi.1983, coll. A. Kay, SCUBA. South-east Queensland: QM GL2747
(fragment NTM 21552): off Mooloolabah, 26"43'S.,153"08'E., 86 m depth, 12.xii.1980, coll. N. Hall,
trawl; QM G300395: Inner Gneerings, off Mooloolabah, 26"38.SS., 153°09.5'E., 14 m depth, 1.v. 1991,
coll. J . Thorogood, SCUBA; QM G300025: Outer Gneerings, off Mooloolabah, 26°39'S.,153010'E.,
25 m depth, 10.xii.1991, coll. J. N. A. Hooper and S. D. Cook, SCUBA.
Description
Colour. Orange-brown alive (Munsell 10R 6/10), beige in ethanol.
Shape. Massive, irregular, lobate, bulbous-digitate sponge, up to 400 mm high, 300 mm
maximum breadth, with small bulbous lobes forming digits (but collapsing and becoming
flattened in preserved material) covering entire mass, and whole sponge excavated by large
oscules; together the excavations and fused bulbous digits produce a honeycombed clathrous
growth form.
Surface. Paler, 'dusty', translucent, skin-like membrane covers entire surface, except
around large oscules; oscules usually distributed on ends of bulbous digits, 10-30 mm
diameter.
Fig. 27. Echinochalina (Echinochalina) intermedia (Whitelegge): A , specimen QM
G300684, choanosomal (auxiliary) quasidiactinal spicules (coring fibres); B, principal
styles (echinating fibres); C, section through peripheral skeleton.
New Caledonian Poecilosclerids
1275
Fig. 28. Echinochalina (Echinochalina) intermedia (Whitelegge): A , specimen ORSTOM
R1278 in situ (photo P . Laboute); B, specimen QM G300684 (ORSTOM R1299) in situ
(photo P. Laboute); C, SEM skeletal structure (magnified 1 0 0 ~ )D,
; SEM fibre characteristics (magnified 8 0 0 ~ ) E,
; preserved specimen QM G300684 (ORSTOM R1299).
Table 12.
Comparison in spicule dimensions between New Caledonian and Australian material of
Echinochalina (Echinochalina) intermedia (Whitelegge)
Measurements are given in micrometres
New Caledonia Great Barrier
Reef
South-east
Qld
22%.
26"s.
NW. Shelf,
W.A.
? Illawarra
Cargados
Carajos,
N.s.w.~
Indian oceanA
20%
16%
34%.
200
3
140-185
4
Latitude
17-19"s.
Coring auxiliary styles
L
W
169-(I 75.3)-189 184-(193.8)-205 184-(196.2)-208 89-(148.2)-180
1.2-(1.9)-2.5
2.0-(3.1)-5.0
1.5-(2.2)-3
1.5-(2.5)-4.0
Echinating oxeas
A
Dendy (1922).
Hallmann (1912).
1276
J. N. A. Hooper and C. L h i
Ectosome. Membraneous, arenaceous surface, without specialised spiculation, although
auxiliary spicules may poke through surface forming sparse, erect bundles, or lying
individually tangential to surface. Primary (ascending) choanosomal fibres also terminate
close to surface, sometimes protruding slightly through ectosomal membrane and forming
low conules. Collagenous spongin in peripheral skeleton heavy, granular, infiltrated with
detritus, including both sand grains and foreign spicules.
Choanosome. Skeletal architecture irregularly plumo-reticulate, with well-developed
spongin fibre skeleton, although fibres only poorly differentiated into primary and secondary
components. Primary fibres (60-100 pm diameter), more-or-less ascending to the surface,
meandering through choanosome, cored by multispicular tracts composed of auxiliary
spicules. Spicule tracts occupy only about 40-60% of fibre diameter, relatively well compacted although becoming more plumose towards periphery. Secondary fibres (30-60 pm
diameter), usually interconnecting primary tracts, uni-, pauci- or rarely aspicular. Fibre
meshes cavernous, irregular, eliptical or oval, 180-440 pm diameter, smaller in choanosomal
region than in peripheral skeleton. Choanosomal mesohyl contains abundant granular
collagenous spongin, heavy deposits of detritus, although detritus occurs only within fibre
meshes and not inside spongin fibres, and few auxiliary spicules dispersed between fibres.
Fibres sparsely echinated by thin principal styles. Choanocyte chambers difficult to observe
due to dense detritus in skeleton, oval, 30-110 pm diameter.
Megascleres (refer to Table 12 for dimensions). Choanosomal (auxiliary) spicules,
coring fibres, long, very thin, straight, with hastate points, rounded smooth bases, varying
from styles, tornostyles, or strongyles in about equal proportions; usually with blackened
axial canals.
Echinating (principal) styles, short, thin, straight, with hastate points and rounded, rarely
subtylote smooth bases.
Microscleres. Absent.
Distribution
Probably widely distributed throughout the Indo-west Pacific, although so far known only
from Great Barrier Reef, Queensland [Direction Is (Burton 1934); Frankland Is, Sudbury
Reef, Stanley Reef (present study)], southern Queensland coast [Mooloolabah (present
study)], New Caledonia (present study), N.S.W., possibly the Illawarra region (Lendenfeld
1888; Whitelegge 1902; Hallmann 1912), Tasmania (Shaw 1927), Northwest Shelf of Western
Australia [Dampier Archipelago (present study)] and Cargados Carajos, Indian Ocean
(Dendy 1922). Depth range in the New Caledonian region is 8-35 m.
Remarks
The New Caledonian population of E. intermedia, a new locality record and most easterly
record for the species, differs from other known populations in having abundant detritus
within both the choanosomal and ectosomal mesohyl, lying between fibre meshes (but not
inside fibres). In all the other specimens examined, and from the published literature,
detritus is never more than minimal in this species, and usually absent completely. New
Caledonian populations of E. intermedia also differ from most other specimens in having
much more slender echinating styles, and in this regard these populations are most similar
to those from Mooloolabah, southern Queensland (Table 12). In all other respects, all
specimens are clearly conspecific in growth form, skeletal architecture and spicule geometry,
although the geometry of auxiliary spicules may range from exclusively styles (seen in the
specimen from the Northwest Shelf, W.A.) to virtually all strongyles [in material from
Mooloolabah, southern Queensland, and also described by Dendy (1922) for material from
Carnados Carajos].
In using the name 'intermedia' for this species, we follow Dendy (1922: 71, pl. 2, fig. 8),
who, according to Whitelegge (1902), was familiar with the type specimen, which is
presently missing, and who describes and illustrates a specimen from the Indian Ocean
that agrees closely with present material. Nevertheless, the synonymy of this species is still
New Caledonian Poecilosclerids
1277
slightly confused, and the validity of the 'type' material of Thalassodendron viminalis and
Echinoclathria intermedia, and their relationships to the concept of Echinochalina intermedia
requires further clarification. For the present, Echinochalina intermedia is used in the sense
of Hallmann (1912) and Dendy (1922).
Echinochalina (Protophlitaspongia) laboutei, sp. nov.
(Figs 29-30, Table 13)
Material Examined
Holotype. QM G 300685 (ORSTOM R1301 +E084; fragment NTM 23870): stn 106, N. entrance,
Rkcif des Cinq Milles, SW. New Caledonia lagoon, 22"293'S.,l66"44.4'E., 8 m depth, 30.iv.1976,
coll. P. Laboute, SCUBA.
Description
Colour. Bright red alive (Munsell 5R 4/10), beige in ethanol.
