Revision of Microcionidae (Porifera: Poecilosclerida: Demospongiae), with description of Australian species

626 Pages·1996·243.4 MB·English

,
REVISION OFMICROCION1DAE (PORIFERA: POECILOSCLER1DA:
OEMOSPONGIAE), WITH DESCRIPTION OFAUSTRALIAN SPECIES.
JOHN N.A. HOOPER
Hooper, J.N.A. 1996 07 01: Revision of Microcionidae (Porifera: Poecilosclerida:
Demosfwini^iac),withdescriptionofAustralianspecies.Memoirsof(heQueenslandMuseum
40: -626. Brisbane ISSN0079-8835.
1
A phylogenctic revision ofthepoccilosclcrid Microcionidae is based on type material, the
worldwide literature, and comprehensive Australian collections. Of73 available generic
names7 generaand ]7 subgeneraarerecognised here. Of561 availablespeciesnames455
areconsideredvalid(1 virtuallyunrecognisable),including52newspecies.TheAustralian
fauna, including Australian Antarctic Territory, contains 14S species (31 new), many new-
records, mostarenewcombinations,andmany illustratedforthe firsttime.Asynonymy of
worldspeciesisprovided.Validtaxainclude:Clalhria(with7subgenera:C (Clathria)(wkh
31 Ausiralian species, S2 other species worldwide, with new species rmtrphyi, noarlun-
gae,biclathratatborealis,burioni,sarai.sarospinifera). C. (Wilsonella)(6,8,abrofhosensis,
ensiat, lindgrem spp. nov.), C. (Mtcrociona) (5, 91. UUnvorme. Itzardensis. sinwt,
hnmdstedi. campecheue. ciaudei, hentscheli, leighensis, stephensac, tunisiae, urizae,
C
lurrlfiiirt spp. nov.), {Dendnnia) (7 speciesendemic to Australia), C. (Axot lella) (6. 6,
pCatu(lTaha.lfyrsoimaosn)fa(e3.6,ge5o3r,giaapehnyslilsa,spcpr.ansopve.d)i,aC^.d(ahrocwiienlelnas)i(^4,f1u.ssreelranvah.iah*astkhinaansnpip,.nhoevs.p)e,rainad,
temalolae. phorbasiformis, sryloprothesis, fingens, wcsselensis, amiranteiensts, herh/eh
spp. nov.); Antho (with 3 subgenera: A. (Antho) (12, 10), A. (Plocamia) (2, 17) and A,
ilsopentctya) (3.1.punuea, saintvincenti spp. nov.); Echinoiiathria (14, 15. bergquistQe,
levii, notialis,parkeri. riddlei spp. nov.); Holopsamma (9 species endemic to Australia, I
indeterminate species);Echinochulina (with 2 subgenera: E iFjiiinochalina) (10, 2, felhi
sp. nov,), £ (Protttphlifospongio) (8 species endemic to Australia and New Caledonia,
callata, favulosa, isitaci, tuberosa spp. nov.)); Artemisimi (4, 10); and Pandoras (incertae
sedis) (0, 2"), Generic keys are provided. Morphometric characters of primary importance
include the origin, geometryanddistribution ofstructural meg3scleres within the skeleton,
modificationofmegasclercsto monactinal ordiactinal forms, the presenceorabsenceofa
specialised ectosomal skeleton, presence ofdetritus incorporated into spongin fibres, and
overallskeletal structure(includingcompressionoftheaxialskeleton anddifferentiation of
axial and extra-axial regions). Briefzoogeographical comparisons are made between con
tinental AustralianandadjacentIndo-westPacificfaunas. Australianspeciescompriseabout
32% of the world's microcionid diversity; about 75% of species arc endemic for the
Australianregion,andtemperatespecies(81%.)havehigherlevelsnfendemismthantropical
M
\pccies{59%).£}Porifera,Demospongiae, Poadosiierido, icrocionidae.familyrevision
netyspecies, Taxonomy, biogeography, Australia,
JuhnN.A. Hooper. QueenslandMuseum. POBQX33QQ, South Brisbane, Queensland.410}
.
