The marine fauna of New Zealand : Porifera: Lithistid demospongiae (rock sponges)

2007·5.8 MB·English

by  Kelly, M

ISSN 1174–0043; 121 (Print)
ISSN 2463-638X; 121 (Online)
TaihNourkou gria n
Cover photo:
Lithistid sponges on the surface of a boulder from hydrothermally active rungapapa Knoll in the Bay of plenty.
the blue nodules and chalice-like sponges are juvenile Reidispongia coerulea (phymatellidae), and the cream knobs
are Macandrewia spinifoliata (Macandrewiidae). the boulder was retrieved using a NIWA epibenthic sled towed at
160 m depth from rv Kaharoa in 2000.
photo: NIWA
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NATIONAL INSTITUTE OF
WATER AND ATMOSPHERIC RESEARCH (NIWA)
The Marine Fauna of New Zealand:
Porifera: Lithistid Demospongiae
(Rock Sponges)
Michelle Kelly
National Centre for Aquatic Biodiversity & Biosecurity
National Institute of Water and Atmospheric Research (NIWA)
Private Bag 109-695, Newmarket
Auckland, New Zealand
[email protected]
NIWA Biodiversity Memoir 121
2007
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Cataloguing in Publication
KELLY, M.
The marine fauna of New Zealand: Porifera: Lithistid Demospongiae (rock sponges) / by Michelle
Kelly—Wellington: NIWA (National Institute of Water and Atmospheric Research), 2007
(NIWA Biodiversity Memoir, ISSN 1174-0043)
ISBN 978–0–478–23277–6
Series Editor: Dennis P. Gordon
Copy edited and typeset by: Geoff Gregory, Word Therapy, Paraparaumu
Printed and bound by: Graphic Press & Packaging Ltd, Levin
Received for publication—30 June 2006
 NIWA Copyright 2007
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CONTENTS
Page
ABSTRACT .................................................................................................................................................................5
INTRODUCTION ......................................................................................................................................................6
THE NEW ZEALAND LITHISTID SPONGE FAUNA ................… ..................................................................7
MATERIALS AND METHODS..............................................................................……………………………… 8
Sample collection .....................................................................................................................……………… 8
Spicule dimensions .................................................................……………………………………………… 8
Registration of type and general materials ..........................................................................……………… 9
Area of study ...........................................................................……………………………………………… 9
Terminology .............................................................................……………………………………………… 9
Abbreviations of institutions ...................................................................................……………………… 10
CHECKLIST OF SPECIES ......................................................................................................................................11
SYSTEMATICS ........................................................................................................................................................12
Lithistid DEMOSPONGIAE (formerly order Lithistida Schmidt, 1870) .........................................................12
Family Theonellidae .....................................................................................................................................13
Genus Discodermia .....................................................................……………………………………… 13
Family PhymaTellidae ......................................................................……………………………………… 15
Genus Neoaulaxinia ...................................................................……………………………………… 15
Genus Neosiphonia ....................................................................……………………………………… 22
Genus Reidispongia ....................................................................……………………………………… 27
Family CorallisTidae ......................................................................................................................................29
Genus Neoschrammeniella .....................................................................……………………………… 30
Genus Herengeria ......................................................................……………………………………… 34
Genus Awhiowhio n. gen. ……………………………………… .........................................................39
Family neoPelTidae .......................................................................................................................................45
Genus Homophymia ...................................................................……………………………………… 45
Genus Callipelta ..........................................................................……………………………………… 47
Genus Neopelta ...........................................................................……………………………………… 50
Genus Lepidothenea ....................................................................……………………………………… 52
Family maCandrewiidae ...................................................................……………………………………… 53
Genus Macandrewia ..................................................................……………………………………… 53
Family Pleromidae ..........................................................................................................................................54
Genus Pleroma ...........................................................................……………………………………… 56
Family isoraPhiniidae ........................................................................………………………………………61
Genus Costifer ............................................................................……………………………………… 61
Family sCleriTodermidae ..................................................................……………………………………… 64
Genus Microscleroderma ..........................................................................……………………………...65
Genus Scleritoderma ................................................................................................................................67
Genus Aciculites .........................................................................………………………………………69
Family azoriCidae ..........................................................................................................................................76
Genus Leiodermatium ............................................................................……………………………… 77
DISCUSSION ...........................................................................................................................................................88
ACKNOWLEDGMENTS .......................................................................................................................................90
REFERENCES ..........................................................................................................................................................90
APPENDIX 1. Key to diagnostic characters ....................………………………………………………………94
APPENDIX 2. Key to diagnostic characters ....................………………………………………………………95
APPENDIX 3. List of stations ............................................………………………………………………………96
TAXONOMIC INDEX ...................................................................……………………………………………… 99
3
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Frontispiece:
Lithistid sponge Aciculites pulchra Dendy (Scleritodermidae), clearly indicated by the noduled surface, on the
surface of a boulder at 120 m off the southern edge of a reef off Great Barrier Island. This photograph was taken
from a Remote Operated Vehicle (ROV) during the Department of Conservation’s Great Barrier Island Deep Reef
Investigation in 2003.