Shape. Branching, digitate, arborescent sponge (220 mm high), stipitate, with holdfast
and short cylindrical stalk (28 mm long, 9 mm diameter), long cylindrical branches (3-6 mm
diameter), frequently bifurcate midway along their length, tapering to rounded tips.
Surface. Macroscopically even; surface slightly bulbous, with concentric swellings and
constrictions at more-or-less regular intervals along branches (2-4 mm apart), superficially
reminiscent of an Zsis gorgonian; surface microscopically porous, finely hispid, with fibre
endings from choanosome protruding slightly through surface. Oscules not visible. Texture
firm, slightly compressible, branches flexible, whip-like.
Ectosome. Auxiliary styles stand perpendicular or lie paratangential to surface, usually
forming sparse bundles, and always lying outside spongin fibres. Ectosomal skeleton
dominated by well-developed peripheral fibre skeleton, with tapering erect fibre endings
protruding about 100-250 pm from the surface of the tangential spongin fibre skeleton.
Peripheral spongin fibres lying perpendicular to surface always have a unispicular core of
larger oxeas, protruding a short distance through fibres, whereas echinating oxeas sparsely
dispersed on peripheral fibres. Mesohyl with very light collagenous spongin seen within
fibre meshes.
Fig. 29. Echinochalina (Protophlitaspongia) laboutei, sp. nov.: A, holotype QM G300685,
choanosomal principal oxeas (coring fibres); B, echinating oxeas; C, ectosomal auxiliary
styles; D, section through peripheral skeleton; E, surface fibres.
1278
J. N. A. Hooper and C. Lkvi
Choanosome. Skeletal architecture regularly reticulate, renieroid, with very tightly
condensed axial skeleton, occupying about 30% of branch diameter, and regularly reticulate
extra-axial skeleton. Organic spongin fibre skeleton emphasised over inorganic spicular
skeleton. Fibres heavy, stratified in cross-section, with differentiated primary and secondary
elements and differences between axial and extra-axial fibre skeletons. Primary fibres in
axis, condensed, 120-180 pm diameter, run longitudinally through branches and are cored
by paucispicular plumose tracts of choanosomal oxeas; these fibres are interconnected by
short, thinner, uncored secondary axial fibres, 50-90 pm diameter, together forming
compressed oval meshes in axial region, 90-180 pm diameter. Primary fibres in extra-axial
region are radial, 50-90 pm diameter, and extend from edge of axis, standing perpendicular
to it to surface, cored by uni-, bi- or paucispicular tracts of choanosomal oxeas, usually
end-on-end, rarely overlapping; primary fibres interconnected by aspicular, rarely unispicular,
transverse secondary fibres, 18-40 pm diameter; fibre meshes in extra-axial and peripheral
regions of skeleton usually elongate, generally larger in periphery (160-210 pm maximum
diameter) than in axial region (100-170 pm maximum diameter). Sparse tertiary fibre
network also present, 10-20 pm diameter, aspicular, subdividing extra-fibre skeletal meshes.
Echinating oxeas heavier in axial than peripheral region, scattered mainly (but not
exclusively) on primary fibres, standing perpendicular to fibres. Mesohyl with very light
collagenous spongin; choanocyte chambers small elongate-oval, 40-60 pm diameter.
Megascleres (refer to Table 13 for dimensions). Choanosomal principal oxeas (coring
fibres) relatively short, straight, stout, with hastate, sharply pointed, telescoped ends.
Echinating oxeas similar in morphology to principal megascleres, but substantially shorter,
marginally thinner, also with hastate points and telescoped tips.
Fig. 30. Echinochalina (Protophlitaspongia)laboutei, sp. nov.: A-B, specimen QM G300685 (ORSTOM
R1301+ E084) in situ, with commensal ophiuroid (photos P. Laboute); C, SEM fibre characteristics
(magnified 8 0 0 ~ )D,
; SEM skeletal structure (magnified 8 0 ~ ) .
'
New Caledonian Poecilosclerids
1279
Ectosomal auxiliary styles long, slender, straight or sinuous, with rounded bases and
tapering raphidiform points.
Microscleres. Absent.
Distribution
Known only from the south-west lagoon of New Caledonia, on coral rubble and coral
reef substrates, 8 m depth.
Remarks
Echinochalina laboutei, sp. nov., is closely related to three other species of Protophlitaspongia from the Indo-west Pacific region: E. bispiculata (Dendy, 1895: 246) from Port
Phillip Heads, Vic., E. oxeata (Burton, 1934: 562) from the Great Barrier Reef and Moreton
Bay, Qld, and another new species from the Noumea lagoon described below, E. bargibanti,
sp. nov. All four species are very similar in several features (more-or-less digitate growth
forms, reticulate skeletal structure with very well developed spongin fibres, possession of
principal and echinating oxeas and subtylote auxiliary styles, all very thin and poorly
silicified), but differ subtly in many other characters (size of fibres, fibre mesh diameter,
presence or absence of an axial skeleton, density of spicules in the skeleton, degree of
silicification of spicules, presence or absence of particular megascleres and microscleres,
specific spicule size and geometry). Nevertheless, these subtle differences are considered here
to justify the distinction of these taxa at the species level, and they provide a good example
of character transformations between allopatric populations of sibling species. These four
species (together with another, undescribed, deeper-water species from southern Queensland),
represent the only known members of the subgenus Protophlitaspongia, which is probably
indigenous to this region.
Echinochalina bispiculata is similar to E. laboutei in its spiculation (smaller oxeas
echinating fibres and larger oxeas of the same geometry coring fibres) and skeletal structure
(subisodictyal-renieroid skeletal structure dominated by spongin fibres), but E. bispiculata
Table 13. Comparison in spicule dimensions between the New Caledonian and Australian species of
Echinochalina (Protophlitaspongia)
Measurements in micrometres
E. laboutei,
sp, nov.
(QM G300685)
E. bargibanti,
sp. nov.
(QM (3301270)
E. bispiculata
(Dend~)
(NMV G23 19)
E. oxeata
(Burton)
(QM G300030)
105-(186.8)-266
2.0-(3.8)-5.0
94-(101.7)-163
1.5-(3.8)-7.5
Coring oxeas
L
52-(75.1)-108
1.8-(2.9)-4.0
55-(73.3)-98
0.8-(1.9)-3.0
W
115-(156.2)-194
1.O-(1.3-2.5
Ectosomal auxiliary styles
144-(216.1)-278
Absent
1.0-(2.4-3.5
142-(181.9)-228
1.5-(2.2)-3
L
W
28-(33 5)-42
2.0-(2.5)-4.0
Echinating oxeas
32-(46.3-58
31-(39.0)-45
2.0-(3.4-5.5
1.O-(2.0)-2.5
67-(76.2)-88
1.5-(2.4)-3 $ 5
L
Absent
Isochelae
14-(15.5)-21
Absent
Absent
L
W
Absent
Absent
Absent
31-(44.9)-62
0.5-(1.1)-1.5
W
L
Toxas
1280
J. N. A. Hooper and C. Ltvi
has a very different growth form (massive, lobate-digitate), a well-developed ectosomal
skeleton composed of choanosomal oxeas (same as those coring fibres) forming relatively
dense, erect, multispicular brushes on the surface, lacks auxiliary styles, has well-developed
extra-fibre collagenous spongin permeating the mesohyl, and the dimensions of coring
spicules are nearly twice as large as those in E. laboutei (Table 13). Echinochalina oxeata
has a very similar arborescent growth form as E. laboutei, but differs in its skeletal
structure, spicule geometry and spicule dimensions (Table 13). Echinochalina bargibanti has
a clumped-digitate growth form, lacks an axial skeleton, differs in spicule geometry and
spicule dimensions (Table 13). Further comparisons between the two New Caledonian species
are provided below.