Australia; received I December 1995.
INTRODUCTION Within the IndtvAustralian region microaonids
arc particularly abundant, with some species
....
dDecMsmc.orcsirpboeocdnieo(imIadxeat,aeM),csooPmonpreriifoesf-ria,tnhgesp,laeacbriogeeusstt(cf8Ha%omoil.p.oi.efersao&ll.. bm,eaai"cnngma,bd<co™mmdhionoSatvn-hTterpc.lor.iAcmtvpeirooantueusrnetwcsooronkftstah.edncmsshgcarclixltbor,iawn-glw.,a.rtrth,ei,r-,
Levt. 1993ft). The family has contained al one <al rccords ° Australian spec.es) ,nc ude:
time oranotherabout 70 genera and 550species. Lamarck (1814, 1815, 1816) Gray ( 1858, 1867,
although fewerthan these arc now recognised as ,869- 1870>- Bowerbank (1864, 1875, 1877), Bar-
valid. The family has a worldwide distribution nard (1879), Kent (1871),Ridley ( 1884a), Ridley
and it is found from the intertidal zone todepths & Dendy (1886, 1887), Lendenfeld (1888. 1889a),
exceeding 2000m. It is clearly one of the more Kieschnick (1896. 1900). Thielc (1898, 1899,
important, ecologically successful groups of 1900, 1903),KirkpatricM1900a,b),Whitelegge.
Porifera. (1901, 1902). Hcntschcl (1909, 1911, 1912),
2
)
MEMOIRS OFTHEQUEENSLANDMUSEUM
Kallmann (1912, I9l4a-e, |yi6a-c. 1920). necessity for dehydration through an alcohol
D1e93n0d/y19&32,Fr1e9d3e3r)i.cBkur(t1o9n24()I,93T4cap)s,eBnetrgq1u1i8s9t7b&, sweerriees.mSaodmeefomriceraocthomsepesceicetsi.onFsracugtmeantt3s0-w3e5rme
Tizard(1967). KeJIy-Borges&.Bergquist(198B), passed through a dehydration series, cleared in
Be9rgqmuist & Froinont (1988). Wiedenmayer toluene or Histosol, and wax embedded for at
a Hooper (1990b). Hooper et al. (1991, least 2 hours. Sections were cut from trimmed
1992), Hooper & Levi 0993a. 1994). A brief wax blocks (cutting from the centre ofthe block
synopsis of the fauna is given by Hooper & to the exterior so as to include both the outer
Wiedenmayer (1994). although some of the surface and inner skeleton relatively intact),
nomenclature and synonymies contained in that placed in clearing agent foran adequateperiodtu
earlierwork are revised here dissolvewaxand/ordewaxingonahotplate,then
Priortothepresentstudymorethan200species soaked in etbanol until perfectly clear, floated
of Mierocionidac had been described in the onto albumen-coated slides, orientated and flat-
Australian fauna(including itsterritorialwatersi, tened, stained with basic fuccsin and mounted.
but many of these were found to be either com- Fragments ofdry specimens (e.g.. type material)
posite (consisting of several sibling species;', or were reconstituted in5% bufferedformalin for 1
synonyms of other species. This study 1) hours, which produced rehydration of the
describes 14* species (31 new), many new- mesohyl and enabled cleaner histological sec-
localityrecords forAustraliaandnew taxonomic tions to be made.
combinations. 2) provide* an annotated
MoRPHOMhTKrc Analyses. Spicules were
synonymy for 3II other species worldwide (in-
cluding 21 new species); 3) revises the mor- measuredwithaslage micrometer,eitherdirectly
phometry* characters used for classification and through a microscope or computer digitiscr.
population variability for particular species; and Twenty five spicules,ofeachspiculecategory', in
4-) determines levels of endemism amongst all specimens were measured. Acanthostyle
width measurements were taken immediately
provincial faunas.
below the base. Toxa lengths refer to choixl
MATERIALS ANDMETHODS length;isocheiacaremeasuredfromapexofalae;
width measurements of other spicules fefc
maximum width.