Photo: Roger Grace, Department of Conservation, Auckland Conservancy.
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NATIONAL INSTITUTE OF
WATER AND ATMOSPHERIC RESEARCH (NIWA)
The Marine Fauna of New Zealand:
Porifera: Lithistid Demospongiae (Rock
Sponges)
Michelle Kelly
National Centre for Aquatic Biodiversity & Biosecurity
National Institute of Water and Atmospheric Research (NIWA)
Private Bag 109-695, Newmarket
Auckland, New Zealand
[email protected]
ABSTRACT
The lithistid Demospongiae fauna of New Zealand has been inventoried from existing and new collections, and is reviewed
here and revised where necessary. Most of the 282 specimens examined were recorded from the largest collection of sponges in
New Zealand, in the NIWA Invertebrate Collection, Wellington. Significant collections were also examined from the Museum
of New Zealand Te Papa Tongarewa. The lithistid Demospongiae (formerly order Lithistida Schmidt, 1870) is represented in
the New Zealand region by nine families, 18 genera (one of which is new to science), and 30 species (12 of which are new to
science): Theonellidae (1 genus, 1 species), Phymatellidae (3 genera, 6 species), Corallistidae (3 genera, 7 species), Neopeltidae
(4 genera, 4 species), Macandrewiidae (1 genus, 1 species), Pleromidae (1 genus, 3 species), Isoraphiniidae (1 genus, 1 spe-
cies), Scleritodermidae (3 genera, 5 species), and Azoricidae (1 genus, 2 species). This work records the first lithistid species,
Neoschrammeniella antarctica n. sp., known from polar regions, and provides the first record of the genus Leiodermatium further
south than the Philippines. Two additional species of Leiodermatium described here are found only in the west-central Pacific
and Micronesian deep waters, but are included for the sake of a complete review of the genus in the Pacific. New species
of the previously monospecific phymatellid genera Neoaulaxinia and Neosiphonia are described, and a new corallistid genus,
Awhiowhio, is recognised from New Zealand waters. All specimens were dredged from between 80 and 1700 m, but were
commonest between 200 and 800 m. With the exception of one specimen from the eastern edge of the Challenger Plateau on
New Zealand’s west coast, and a new species from the Ross Sea, Antarctica, all were found north of the northern edge of the
Chatham Rise and in New Zealand’s northernmost waters. Known and new species are redescribed from representative New
Zealand material and, in some cases, the characters used to define genera and species are redefined and clarified. In particular,
ornamentation of the desma skeleton and morphology of the augmenting microscleres are emphasised for distinction at the
species level.