Etymology
Named in honour of M. Pierre Laboute, ORSTOM, who has been studying and photographing the marine fauna of New Caledonia since 1971, for his significant contributions
to the marine biology of New Caledonia.
Echinochalina (Protophlitaspongia) bargibanti, sp. nov.
(Figs 31-32, Table 13)
Material Examined
Holotype. QM G301270 (ORSTOM R1251): stn 271, between Ilot Tere and Ilot N'Da, S. New
Caledonia, 22"48.5'S.,166"51.4'E., 30 m depth, 1.iv.l98O, coll. P. Laboute, SCUBA.
Description
Colour. Dark red alive (Munsell 5R 5/10), dark grey-brown in ethanol.
Shape. Lobate-digitate mass (65 rnm high, 57 mm wide), composed of clumps of
partially fused, small, thick digits, joined to a common base encrusting on a dead bivalve.
Digits are bulbous, cylindrical (18-35 mm long, 8-15 mm diameter), thicker at apex than
at point of attachment to base, usually single or bifurcate, sometimes trifurcate at apex,
often partially fused with adjacent digits midway along stem, together producing a vaguely
reticulate, excavated, honeycombed mass.
Surface. Digits with even, rounded tips, either unornamented or with few, large conules
(or secondary digits) near apical margins; conules 6-12 mm high, 4-9 mm diameter, approximately 3-7 mm apart. Oscules small, 1-2 mm diameter, located mainly on ends of, or
between, secondary digits (surface conules). Oscules have slight subectosomal sculpturing
(astrorhizae), converging towards each pore, but this is not extensively developed. Texture
rubbery, compressible, difficult to tear.
Ectosome. Sparse, tangential or paratangential tracts of auxiliary subtylostyles dispersed
over ectosomal region. Where thickest auxiliary subtylostyles may form sparse plumose
brushes, but usually these spicules are scattered. Terminal, ascending primary fibres protrude
through ectosomal spicule skeleton for up to 150 pm, containing a core of principal oxeas,
the ends of which barely protrude through ends of fibres. Relatively sparse clumps of lightly
pigmented collagen usually associated with ectosomal spicule skeleton.
Choanosome. Skeletal architecture regularly reticulate throughout, without any differentiation of axial or extra-axial/peripheral regions. Spongin fibres dominate skeleton, and
spicules both greatly reduced in abundance and poorly silicified (all with blackened axial
canals). Spongin fibres divided into primary ascending and secondary transverse (connecting)
components, with a very thin tertiary network also apparent. Primary fibres, 40-60 pm
diameter, meandering throughout skeleton but more-or-less ascending to surface, terminating
in small digitate microconules, cored by uni- or paucispicular tracts of choanosomal principal
oxeas, occupying only a small proportion of fibre diameter. Secondary connecting fibres
usually transverse, 15-25 pm diameter, aspicular or unispicular, and tertiary fibres, 8-20 pm
diameter, invariably aspicular and interconnecting both secondary and primary fibre system.
Fibre meshes cavernous, square or rounded, 80-180 pm diameter, containing very little
New Caledonian Poecilosclerids
1281
visible collagen, although distinct clumps of granular collagen plus principal oxeas and
isochelae scattered within choanosome, particularly around major fibre nodes. Echinating
acanthostyles sparse, usually associated with small fibre nodes protruding from a primary
or secondary fibre. Choanocyte chambers small, elongate-oval, 35-60 pm diameter.
Megascleres (refer to Table 13 for dimensions). Choanosomal principal oxeas (coring
fibres) relatively short, slender, with hastate, sharply pointed ends, occasionally telescoped
points.
Ectosomal auxiliary subtylostyles long, slender, invariably straight, with enlarged subtylote
bases, rounded or slightly pointed, and tapering to sharply pointed or slightly rounded tips.
Echinating oxeas rare, similar in morphology to principal oxeas, but marginally shorter
and of similar thickness.
Microscleres. Palmate isochelae moderately large, common, usually found aggregated
in clumps of collagen in choanosome.
Toxas are absent.
Fig. 31. Echinoclathria (Protophlitaspongia) bargibanti, sp. nov.: A, holotype QM G301270,
ectosomal auxiliary subtylostyles; B, choanosomal principal oxeas (coring fibres); C, echinating
oxeas; D, palmate isochelae; E, section through peripheral skeleton.
1282
J. N. A. Hooper and C. L6vi
Distribution
Known only from the south-west lagoon of New Caledonia, on coral rubble substrate,
30 m depth.
Remarks
If a phylogenetic study of Protophlitaspongia eventually shows E. laboutei (New
Caledonia) and E. bispiculata (south-east Australia) to be sister-species, then it is quite
probable that E. bargibanti (New Caledonia) is also a sister-species of E. oxeata (north-east
Australia). Both pairs of species have comparable similarities in their skeletal structure and
fibre characteristics, although they differ subtly in many other ways.
Echinochalina bargibanti differs from the other three described sister-species in having
isochelae microscleres (whereas E. laboutei has toxas, and the others have no microscleres
Fig. 32. Echinoclathria (Protophlitaspongia) bargibanti, sp. nov.: A, holotype
QM G301270 (ORSTOM R1251), SEM fibre characteristics (magnified 4 0 0 ~ ) ;
B, SEM skeletal structure (magnified 100 x ) ; C, three specimens in situ (holotype
in right foreground) (photo P. Laboute); D, holotype; E, fibre structure, showing
coring spicules and sparse aggregates of collagen plus spicules located near fibre
nodes (indicated by arrows) (scale =200 pm).
New Caledonian Poecilosclerids
1283
at all). This species also differs from E. laboutei in growth form: lacking a condensed axial
skeleton, having instead undifferentiated axial and extra-axial regions; having a more
cavernous skeleton, with rounded or square fibre meshes, whereas E. laboutei has elongate
meshes; echinating oxeas are very scarce in the present species and the geometry of auxiliary
spicules differs between the two species. Spicule measurements of all these species are
compared in Table 13.
Etymology
Named for M. Georges Bargibant, ORSTOM, in recognition of his substantial contributions in documenting the marine biology of the New Caledonian region.
Family RASPAILIIDAE Hentschel
Definition
Poecilosclerida usually with very hispid surface, specialised ectosomal skeleton of brushes
of small thin styles or oxeas, surrounding individual long thick styles or oxeas; choanosomal
skeleton varies from compressed axial skeleton, to plumo-reticulate or exclusively reticulate
structures; spongin fibres enclose choanosome styles, oxeas or both; spined styles (acanthostyles) or highly modified styles echinate fibres; microscleres usually absent, although single
raphides or bundles (trichodragmata) occur in some genera (from Hooper 1991b).
Remarks
The present work brings the total number of raspailiid species in the New Caledonian
region to seven, three apparently endemic and four also recorded from other Indo-west
Pacific localities. These species are: Raspailia wilkinsoni Hooper (Great Barrier Reef),
Ceratopsion clavata Thiele (Moreton Bay, Qld, and Japan), C. palmata Hooper (tropical
western and northern Australia), Aulospongus clathrioides LCvi (endemic), A. gardineri
(Dendy) (Amirante, W. Indian Ocean), Plocamione pachysclera ( L h i & LCvi) (endemic,
deep water), and Lithoplocamia dolichosclera LCvi & LCvi (endemic). Three of these species
(R. wilkinsoni, C. clavata and C. palmata) are new records for the region. The taxonomic
system used here follows Hooper (1991b).