COLLECTION AND HISTOLOGICALTECH-
Spiculedimensions were sortedandstatistical-
NIQUES Material examined in this study was ly compared forvariousparameters(e.g., Season,
predominantly collected using SCUBA (0-40m locality, depth)* including one* and two-way
depth) or dredging anil uawls (3O»360m depth). ANOVs with replication, two-way ANOV with
Seasonal sampling for reproductive periodicity unequal replication, means differentiated using
a.is conductedover Iwn years in the Darwin and two-tailedttests.Line-drawingsweremadeusing
Cobourg Peninsula regions, NT. Immediately a calibratedcamera lucida, and microphoiographs
aHer collection specimens were either fixed in were taken withan Olympus microphotosystem.
80-100% methylatedcthanol or frozen (which 10 Taxonomic keys were constructed using ordered
some extent fixes the pigments), and later multistats disordered multistats roorphometnc
preservedin70%alcohol.Reproductiveproducts and binary character utilising the DELTA sys
were searched forin fresh or frozen tissue. tem(Da)lwitz& Paine, 1986),
Nitricacid spicule preparations, thick-sections
and thin-seciion mounts iveft iruimely made as SCANNING ELECTRONMICROSCOPY. Sec-
follows. Fragments of each sponge, including tions were prepared as follows:
ectosomalandchoanosomalregions,wereheated 1 Cut at 1-1.5mmthick,ensuring thatboth ihe
directly on a glass microscope slide in several cctosomc and choanosomc were represented.
drops in nitric acid (the solution was evaporated 2) Placed in a cavity block and covered with
nither than boiled, using low heat), and mounted several drops of sodium hypochlorite toetch the
in Canada balsam once completely dry, and mesohyl matrix from the skeleton. The etching
coujed. Thick, hand-cut sections were made per- process was monitored through a dissecting
pendicular to the surface, soaked in a saturated microscope in orderto prevcnl the skeleton fall-
solutionofphenolandxylene(forapproximately ing apart. Delicate structures (plumose,
24 hours), and mourned in Durcupan (ACM hahchundroid. hymede-smoid skeletons) only re-
FlukaPnidu<'i\)usrnggbvsslivcrxv>r<:arthoiaise quired a few seconds treatment with bleach;
the GOVertlip level Phemd-xylene precluded the robust skeletons (reticulate, fibrous^ articulated
REVISIONOFMICROCfONJDAE-:
skeletons)required several minutes: bul general- 6) These steps were repeated if any coll
ly 30 seconds was adequate. remained.
3) Bleach was pipetted off at the appropriate 7] Spicules were resuspended firstly in
lime and 70% ethanol immediately added. Sec- demineralised water, 70% ethanol, then two
tions were left to stand for several minutes to series of 100% ethanol solutions, oentrifuging
ensure bleachwas completelyneutralised. and decanting the supernatant between each
4) Steps 2-3 were repeated, without removing changeofsolution,finallyendingwithsuspended
section from cavity block Substituting con- spicules ina solution ofabsolute ethanol.
centrated hydrogen peroxide in place ofsodium 8) Amicro-coverglasswasadheredtoanSEM
hypochlorite, finally rinsing in etham.il The stubusingdouble-sidedlaporcopperdag.several
hydrogen peroxide slep was omitted foi very dropsofsuspendedspiculesplacedontothecover
delicate sections. glass, the alcohol-spicule solution ignited and
5) Sections were placed on clean microscope spreadacrosstheglasswithaglassrodorforceps
slides and letdry completely. until all ethanol was vaporised. Spiculesbond to
6) Sections mounted on SEM stubs using glass relatively firmly, bulexcess spicules could
double-sided tape, copper dag. or 'Supa Glue* be blown off glass using compressed air, or
(Supa Glue, Selleys Chemical Company. spread out over the glass by adding further
Padstow). An alternative method used to fix ethanol and igniting. The distribution ofspicules
samples to stubs was to cover stub with on the cover glass was monitored under com-
'Aquadhere' wood glue (Aquadhere. Scllevs pound or dissecting microscope (magnification
Chemical Company, Padstow), letdrycomplete- dependingonspiculesizeK Moredropsofspicule
ly (usually Severe! days), then prior to use ex- solution added and this slep repeated if too few
poseddrygluetovigoroussteam(whichsoftened spicules were present, ensuring not toovercrowd
the set glue), and placedthe section on top ofthe field of view lor SEM photographic purposes.