Keywords: Porifera, lithistid Demospongiae, Lithistida, Theonellidae, Phymatellidae, Corallistidae, Neopeltidae, Macan-
drewiidae, Pleromidae, Isoraphiniidae, Scleritodermidae, Azoricidae, Desmanthidae, polyphyletic group, systematics, sponges,
taxonomy, new species, New Zealand
5
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INTRODUCTION
Mayer & Hamann 2004; Piel et al. 2004). A recent survey
Lithistid demosponges are a polyphyletic group com-
of the literature revealed some 40 papers reporting bio-
prising 13 extant families; 36 genera are included in
logically active compounds and their synthesis from the
the most recent classification (Pisera & Lévi 2002a) but
Southwest Pacific genera Aciculites, Callipelta, Discoder-
five genera remain of uncertain status. They differ from
mia, Microscleroderma, Neosiphonia, Pleroma, Reidispongia,
other demosponges in that the dominant structural spi-
and Scleritoderma. Of these genera, the first example of
cules (desmas) are articulate, forming in most species a
double bioalkylation of the sterol side chain at position
solid, rigid, heavily siliceous skeleton. These desmas are
26 was reported from New Zealand’s Aciculites pulchra
highly diverse morphologically; the overall architecture
(Crist et al. 1983). Furthermore, Pleroma menoui Lévi &
of the desma, the ornamentation of the desma surface,
Lévi, Reidispongia coerulea Lévi & Lévi, and Neosiphonia
and the pattern of articulation with adjacent spicules, are
superstes Sollas from New Caledonia were the subject of
diagnostically important (Schrammen 1910, see Kelly
considerable attention for their production of bromind-
2000a; Pisera & Lévi 2002a).
oles (Guella et al. 1989) and antiviral (Laille et al. 1998),
Lithistid sponges were traditionally placed in the
antifungal (D’Auria et al. 1995), and cytotoxic macrolides
single order Lithistida Schmidt, 1870 owing to the com-
(D’Auria et al. 1996; Zampella et al. 1997; Carbonelli et
mon possession of desmas, even though these display
al. 1999; Bassarello et al. 2000).
considerable morphological diversity. For some time,
Lithistid demosponges are of particular interest to
lithistids have been recognised as polyphyletic, with
paleontology because many extant genera and species
several points of origin within the Demospongiae (de
are relict from sponge faunas that were more abundant
Laubenfels 1936; Reid 1963, 1970; Kelly-Borges & Pom-
before the present. Lithistid demosponges are known
poni 1994; Pisera & Lévi 2002a). An indication of the
from the Ordovician, Silurian, Devonian, Permian, Late
polyphyletic nature of lithistid sponges is revealed in the
Jurassic, Late Cretaceous, and Eocene (Pisera 2002,
wide range of microscleres and ectosomal megascleres,
2006; Reid 2004). The Paleozoic fauna was dominated
and desma axial geometries, which include tetraxial,
by Orchocladina (extinct in the Permian) and sphaero-
monaxial, polyaxial, and anaxial forms.
cladinid sponges (solely represented in the Recent fauna
Familial and ordinal affiliations of lithistids with
by Vetulina stalactites from the tropical Atlantic). The
each other and with non-desma-bearing sponges are
Mesozoic (260–60 Ma) lithistid fauna was dominated by
uncommon but can be found in the megasclere and
Tetracladina, Megamorina, and Rhizomorina, groups
microsclere components of the skeleton (see Pisera &
that are still relatively common in Tertiary and Recent
Lévi 2002a). The presence of triaene megascleres and
lithistid faunas (Pisera 2002, 2006). These sponges fos-
asterose microscleres in some lithistid genera clearly
silise well because of their rigid silica skeletons; 13
indicates affinity with demosponge Astrophorida, and
suborders, 34 families, and more than 200 fossil genera
this has been supported by recent DNA studies (Kelly-
have been recently revised (Pisera 2002, 2006), and many
Borges & Pomponi 1994; Chombard et al. 1998; McIn-
more nominal genera are known.
erney et al. 1999). The exotylostyles of Gastrophanella
Lithistid sponges are known from almost all temper-
are considered to demonstrate affinity with the non-
ate and tropical oceans but are generally restricted to
lithistid demosponge order Hadromerida (Van Soest
depths greater than about 80 m in South Pacific locations
& Stentoft 1988).