Genus Raspailia Nardo
Raspailia Nardo, 1833: 522 [type species Raspailia typica Nardo, 1833: 5221 [fullsynonymy and discussion
given by Hooper (1991b: 1195)l.
Diagnosis
Raspailiidae with axial compression and differentiated axial and extra-axial skeletons,
or sometimes reduced to reticulate structures (subgenus Clathriodendron); axial skeleton
typically condensed, reticulate, with fibres and spicules forming distinct tracts; extra-axial
skeleton plumo-reticulate, perpendicular to axis, cored by large styles or oxeas in tracts, or
plumose, with single extra-axial spicules; specialised ectosomal skeleton present, with bundles
of erect, small styles surrounding bases of protruding extra-axial megascleres; choanosomal
spongin fibres echinated by club-shaped acanthostyles (subgenus Raspailia), acanthose
rhabdostyles (subgenus Raspaxilla), modified cylindrical styles with greatly enlarged bases
(subgenus Hymeraphiopsis), or secondarily lost (subgenus Syringella); microscleres absent
(from Hooper 1991b).
Remarks
This genus has been recently revised, mainly on the basis of Australian material, and five
subgenera were recognised (as defined above). It is possible, however, that several more
subdivisions within the genus may be appropriate eventually, although more extensive
studies of other faunas are first required. Prior to the present work, no species of Raspailia
had been recorded from the New Caledonian region, although Plocamione pachysclera was
initially referred to the genus by LCvi and LCvi (1983b).
1284
J. N. A. Hooper and C. LCvi
Raspailia (Raspailia) wilkinsoni Hooper
(Figs 33-34, Table 14)
Raspailia wilkinsoni Hooper, 1991b: 1220-22, figs 17-1 8.
Material Examined
Holotype. NTM 22734: Back Reef, Davies Reef, off Townsville, Great Barrier Reef, Qld,
18"5O'S.,147"39'E.,15 m depth, ll.viii.1989, coll. C. R. Wilkinson (AIMS ref. R22).
New Caledonian material. QM G300744 (ORSTOM R766): stn 206, Banc de la Torche, Ile des
Pins, 22°51.1'S.,167040.0'E., 36 m depth, 8.iii.1978, coll. P. Laboute, SCUBA; QM G300747
(ORSTOM R322): stn 196, barrier reef M'Bkre, false passe de Uitoe, 22°18~1'S.,166011.1'E.,45 m
depth, 9.xi.1977, coll. J.-L. Menou, SCUBA; ORSTOM unregistered: RCcif exterior de Passe Boulari,
60 m depth, l.vi.1977. coll. P. Laboute, SCUBA.
Comparativematerial. ORSTOM TI: precise locality unknown, Tanna, S. Vanuatu (New Hebrides).
Description
Colour. Dark orange-brown alive (Munsell 5YR 5/8), dark brown in ethanol.
Shape. Arborescent, digitate, bushy, 80-200 mm high, 60-110 mm maximum breadth
of branches, with short or long cylindrical stalk and basal holdfast, 12-85 mm long, 4-8 mm
diameter, and thick, bifurcate, cylindrical branches, 25-95 mm long, 5-1 1 mm diameter,
tapering to sharp points at apex.
Surface. Shaggy, prominently microconulose surface on branches, more compact on
stalk. Surface conules up to 3 mm long, narrow, close-set. Oscules not observed; inhalent
pores small, up to 0 . 5 mm diameter, scattered between surface conules. Texture firm,
compressible, with easily flexible branches and less flexible stalk.
Ectosome. Membraneous, without specialised raspailiid skeleton, although smaller,
thinner, wispy auxiliary spicules scattered in peripheral skeleton mostly tangential to surface.
Subectosomal region dominated by micro-digitate projections, 1 5-3 mm long, predominantly
cored by ascending, plumose, multispicular tracts of long, thick, subectosomal auxiliary
spicules, protruding through surface for about 100 pm, usually in bundles. Ascending
subectosomal spicule tracts interconnected by transverse spicule tracts composed of
choanosomal principal oxeas. Whole peripheral skeleton cavernous, with widely spaced
extra-axial spicule tracts producing more-or-less rectangular meshes, up to 500 pm diameter.
Peripheral spicule tracts multi- or paucispicular, heavily echinated by acanthostyles, more
so than in axial region. Collagenous spongin abundant in mesohyl of peripheral skeleton.
Choanosome. Skeleton reticulate, with well differentiated compressed axial and reticulate
extra-axial regions; axial skeleton tightly compressed, composed of close-set, reticulate, heavy
spongin fibres, 100-150 pm diameter, with stratified spongin fibres producing tight meshes,
up to 150 pm diameter. Fibres cored by choanosomal oxeas in pauci- or multispicular tracts,
and lightly echinated by acanthostyles. Mesohyl in axial region only lightly invested with
collagenous spongin.
Megascleres (refer to Table 14 for dimensions). Choanosomal principal oxeas long, thin
or thick, usually symmetrically curved, tapering to sharp points.
Subectosomal auxiliary spicules predominantly oxeas, less commonly styles (cf. holotype),
long, thick, straight or slightly curved, often anisoxeote, with rounded bases (for styles)
and tapering to sharp points.
Ectosomal auxiliary spicules short, thin, whispy, varying from symmetrically curved oxeas
to sinuous anisoxeas, tapering to sharp points.
Echinating acanthostyles short, thixf, straight, with subtylote bases and tapering rounded
blunt tips; spines long, slender, conical, sharply pointed, concentrated mainly on bases
and points.
Microscleres. Absent.
New Caledonian Poecilosclerids
1285
Distribution
Central region of the Great Barrier Reef, south-west lagoon of New Caledonia and
Tanna, southern Vanuatu, depth range 15-60 m, on live and dead coral substrates.
Remarks
The material described above from New Caledonia and Vanuatu is only the second record
of this species, which was previously known only from the holotype. There are a number
of differences between these three populations, but these are considered here to be relatively
minor and in our opinion cannot justify their differentiation at the species level.
Both the holotype and the Vanuatu specimen lack any sort of ectosomal spiculation,
whereas New Caledonian specimens have sparse, wispy tracts of auxiliary oxeas (or
anisoxeas) mostly lying tangential to the surface. These ectosomal auxiliary spicules do not
form a specialised ectosomal skeleton, synapomorphic for the Raspailiidae, and hence in this
regard the original concept of the species is retained. Similarly, the larger subectosomal
auxiliary spicules in both the holotype and the Vanuatu specimen are exclusively styles,
Fig. 33. Raspailia wilkinsoni Hooper: A, specimen QM G300744, choanosomal principal oxeas;
B, subectosomal auxiliary oxeas and styles; C, ectosomal auxiliary oxeas/ anisoxeas; D, echinating
acanthostyles; E, section through peripheral skeleton.
J. N. A. Hooper and C. L6vi
1286
Fig. 34. Raspailia wikinsoni Hooper: A , specimen QM G300744 (ORSTOM
R766), SEM skeletal structure (magnified 200x); B, preserved specimen; C,
unregistered ORSTOM specimen in situ (photo P . Laboute); D, specimen QM
G300744 (ORSTOM R766), SEM fibre characteristics (mangified 600 x ); E,
preserved specimen QM G300747 (ORSTOM R322).