stub(it would sink in a shortwaybutwasbonded 9) An alternative method was used to produce
reasonablywelltothestub,andhadtheadvantage a perfectly smooth background, UStrig an
ofproducing aperfectly smooth background). *Aquadhere* glue-coated stub, dried for se
7) The stub was sputter-coated well to ensure days then softened with steam, and sfpici
thai all fibres were well coated to reduce placed directly ontosoftglue tin thiscaseethanol
'charging'. In some cases uncoated sections was no* burnt bul evaporated). Single spicules
could be viewed successfully under low ac- would sink intogluetoofarifitwasloosoft (i.e.,
celeratorvoltage, butbetterresults were general- leftin steam too long).
ly obtainedon coated specimensat higher voltage. 10) Spicule coated >tubs were sputter coaled
Typical viewing conditions used were 25kV, at briefly and viewed id 25kV, minimum working
close working distance lo provide best depth of distance and smallest apperture for best resolu
field and focus, and at low magnifications. lion.
Spicule were prepared as follows:
1) Thinly cutsections including both cctosmc ABBREVIATIONS
and choanosome were placed in a durham tube
(micro-test lube), to which drops ofconcentrated AAT, Australian! Antarctic Territories; ABIP,
ftilric ftcldarcadded, usingdrop-hy-dropaddition Ccntro dc Esiudios Avan/ados dc Blanes. In-
soastocontroltheoxidationreactionandproduc- stitute dc Invcstiguciones Pesqueras Barcelona,
tion of by-product oxides. Aquarium dc Blanes.Gerona. ABRS, Australian
?) Upon completion ofacid digestion the dur- Biological Resources Survey, Canberra, AFZ,
ham tube was half filled with acid and gently Australian Fishing Zone; AHR Alan Hancock
heated over an alcohol Ilame, ensuring thai onlv Foundation, University of Southern California,
smallbubbles form (lowheat, noboiling), for 1-2 Los Angeles. AIMS. Australian Institute of
minuics. Marine Science, Townsville; AM, Australian
3) Solution was let stand lo cool, then Museum, Sydney; AMNH, American Museum
ccntrifuged (approximately 40O0rpm for 30 of Natural History, New York; BMNH, The
seconds). Natural History Museum, London; BPBM, Ber-
4) Nitricacid waspipettedoffleavingaspicule race P. Bishop Museum, Honolulu; CP,Cobourg
mass at the bottomof the tube, undisturbed. Peninsula, NT; CPMNP, Cobourg Peninsula
5) Spicules were resuspended in fresh nitric Marine National Park. NT; C5IR0, Commotl
acid Hid gcitify stirred usingclean,fine,gla^srod wealth Scientific and Industrial Research Or-
MEMOIRS OFTHEQUEENSLAND MUSEUM
ganisation, Marine Laboratories, Hobart, Queensland Museum, Museum of Tropical
(leveland and Penh; DAR, Darwin region, NT; Queensland, Townsville: NCIQ66C-, United
DELTA, Description Language for Taxonomy States National Cancer Institute, Australian In-
computer software (Dallwitz & Paine, 1986); stitute of Marine Science shallow water collec-