including New Zealand and the Norfolk Ridge, and 150
A greater difficulty arises when taxa do not have
m in the tropical Western Atlantic region (Pomponi et
accessory spicules, or the spicules that they have bear
al. 2001). Apart from the proliferation of the genera Ac-
only superficial resemblance to those in non-lithistid
iculites, Neoschrammeniella, and Pleroma in the Southwest
demosponges. Species of Aciculites, for example, contain
Pacific, Corallistes and Discodermia in the western tropical
only acanthose anisostrongyles as accessory spicules,
Atlantic, and Theonella in the Indo-Pacific, most genera
and Leiodermatium and Vetulina possess only desmas
are known only from fragments or single specimens
and diactines, and desmas, respectively. In all three
that have been rarely, if ever, recollected (e.g. Lyidium
taxa, molecular evidence was required to indicate their
[= Pleroma] torquilla Schmidt, 1870 from Cuba). Lithistid
phylogenetic relationships (Kelly-Borges & Pomponi
sponges are rarely collected from below about 1700 m
1994).
depth.
The lithistid sponges are of interest to biomedical
Prior to the present work, two major regional fau-
science because of the great variety of pharmaceutically
nas were known worldwide: the continental shelf and
relevant biological activities of their chemical extracts
slope fauna of the tropical western Atlantic (Schmidt
(see Pomponi 2001; Bewley et al. 1998; Munro et al. 1999;
6
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1870, 1880; van Soest & Stentoft 1988; Kelly-Borges et 2005). In both locations, lithistid sponges dominate the
al. 1994; Kelly-Borges & Pomponi 1994; Lehnert & van sponge fauna (Lévi 1991; Reed & Pomponi 1997; Richer
Soest 1996; Pisera 1999; Pomponi et al. 2001) and the de Forges et al. 2000) between 150 and 1800 m but the
seamount fauna of the Southwest Pacific including the structure and taxonomic composition of the communi-
seamounts of the New Caledonian Norfolk Ridge (Lévi ties differ considerably (Pomponi et al. 2003).
& Lévi 1983, 1988; Lévi 1991; Schlacher-Hoenlinger et al.
THE NEW ZEALAND LITHISTID SPONGE FAUNA
Prior to 1991, when research on the lithistid fauna of tetracladine lithistids (which are comparable to those
New Zealand commenced, only two Recent lithistid of the Recent family Phymatellidae) and described a
species had been described, although several more new species of Vetulina, V. oamaruensis, based upon
were known (P.R. Bergquist pers. comm.). Arthur the sphaerocladine desmas. These desmas strongly
Dendy described the first lithistid sponges from New resemble those found in a species of Crambe from
Zealand―Aciculites pulchra Dendy, 1924 and Lepi- Spirits Bay, Northland and their conspecificity cannot
dothenea incrustans (Dendy, 1924) from the vicinity of be discounted (Kelly et al. 2003).
North Cape and the Three Kings Islands, respectively. Whole-body fossils of lithistid sponges are also
Bergquist (1968) redescribed this material but recorded known from the mouth of the Kakanui River in the
no further species. No further species were added by Oamaru district (Kelly et al. 2003). These fossils occur
Dawson (1993) in his comprehensive index to the New in a volcaniclastic Ototara Limestone bed of lower
Zealand Porifera. Whaingaroan age just above the top of the Mineral
From 1991 onwards, many of the lithistid species Breccia Member of the Deborah Volcanics (31.6 Ma)
described by Lévi and Lévi (1983, 1988) from the south (Dickey 1968; Daesch et al. 1970). The body fossils were
New Caledonian slope and seamounts were progres- found to be very similar morphologically to the living
sively discovered in the New Zealand region (Kelly et pleromid sponge Pleroma aotea Kelly when compared
al. 1999; Kelly 2000b; Kelly 2001a,b; Pomponi et al. 2001; morphometrically to known extant sponges from New
Kelly 2003; Kelly et al. 2003; Kelly & Buckeridge 2003; Zealand and New Caledonia. Kelly et al. (2003) consid-
Kelly 2004; Kelly & Tubbs 2006; Kelly et al. in press), ered them to be conspecific with living sponges from
along with several undescribed species. A voyage to deepwater seamounts and banks off northeastern New
the seamount region south of New Caledonia in Au- Zealand, citing several additional examples from the
gust 1999, courtesy of IRD, Nouméa, assisted greatly substantial record of lithistid sponges in the Oamaru
in the author’s understanding of the faunal relation- Diatomite.