Table 14. Comparison in spicule dimensions between New Caledonian, Vanuatuan and
Great Barrier Reef populations of Raspailia (Raspailia) wilkinsoni
-
Measurements are given in micrometres; comparative data from Hooper (1991b)
- New Caledonia
Vanuatu
Great Barrier Reef
-
-
-
-
Choanosomal oxeas
L
W
188-(238.1)-285
4-(6.2)-10
650-800
6-8
151-(214.0)-346
4-(6.6)-10
Subectosomal auxiliary oxeas and styles
L
W
255-(382.9-524
8-(9.3-12
L
W
Ectosomal auxiliary oxeas and anisoxeas
106-(159.6)-208
Absent
I-(]. 3-2.5
250-350
10
Echinating acanthostyles
267-(353 ~3)-542
3-(5.6)-10
Absent
New Caledonian Poecilosclerids
1287
whereas in New Caledonian material these spicules are mostly oxeas, less commonly styles,
occasionally anisoxeas. Nevertheless, the dimensions of these spicules between Australian
and New Caledonian material are closely comparable (Table 14). Similarly, transverse
sections of the subectosomal spicule and fibre skeletons in New Caledonian specimens are
thicker, and contain a larger core of multispicular tracts, whereas those in the holotype are
much less robust.
Genus Ceratopsion Strand
Ceratopsion Strand, 1924: 33 [type species Ceratopsis expansa Thiele, 1898: 571 [full synonymy and
discussion given by Hooper (1991b: 1327)l.
Diagnosis
Raspailiidae with well-differentiated axial and extra-axial skeletons, and condensed
reticulate axial skeleton cored by styles, anisoxeas or oxeas, often sinuous; extra-axial
skeleton radially arranged long styles, anisoxeas or strongyles, sinuous or straight, projecting
through surface, surrounded by specialised raspailiid ectosomal skeleton of oxeas or styles;
echinating megascleres absent; microscleres absent (from Hooper 1991b).
Remarks
Eleven species of Ceratopsion have been described (Hooper 1991b), extending in distribution from Japan, New Zealand, southern Indonesia, northern and eastern Australia, South
Africa and the Mediterranean. The present records of C. clavata and C. palmata from
New Caledonia are the easternmost records of the genus in the Pacific.
Ceratopsion clavata Thiele
(Figs 35-37, Table 15)
Ceratopsis clavata Thiele, 1898: 57, pl. 5, fig. 23, pl. 8, fig. 42a-c.
Ceratopsion clavata. -Hooper, 1991b: 128, table 18.
Material Examined
Holotype. Unknown, possibly ZMB: Enoshima region, Sagami Bay, Japan, 139°30'N.,35010'E.,
130 m depth.
New Caledonian material. QM G300743 (ORSTOM R685): stn 202, NW. Rtcif Ana, 21°22. l'S.,
166"01.11E., 35 m depth, 8.ii.1978, coll. G. Bargibant, SCUBA; QM G300745 (ORSTOM R755)
(4 specimens): stn 209, NNE. I16t Ugo, Ile des Pins, 22°28.1'S.,166055.2'E., 25-35 m depth, l0.iii.
1978, coll. P. Laboute, SCUBA; QM G300018 (ORSTOM R1260): stn 282, 'travers de la cascade',
Tao, 20°32~1'S.,166"49~0'E.,37 m depth, 26.v.1980, coll. P. Laboute, SCUBA; QM G300720
(ORSTOM R328): stn 114, NE. Ili3t Tareti, Banc Gail, 22°221'S.,166039.0'E., 35 m depth, 29.vi.
1976, coll. P. Laboute, SCUBA.
Comparative material. QM G301289 (2 specimens): Hanlon Light, Moreton Bay, Qld, 27"28+5'S.,
153"20.9'E., 10 m depth, 3.xi.1992, coll. J. N. A. Hooper and S. D. Cook, SCUBA.
Description
Colour. Yellow (Munsell 2.5Y/8-10) or pale orange alive (5YR 7-10), pale beige to
light brown in ethanol.
Shape. Arborescent digitate, variable morphology ranging from bushy to elongate
digitate, whip-like, 60-360 mm maximum height, 25-115 mm maximum breadth (width of
branching), with short, thick, cylindrical basal stalk, 5-28 mm long, 2-11 mm diameter,
enlarged basal holdfast, and long, bifurcate, bulbous-cylindrical branches, 25-125 mm long,
5-12 mm diameter (inclusive of surface conules), tapering towards ends.
Surface. Prominently conulose surface, although varying between specimens (poorly
developed in G300720); conules close-set, irregular, sharply pointed, shaggy, up to 5 mm
long, 3 mm basal diameter.
1288
J. N. A. Hooper and C . L h i
Ectosome. Predominantly membraneous ectosomal skeleton, with subectosomal auxiliary
spicules protruding for up to 200 pm from surface, and with sparse tangential tracts of
ectosomal auxiliary oxeas dispersed around the vicinity of larger subectosomal spicules,
although not producing a specialised raspailiid skeleton. Heavy collagen in peripheral region,
with large areas completely clear of any spicules.
Choanosome. Skeleton divided into three distinct components: compressed axial core,
consisting of short, tightly meshed, heavy spongin fibres, 60-100 pm diameter, producing
small oval meshes, 30-70 pm diameter, sparsely to moderately cored by sinuous choanosomal
principal spicules; radial extra-axial skeleton, consisting of subectosomal auxiliary spicules,
occurring individually or in bundles, embedded in and standing perpendicular to axial fibres,
Fig. 35. Ceratopsion clavata Thiele: A , specimen QM G300743, choanosomal principal
strongyles/ strongyloxeas/ anisoxeas; B, subectosomal auxiliary oxeas/ anisoxeas/ styles;
C , ectosomal auxiliary oxeas; D, section through peripheral skeleton.
New Caledonian Poecilosclerids
1289
in parallel rows, and extending outwards for only one spicule length; and plumose extra-axial
skeleton, consisting of large (up to 5 mrn) plumose (or less obviously plumo-reticulate),
multispicular tracts of subectosomal auxiliary spicules, and including intermingled sinuous
coring spicules from choanosome; latter part of skeleton most prominent, corresponding
to large surface conules. Mesohyl in axial region only lightly invested with collagenous
spongin, whereas closer to periphery there are heavier deposits of more granular collagen.
Choanocyte chambers oval, up to 120 pm diameter.
Megascleres (refer to Table 15). Choanosomal principal megascleres, predominantly in
axial skeleton, usually sinuous, sometimes merely curved, rarely straight, ranging from
strongyles with evenly rounded ends, strongyloxeas with telescoped, mucronate ends, to
anisoxeas with a combination of rounded and telescoped ends. Queensland specimens
predominantly with sinuous principal spicules; New Caledonian specimens predominantly
with curved principal spicules.
Subectosomal auxiliary megascleres, forming extra-axial skeletal tracts, range in size and
geometry from small wispy oxeas, anisoxeas or styles, to large thick styles and anisoxeas;
most spicules have telescoped ends, but larger ones may have evenly rounded ends; spicules
straight, symmetrically or asymmetrically curved or less often sinuous. Queensland specimens
predominantly with thinner, sharply pointed telescoped oxeas; New Caledonian specimens
predominantly with thicker, rounded telescoped oxeas.