EIS, Environmental Impact Study;CSIROEMG, tion contract (1984-91), Townsville (primary
CSIRO Food Research Laboratory. Division of voucher samples now lodged in QM, others in
Food Processing, North Ryde. Sydney. EMU, NTM and USNM); NCIOCDN-. United States
Ensight (formerly Environmental Management National Cancer Institute, Coral Reef Research
Unit), Water Board (Sydney, lilawarra, Blue Foundation shallow water collection contract-
QM
Mountains), Sydney; EPA. Environment Protec- ChuukState (vouchersampleslodgedin and
tion Authority, Sydney; EPALR, East Point USNM); NEQ, northeast Queensland (Bowen to
AquaticLifeReserve,DudleyPoint,DarwinHar- Cocktown); NM, Natal Museum, Pietei
bour, NT: FNrQ, farnorthern Queensland (Cook- marit/tMirg, NMB, Naturhistorisches Museums
town to Torres Straits); FUB. Freie Universitat zm Basel, Basel; NMCIC, National Museum of
Berlin; GBR, Great Barrier Reef, Queensland, Natural Sciences, National Museums ofCanada,
HNLTK, Natural History Museum, Flam Nam Ottawa; NMNZ. National Museum of New
University. South Korea; ICBUC. Institute] Zealand (formerly Dominion Museum), Wel-
Centralde Biologia,UniversidaddeConception, lington; NMV, Museum of Victoria (formerly
Chile, ICZK, International Code of Zoological National Museum of Victoria), Melbourne;
Nomenclature (sec Anonymous, 1985); 1M, In- NSM, National Science Museum. Tokyo, NSVv.
dian Museum (Zoological Survey ofIndia). Cal- New SouthWales; NT,NorthernTerritory; N
cutta; IMZUB. Islitulo c Museo di Zoologia ed NorthernTerritoryMuseumofArtsandSciences,
Anaconda Comparata dclla Univcrsita dt Ban, Darwin, NTU. Northern Territory University,
Ban; 1MZUN. Instituto e Museo di Zoo Darwin; NWS. Northwest Shelf region, Wcsicm
deirUnlversititdi Napolt,Naples; fNM.Nal ralia; PAUP, Plvyloeenetie Analysis Using
Museum nf Ireland, Dublin; IZUG, Muscn Parsimony (seeSwofford, 1991);PIBOC,Pacific
cp di Stona Na'.urale di Genova, Genova; Institute nf Bio-organic Chemistry, Far
JCU, James Cook University of North Scientific Centre, Academy of Sciences of the
Queensland Townsville; KFAU. Zoologiscben USSR.Vladivostok;PMJ,PhyletischesMuseum,
imlung der Universitii! Erlungen-Numbcrg, Jena, PNG, Papua New Guinea; QFS,
L'rlangen; LFM, Merseyside County Museums Queensland Fisheries Service, EVepartment of
(formerly Liverpool Free Museum), Liverpool; Primary Industries. Brishane and Caims, QLI),
LMJG, Abteilung fur Zoologic am Landcs- Queensland; QM, Queensland Museum, Bris-
nii.cum Joanncum (1 Museum Jubileum bane; QVML, Queen Victoria Museum and Art
Graz), Graz, MABA. Museo An ninn de Gallery, Launceston: RMBS, R0$COiT M;
t (
cias Naturales 'Bernardino Rivatfavia*, x»ucnns Biological Station. Roscoff, Fftncc; RMNI1,
Aires; MCNP,Div. Invest,del MuseodcCicnt:us Rijksmuseum van Natuurlijke Historic video.