ships between the two countries and of lithistid sponge Lithistid microfossil spicules and partial body
ecology in general. fossils are also known from the Tutuiri Greensand
Homophymia stipitata Kelly, 2000a, discovered in (Teurian–basal Waipawan) in the Chatham Islands
New Zealand waters, was only the second known spe- (Buckeridge in Campbell et al. 1993; Buckeridge &
cies of the genus, which was known previously only Kelly 2002; Kelly & Buckeridge 2005). The spicules
from a single species from Madagascar and Réunion. are trapped within the skeletons of fossil hexactinel-
Also recently recognised in national collections was lid sponges. These are remarkable for their diversity,
Pleroma aotea Kelly, 2003a, the third species to be and the combination of lithistid, astrophorid, and
described in a genus principally known from New hexactinellid taxa indicates a paleoenvironment very
Caledonian waters. similar to that found today at depths of 500–800 m on
Prior to the work of Dendy (1924), Hinde and the Chatham Rise (Buckeridge & Kelly 2005) and in the
Holmes (1892) described a species-rich assemblage tropical Atlantic (cf. Pomponi et al. 2001).
based on siliceous spicules from marine diatomaceous Reasonably well-preserved sponge body fossils
sediments now known to be from the early Runangan resembling lithistid Corallistidae, Isoraphiniidae,
(Late Eocene) horizon within the Oamaru Diatomite Pleromidae, Phymatellidae, and Scleritodermidae,
Member of the basaltic Waireka Volcanics (c. 35 Ma) at have also been found in the Red Bluff Tuff (Teur-
Oamaru in North Otago (Suggate et al. 1978; Edwards ian–Waipawan) on Chatham Island (Buckeridge in
1991). Amongst the many non-lithistids represented Campbell et al. 1993; Buckeridge & Kelly 2006), provid-
were microfossil spicules of what were considered ing an interesting record for this group that straddles
to be a species of Lyidium [= Pleroma], and species of the Mesozoic–Cenozoic boundary (Maastrichtian to
Corallistes, Discodermia, and Theonella were illustrated. early Ypresian) (Kelly et al. 2006).
Hinde and Holmes (1892) also illustrated desmas of
7
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MATERIALS AND METHODS
SAMPLE COLLECTION Specimens were either frozen immediately upon
collection or preserved in 50% isopropanol and pre-
Sponges were collected by rock dredge from north-
pared for histology and scanning electron microscopy
eastern New Zealand waters by the National Institute
as in Kelly-Borges and Vacelet (1995). Specimens are
of Water & Atmospheric Research (NIWA) research
stored for the long term in 70% ethanol.
vessels Tangaroa (1967–1989) and Kaharoa (1998) (Fig.
1). Specimens of known New Zealand species collected
SPICULE DIMENSIONS
by the author on the IRD cruise LITHIST (RV Alis,
August 1999) on the New Caledonian Norfolk Ridge
Whole desmas are notoriously difficult to measure
seamounts have been included in the study insofar as
because of their intimate articulation with adjacent
they represent a natural northern extension of the range
spicules and their frequent irregular morphology. The
of the New Zealand species. NIWA, NMNZ, and IRD
size of the desma is expressed here as the approximate
station data are listed in Appendix 3; only the station
size of the overall spicule, and as the mean clone length
name is listed in the sections describing material exam-
and thickness. All spicule measurements are expressed
ined. Data for other stations are listed in the text.
in µm. Microsclere dimensions recorded in this work
0 0
4o·s
~
(l
!i'
"')
_)
c,
"
~-
-;,f (>
50-S
1so·E
Figure 1. Locality maps, showing the
collection stations for rock sponges in
New Zealand and Antarctic waters.
8
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