Ectosomal auxiliary spicules thin whispy oxeas, slightly curved, usually asymmetrical
curvature, with tapering fusiform points.
Echinating megascleres absent.
Microscleres. Absent.
Fig. 36. Ceratopsion clavata Thiele: A-E, New Caledonian material; A , specimen QM G300720
(ORSTOM R328) in situ (photo P. Laboute); B, 3 specimens QM G300745 (ORSTOM R755); C ,
specimen QM G300743 (ORSTOM R685); D, specimen QM G300018 (ORSTOM R1260), SEM axial
and extra-axial skeletons (magnified 100x); E, SEM extra-axial skeleton (magnified 3 0 0 ~ ) .
1290
J. N. A. Hooper and C. LBvi
Distribution
Sagami Bay, Japan; Moreton Bay, Queensland; and south-west lagoon of New Caledonia.
Remarks
This is the first record of the species since it was initially described from Japan by Thiele
(1898). The species shows remarkable variability in the extent to which surface conules are
developed (Figs 36A-C, 37A), in spicule size (Table 15), and in the terminations on sub-
Fig. 37. Ceratopsion clavata Thiele: A-C, Moreton Bay material; A , specimen
QM G301289 in situ (photo J . N . A. Hooper); B, SEM peripheral skeleton (magnified
200 x ) ; C , SEM transverse section through branch (magnified 80 X ) (scale = 500 km).
New Caledonian Poecilosclerids
1291
Table 15. Comparison in spicule dimensions between specimens of Ceratopsion clavata Thiele, 1898
The great range of spicule dimensions within this species is illustrated; measurements are given in
micrometres; comparative data from Hooper (1991b)
Japan
~ o l o t y ~ eQM
~ G300720
New Caledonia
QM G300743
QM G300745
QM G300018
Moreton Bay
QM G301289
Choanosomal strongyles/anisoxeas
L
W
690-1500
10-30
294-(405 '3)-680 252-(439.1)-790 251-(447.8)-868 284-(577.8)-912
2.0-(2.8)-4.0
1.5-(2.9)-4.0
2.0-(3.6-4.5
4-(5.6)-9
L
W
550-1500
10-50
234-(310.8)-418 258-(381.2)-590 172-(317.4)-453 232-(319.8)-422
3-(4.7)-6
4-(6.3)-8
6-(7.8)-10
7-(10.3-13
L
W
60-1 10
3
192-(251.4)-302 126-(193.3)-263 165-(189.6)-232 163-(213.0)-246
1.0-(1.7)-3.0
0.8-(1.4)-2.0
0.8-(1.3)-2.0
1-(2.1)-4
422-(573 '3)-805
2.0-(3.9)-6.0
Subectosomal auxiliary styles/anisoxeas
216-(258.0)-288
4.0-(5.4)-8.0
Ectosomal auxiliary oxeas
A
142-(202.4)-268
0.8-(2.1)-3.0
From Thiele (1898: 57).
ectosomal auxiliary spicules (oxeas with rounded ends, telescoped ends, sharply pointed,
etc.; Fig. 35A-C). In growth form, one specimen (QM G300720) has greatly elongated and
spindly branching, with nearly vestigial surface sculpturing, whereas others (e.g. QM G300743
from New Caledonia and QM G301289 from Moreton Bay) are very bushy, and also have
prominent close-set conules, as in the holotype. In spiculation, one specimen from New
Caledonia in particular (QM G30018) has generally larger and thicker spicules than any of
the other New Caledonian specimens, although these appear to vary greatly between all
known specimens, and spicule thickness seems to be highly variable in the species. Moreover,
all specimens described above from New Caledonia and southern Queensland have substantially smaller spicule dimensions than those published for the Japanese holotype (Thiele
1898). Nevertheless, resemblance is very close in all other features between the present
material and Thiele's (1898) original description, and this material is considered here to
be conspecific.
Ceratopsion clavata differs from most other species of the genus in lacking a clearly
defined, specialised raspailiid ectosomal skeleton: specialised ectosomal auxiliary spicules are
present in this species, but these are obviously vestigial and do not form plumose surface
structures. By comparison, in all other species examined of Ceratopsion, this feature is
consistently present (cf. Hooper 1991b: 1327-39). Ceratopsion clavata has sinuous axial
strongyles (some modified to anisoxeas or strongyloxeas, cf. the holotype description)
and extra-axial styles (many modified to anisoxeas or oxeas, usually with telescoped
ends). In this regard, the species shows some similarities to species from South Africa
[C. microxephora (Kirkpatrick, 1903)l and the Mediterranean (C. minor Pulitzer-Finali,
1983). In growth form it resembles C. dichotoma (Whitelegge, 1907) most closely, although
surface sculpturing is usually more prominent in the present species. The species is contrasted
further with other members of the genus in Hooper (1991b).
Ceratopsion palmata Hooper
(Figs 38-39, Table 16)
Ceratopsion palmata Hooper, 1991b: 1337-9, figs 78-9, table 18.
Material Examined
Holotype. N T M 23051: N, of Amphinome Shoals, Northwest Shelf, W.A., 19"19.7-23.3'S.,
119"08.8-12~2'E.,50 m depth, 19.vii.1987, coll. J. N . A. Hooper, USSR RV 'Akademik Oparin',
beam trawl.
1292
J. N. A. Hooper and C. L6vi
New Caledonian material. NTM 23899 (fragment QM G300698): exterior of Grand Rkif
Mbere, off Noumea, 22°20'S.,166013'E., 28 m depth, 27.ix.1990, coll. J. N. A. Hooper, SCUBA,
stn JH-90-021.
Comparative material. Refer to Hooper (1991b: 1337).
Description
Colour. Yellow-beige alive (Munsell 7.5YR 7-10), beige in ethanol.
Shape. Small fan, 240 mm high, 150 mm wide, 8 mm maximum thickness, with folded
margins and short stalk, 12 mm long, 5 mm diameter.
Fig. 38. Ceratopsionpalmata Hooper: A, specimen QM G300698, choanosoma1 principal oxeas; B, subectosomal auxiliary styles/ oxeas; C,ectosomal
auxiliary oxeas; D, section through peripheral skeleton.
New Caledonian Poecilosclerids
1293
Surface. Optically even, microscopically villose and hispid,surface; oscules not seen;
texture firm, rubbery.
Ectosome. Darkly pigmented collagenous surface, with long extra-axial styles protruding,
more-or-less surrounded by sparse bundles of thin ectosomal oxeas but not necessarily
confined to the region at the point of insertion of the extra-axial spicules.
Fig. 39. Ceratopsion palmata Hooper: A , specimen NTM 23899 in situ; B, specimen
QM G300698 in situ (photo J. Hooper); C, SEM skeletal structure (magnified 8 0 ~ ) .