I
Naiurales de la Pk>i». Argentina. MCZN, RRIMP. Roche Research Institute of Marine
Museum of Comparative Zoology, liar Pharmacology, Sydney (discontinued; sponge
University, Cambridge iMa- -..:-; MEQ, mid east collectionsnowheldinAM); RSME, Royal Scot-
fern Queenst;:; Lstune to tfowen); MHNG. tish Museum. Edinburgh; SA South Australia;
Museum d'Histoirc Nature]In de Gen4 SAM,SoiuhAustralian Museumt.Adelaide;SEQ,
Geneve; MLUM, Marine Laboratory of the southeast Queensland (Tweed River to
v crsity of Mfjiitii. Miami; MMBS, Gladstone). SM.Muscc Zoologique, Strasbourg;
Mukaishima Marine Biological Station, Faculty SME, Station Marine d'Endoume, Marseille;
of Science, Hiroshima University, Onomrehi; SMF, Natur-Muscum und Forschungsinstitut
MNI1N, Museum National d'llistoire Naturelle, Senckenbcrg, Frankfurt; TAS, Tasmania; TM.
Laboratoirc de Biologic des lavatebres Marins Museo t Istiiuto di Zoologia Sistematiea dclT-
et Malacologn Puna <DT. Topsenl collections. Universiu di Torino,Torino: TMAG.Tasmanian
.
DCL, Levi coJJectioas; DJV, Vacclet collections; Museum and Art Gallery, Hobart; UAZD,
DNBE, Boury-Esnaull collections); MOM, University of Auckland, Zoology Department,
MMuRnAcCc,OeKeoanniognrkalpihjiktnMirusdeeuMmOVvMoCaOi, MMi.>didi;eknh-, UAuCcTk.laSonudt;h AUfBr,icaIn'bMeuis-reeumMuosfeNautmu,ralBHriesmtoerny..
Afrika, Tervuren; MRHN, MU9& Royal d'- CapeTown; VQ, UniversityofQueensland, Briv
Histoire Naturcllcdc Bclgiquc, BruxcIIcs; MTQ, bane; VSC, University 0* Southern California,
REVISIONOFMICROCIONIDAE
Los Angeles; USNM, National Museum of (1983-1989) and Heritage Commission of the
Natural History, Smithsonian Institution. Northern Territory (Darwin) provided additional
Washington DC; UZM, Zoologisk Museum, fundingto visitremotelocalitiesandthescattered
Universitetsparkem Copenhagen; VIC, Victoria; collections in Australian museums.
WA, Western Australia; WAM, Western I amparticularlygrateful toLeonie Hooperfor
Australian Museum, Penh; YPM, Peabody linedrawings andJohn Kennedy formany ofthe
MNuesweHuamveonf(NCoantnu.r)a;lZIHiLs,toZroyo,loYgaiclaelUInnistvietrustietyo,f WSEeMbsb.(1NTaUl)so. tahnadnkGiBvoebWHialkridnyso(nUQ()A,IMChSa)rlfeosr
Leningrad, Academy of Sciences Museum of providing assistance with SEM photography
Zoology, St Petersburg; ZMA, ZooJogisch For financial or logistic assistance with
dMaums;euZmM.BU,nivMerussiteeuitmvafunrANmasttuerrkduanmd,eAmasntedre-r Vfiaekl.ddwyoarkKrIasaocckhnino,wleVdgYea:koGveleovr,geUSElSvRakoRvV.