Table 16. Comparison in spicule dimensions between specimens of Ceratopsion palmata
Hooper, 1991
Measurements, given in micrometres, are presented as range (mean) of lengths and widths;
comparative data from Hooper (1991b)
Holotype
Australia
(n = 5)
New Caledonia
(n=1)
L
W
229-(348.4)-391
4-(12.9)-18
Choanosomal oxeas
231-612
3-20
269-(293.0)-3 12
6-(9.9)-13
L
W
521-(722.3-808
5-(10.3)-18
Subectosomal styles
583-2232
3-49
549-(645.4)-751
5-(6.6)-8
L
W
202-(340.1)-442
1-5-(3.5)-6
Ectosomal oxeas
191-741
0.5-15
194-(247.9)-357
2-(3.3)-4
1294
J. N. A. Hooper and C. Ltvi
Choanosome. Axial and extra-axial skeletons clearly differentiated and extra-axial
skeleton composed of two components. Axis compressed, consisting of bundles of oxeas
mostly running longitudinally through the sponge, bound together by primary spongin fibres,
60-1 10 pm diameter, in vaguely cylindrical bundles, interconnected with thinner secondary
fibres. Extra-axial skeleton perpendicular to axis, running to surface, composed of plumose,
ascending multispicular tracts interconnected by fewer transverse, uni- or paucispicular tracts,
both sorts of fibres cored by choanosomal oxeas, producing a plumo-reticulate extra-axial
skeleton. Embedded in the ends of extra-axial plumose tracts are individual subectosomal
megascleres, standing perpendicular to surface. Mesohyl contains abundant, darkly pigmented, granular collagen. Choanocyte chambers small, oval, up to 70 pm diameter.
Megascleres (refer to Table 16 for dimensions). Choanosomal principal oxeas, in both
axial and plumo-reticulate extra-axial skeletons, usually robust, slightly curved, rarely
straight, with sharply pointed or faintly telescoped ends.
Subectosomal auxiliary styles, in peripheral extra-axial skeleton, long, slender, slightly
curved near base, tapering to fusiform points, with evenly rounded bases or occasionally
modified to oxeas.
Ectosomal auxiliary spicules are symmetrical oxeas, relatively long, slender, slightly curved
or straight.
Echinating megascleres absent.
Microscleres. Absent.
Distribution
Tropical Western Australia, northern Australia and New Caledonia; coral reef and interreef regions, 5.5-76 m depth.
Remarks
The present specimen is the most easterly record for the species, and there are some
slight differences between it and the Australian population. The New Caledonian specimen
is yellower in coloration; it is a flat fan with folded margins rather than with digitate
projections on the margins; the ectosomal skeleton is more poorly developed, with only few
ectosomal bundles seen; extra-axial styles are sometimes modified to oxeas; and spicule sizes
differ slightly from other material (Table 16). However, these differences are minor and the
species appears to be relatively homogeneous across its broad distribution. Relationships
between C. palmata and other species are discussed in detail by Hooper (1991b).
Genus Aulospongus Norman
Aulospongus Norman, 1878: 267 [type species Haliphysema tubulatus Bowerbank, 1873: 291 [full
synonymy and discussion given by Hooper (1991b: 1307)l.
Diagnosis
Raspailiidae with specialised growth form of fused plumose skeletal columns, without or
with greatly reduced reticulate skeletal structure; axial and extra-axial skeletons poorly
differentiated, without axial compression; choanosomal skeletal tracts plumose or bifurcate
microcionid-like; extra-axial skeletal tracts radial, plumose, diverging at periphery;
choanosomal megascleres rhabdostyles; extra-axial skeletal columns echinated by acanthorhabdostyles or acanthostyles; specialised raspailiid skeleton present or absent; microscleres
may include raphides (from Hooper 1991b).
Remarks
Two species of Aulospongus have been previously recorded from the New Caledonian
region, one deeper-water species [A. gardineri (Dendy) from 300-430 m depth] and one
shallow-water species (A. clathrioides LCvi). The former species is also known from
Amirante, whereas the latter is endemic and relatively abundant in the lagoon.
1295
New Caledonian Poecilosclerids
Aulospongus clathrioides LCvi
(Figs 40-41)
Aulospongus clathrioides LBvi, 1967: 21, text-fig. 5, pl. 2, fig. c. -Hooper, 1991b: 1307, fig. 67a-d.
Material Examined
Holotype. MNHN DCL 823: Canala, Melasceu, New Caledonia, 04.i.1962, coll. Mission SingerPolignac.
Other New Caledonian material. QM G300721 (ORSTOM R714): stn 198, pinnacle S. of Canyon
Central, Chenal des Cinq Milles, 22"30.4'S.,166"45.1'E., 35 m depth, 153.1978, coll. G. Bargibant,
SCUBA; QM G300742 (ORSTOM 'cfR714'): stn 106, N. entrance, RBcif des Cinq Milles, SW. New
Caledonia lagoon, 22°29.3'S.,166044.4'E., 8 m depth, 16.ii.1978, coll. P. Laboute, SCUBA; QM
G300746 (ORSTOM R515): same locality, 30.iv.1976, 8 m depth; ORSTOM R854: stn 198, pinnacle
S. of Canyon Central, Chenal des Cinq Milles, 22°30.4'S.,166045.1'E., 40 m depth, ll.iv.1978, coll.
P. Laboute, SCUBA; ORSTOM R696: Plateau des Cinq Milles, 22°29.3'S.,166044.4'E., 35 m depth,
16.ii.1978, coll. P. Laboute, SCUBA.
Fig. 40. Aulospongus clathrioides L h i : A , specimen QM G300746, choanosomal principal
styles/ anisoxeas; B, echinating acanthostyles; C , section through peripheral skeleton.
1296
J. N. A. Hooper and C. LCvi
Description
Colour. Yellow orange alive (Munsell 7.5YR 7/10), khaki brown in ethanol.
Shape. Arborescent, bushy, 55-180 mm high, 40-110 mm maximum branching width,
with short woody stalk, 22-40 mm long, 2-8 mm diameter, and enlarged basal attachment;
branches prolific, usually repeatedly bifurcate and anastomosing, producing a reticulate
bushy mass. Branches thinly cylindrical or slightly flattened, up to 6 mm diameter, tapering
to rounded bifurcated tips.
Surface. Thin translucent skin-like ectosomal membrane stretched between surface
conules, intact in only few places in preserved material. Branches with uneven, shaggy, very
prominent microconulose surface, and macroscopically hispid; basal stalk woody, more
darkly pigmented; branch tips even, hispid only. Oscules occasionally seen on lateral sides
of branches, between surface conules, up to 2 mm diameter. Texture of branches soft,
flexible, spiky; stalk firm, flexible.
Ectosome. Without specialised spiculation, with plumose brushes of choanosomal
principal styles protruding up to 150 pm from peripheral fibres. Peripheral skeleton
dominated by swollen spongin fibres, with fibre nodes protruding through surface and
forming surface conules; fibre nodes 350-600 pm long, up to 250 pm maximum diameter.
Mesohyl in peripheral skeleton with heavy collagenous spongin.
Choanosome. Skeleton predominantly plumose, although a slightly compressed axial
skeleton also present; both axial and extra-axial skeletons dominated by well-developed
spongin fibre system, with spicule skeleton proportionally reduced. Axial fibres small,
close-set, with vestigial core of choanosomal principal styles; axial fibre meshes 60-90 pm
diameter, with only moderate quantities of collagen. Extra-axial fibres occupy most of the
branch diameter, with primary ascending spongin fibres, 60-200 pm diameter, cored by
multispicular plumose columns of choanosomal principal styles (although spicule columns
Fig. 41. Aulospongus clathrioides Lhi:
A , specimen QM G300721 (ORSTOM
R714) in situ (photo G . Bargibant); B,
specimen (ORSTOM R854) in situ (photo
P . Laboute); C, specimen QM G300742
(ORSTOM 'cfR714'), SEM of peripheral
skeleton (magnified 8 0 ~ ) D,
; SEM of
fibre characteristics (magnified 1 0 0 0 ~ ) .
New Caledonian Poecilosclerids
1297
occupy less than 50% of fibre diameter, and plumose brushes composing columns are
relatively sparse); secondary fibres short, thin, 100-150 pm long, 30-60 pm diameter, cored
by 0-2 choanosomal styles, and anastomoses between primary and secondary fibres produce
elongate cavernous meshes, 200-550 pm diameter. Echinating acanthostyles moderately
sparse, dispersed relatively evenly over primary and secondary fibres, but probably more
abundant in extra-axial rather than axial skeleton. Collagenous spongin heavy in extra-axial
region; choanocyte chambers small, oval, 15-40 pm diameter.
Megascleres. Choanosomal principal styles of a single category only, although length
and thickness variable; usually long, slender, slightly curved, occasionally sinuous, with
rounded smooth bases, sometimes anisoxeote, tapering fusiform points. Length 145-(249.3)454 pm, width 3-(5.1)-7 pm.
Subectosomal and ectosomal megascleres absent.
Echinating acanthostyles long, slender, slightly curved towards basal end, with rounded
or slightly swollen, mostly smooth bases, pointed or slightly swollen tips, and evenly covered
with small granular spines (except for basal region). Length 58-(70.5)-82 pm, width 1.5(2.4)-4.0 pm.
Microscleres. Absent.
Distribution
Known only from the New Caledonia lagoon, inter-reef region, coral and sand substrate,
8-40 m depth.
Remarks
Aulospongus clathrioides is slightly unusual to the genus in retaining the rudiments of
an axial skeleton, although this structure is greatly reduced in relation to the extra-axial
region. By comparison, most species of Aulospongus lack axial compression or any marked
differentiation between axial and extra-axial regions. Similarly, although typical members
of the genus have a peculiar growth form consisting of fused bundles of plumose fibres,
together forming rounded, massive and clathrous structures, A. clathrioides is digitate
(Fig. 41A), and as such, more similar to many species of Raspailia. However, the prominent
surface fibre bundles in this species, consisting of large, swollen spongin fibres protruding
through the peripheral skeleton, are distinctly homologous to the typical Aulospongus
condition, and in this regard there are also clear affinities with other members of the genus.
Further discussion of A. clathrioides, and its relationships to other Aulospongus, has been
presented by Hooper (1991b: 1307-8).
Discussion
From our relatively rudimentary knowledge of the sponge fauna of the New Caledonian
region, based exclusively on the recent works of LCvi (1967 et seq.), Ltvi and LCvi (1978
et seq.) and Desqueyroux-Faundez (1984, 1987), it has been suggested that levels of
endemism for the whole sponge fauna in this region are relatively high (about 60% for all
Demospongiae). However, most of these 'apparent endemics' were restricted to the deeperwater shelf and slope fauna, with ancestral links to the Cretaceous fauna of northern New
Zealand (LCvi and LCvi 1978): of the 76 species described from deeper-waters, 55 have not
been recorded from any other province (72% endemism). In contrast, it was previously
thought that the shallower-water fauna contained far fewer indigenous species, having a
'widespread Indo-Pacific' fauna (Ltvi 1979), in common with many other groups of marine
invertebrates (e.g. Crosnier and Bouchet 1991). Of the 68 shallow-water species described
prior to the present study, only 25 had not been recorded from any other province (3770
endemism).
The present results, concentrating on the shallow-water Poecilosclerida, generally support
these trends, although levels of endemism were substantially higher for this fauna than had
been predicted on the basis of Ltvi's (1979) preliminary analysis of the species composition.
These data probably reflect the high diversity of Poecilosclerida in shallow waters. Thirty
1298
J.
N. A. Hooper and C . Ltvi
species o f Myxillidae, Crellidae, Desmacellidae, Microcionidae and Raspailiidae are now
known from the New Caledonian region, including 13 new species and six new locality
records for the fauna provided here. Eighteen o f these species (60%)are apparently endemic
to this fauna: seven species are restricted to the deeper-water fauna ( o f which 5 are
endemic), whereas 21 are predominantly or exclusively shallow-water species ( o f which
13 are endemic). The non-endemic, deeper-water sponge fauna shows similarities to the
northern New Zealand sponge fauna, whereas the non-endemic species found in the shallowwater fauna o f New Caledonia also occur in both the temperate (Peronian and Maugean
provinces) and tropical Australian faunas (Solanderian province). In most cases, shallowwater species living in both New Caledonian and Australian waters represent the easternmost
extent o f species' distributions in the Indo-west Pacific, and, accordingly, many o f these
species were substantially different in some o f their characters. Furthermore, many o f the
endemic New Caledonian species were also usually immediately recognisable as (transformed)
sister-species o f Australasian species. More detailed biogeographic affinitieso f this fauna will
be discussed in a separate contribution.
A summary o f the New Caledonian poecilosclerid fauna (for the five families indicated)
is presented below. S , predominantly or exclusively shallow-water species (0-100 m depth);
D, predominantly or exclusively deeper-water shelf species (100-500 m depth); species
marked with an asterisk are presently thought to be endemic to the region.
Myxillidae (only 3 species have been described, all 'apparently endemic'):
Lissodendoryx bifacialis LCvi & LCvi, 1983b [Dl*
L. stylophora LCvi & LCvi, 1983b [Dl*
Acarnus caledoniensis, sp. nov. [S]*
Desmacellidae (only 1 'endemic' species has been described; the hypercalcified 'sclerosponge'
Merlia is also included here, after van Soest (1984b) and others]:
Neofibularia hartmani, sp. nov. [S]*
Merlia normani Kirkpatrick, 1908 [S]
Crellidae (only 2 species described, 1 'apparently endemic'):
Yvesia acanthosclera LCvi & LCvi, 1983b [Dl*
Crella spinulata (Hentschel, 191 1 ) [ S ]
Microcionidae (14 species are known, 10 o f which are 'apparently endemic'):
Antho novizelanica (Ridley, 1881) [Dl
Clathria (Clathriopsamma) rugosa, sp. nov. [S]*
C. (CI.) litos, sp. nov. [SI*
C. (Thalysias) vulpina (Lamarck, 18 14) [S]
C. (T.) flabellifera, sp. nov. [SI*
C. (T.) corneolia, sp. nov. [S]*
C. (T.) araiosa, sp. nov. [S]*
C. (T.) hirsuta, sp. nov. [S]
C. (Clathria) kylista, sp. nov. [S]*
C. (C.) bulbosa, sp. nov. [S]*
C. (C.) menoui, sp. nov. [S]*
Echinochalina (Echinochalina) intermedia (Whitelegge, 1902) [S]
E. (Protophlitaspongia) laboutei, sp. nov. [SI*
E. (P.) bargibanti, sp. nov. [S]*
Raspailiidae (7 species have been described, 3 o f which are 'apparently endemic'):
Raspailia wilkinsoni Hooper, 1991b [S]
Ceratopsion clavata Thiele, 1898 [S]
C. palmata Hooper, 1991b [ S ]
Aulospongus clathrioides LCvi, 1967 [S]*
A . gardineri (Dendy) [Dl
Plocamione pachysclera (LCvi & Lkvi) [Dl*
Lithoplocamia dolichosclera LCvi & LCvi [Dl*
New Caledonian Poecilosclerids
1299
Acknowledgements
This paper benefited greatly from the comments of an anonymous reviewer. 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 Barbigant, Jean-Louis Menou, and other staff of ORSTOM,
Noumea, for the hospitality, helpful assistance and cooperation during this project.
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Manuscript received 2 December 1992; accepted 14 April 1993