ZHuomobloolgdits-kUniMvuesrsehuamt.zuCoBeprelinnh.aBgeernli;n;ZZMMHC,, 'Akademik Oparin* fPlBOC); Alice Kay. Lester
Cannon (QM) and Queensland FisheriesService;
Zoologiseht. tute und Zoologisches Mick Ready (FV 'HydronauO: Peter Murphy,
ZMuMsUeUu,m dUeprpsUanlivaersUintiavterHsaimtbeutrsg.ZooHlaomgbiusrkga; Martin Riddle, Shirley Sorokin. Rob McCauley
and other members of the NCI team (AIMS);
Muscct, Zoologiska Institute!, Uppsala; ZSN. Danny Roberts (EPA); Scott Chidgey, Calwell
Aquarium c Muse© della Slazione Zoologies di Connor and Associates: Patricia Byers (FV
NSatpooclkih,olNma.ples; ZRS, Zoologiska Rijkmuseum, T'rSekevloertoWnar)dB(i!CISRIuRdOmFainshe(rAiMes).;RLVm'PSooien!car\aRnVd
'Sprightly", FV Clipper Bird*, RV Southern
ACKNOWLEDGEMENTS Surveyor*); Martin Riddle and Lisa Miller
(EMU.C Rob Cupon (University ol Melbourne);
Patricia Bcrgquist (UAZD) encouraged this Alan Butlei (University ofAdelaide);ClayBrycc
siudy and provided UwixaUon to pc«eveiifi [WAM); Neville Coleman (Australian Marine
ihc long learning curve associated with sponge Photographic Index, Briibanej; Conservation
taxonomy. Felix Wicdcnmaycr (NMB) provided Commission of the Northern Terntory. Darwin;
access to his numerous unpublished personal Dave Ramra and Anne Coleman
notes on museum collfi Sponge Arch: INTFisheriesDarwin);Neil Smit(NTUniversity
NMV; Wiedenrnayer. 1989; Hooper* Wieden- Darwin) Barry RussellandHelen LarsoniNTM);
mayer. 1994). Rob Van Soest iZMA) provided Lu INMV); C6cite Debieus, George Bar-
numerous discussions on sponges, alternative gibunl, Je;in 1-onis Mcnou> Pierre Laboute
views on the diagnostic importance and polarity (ORSTOM Noumea); Pat and Lori Colin ("NCI
ofcharacters, andpossible relationships between CRRF Chuuk and Palau); and Tan and Pam Low
igc groups. Michelle Kclly-Borgcs (BMN1I) [FV 'Rachel'),
andPeterJellCQM)l>rovi(Iedn»an) positive, lamgrateful forcompetentfield assistance and
menis on the manuscript. laboratory technical assistance, during various
(MINIiiImN)pafrotricpurloavrildyinggraatepfouslt-tdooctColraau!defelIlow- parts of this sHtoudbyh,s,froAmleJnodiHeoBwaaxrtedr,. CStaeivheyn
ship at Ihc MNHN and giving mc access to its Johnston, John Kennedy. Daniel Loy Choy,
lypc collections, including all the Lamarck Anne-Marie Mussig, Paula Tonkins and Rex
material, the Sir Winston Churchill Memorial Williams 1 alsothank Phil Aldcrslade (N
Trust (Canberra), the Australian Biological assistance in developing computer digitising
Resources Study (Canberra), Klaus Riitzlex software,and Russell H.mley,NTM)foridentify-
(USNM). Willard Hartman (YPM), Joachim ingcommensal polychactcs.
Rcitner (HUB. now Gottingen) and the Trustees 1alsothankmanypcoplcforprovidingm<ilcirjl
of the Queensland Museum (QM) for providing forexamination, or for otherinformationcitedin
grants af various times, enabling me toexamine the text; Pennv Barents (AM), Nicole Bnury-Es-
major Museum collections and to interact with nauli <SME}.' Beatrice Buxch (BPBM). Susan
colleagues at several international forums (fa^m Chambers(RSME).FrankClimo(NMNZKRuth
which many of the ideas in this present volume Desqueymux-Fauudez (MHNG and LMJG),
matured). The Northern Territory University JaneFromonKJCU;. Manfred Grasshoff(SMF),
Planning Amhority (1983-1985). the Museums Jan Den HarTOg (RMNH), Takomura Hoshino
and Art Galleries Hoardofthe NorthernTerritory (MMBSfc Prank -.on Knorrc (PMJ). Dciter
MFMOIRSOFTHEQUEENSLAND MUSEUM
Kuhlmann (ZMB), Vladimir Krasochin when used alone skeletal architecture is not
(P1BOC). Romelv UwSkyoi (Cootamundra necessarily a reliabLe indicator of relationships.
lis Survey team, UK). Susan Boyd (NMv Hooper (1991, herein) noted that the so-called
Liz McCaffrey (UQ Brisbane), A.K. Mandal typical, compressed, axinellid-like skeleton of
<IM). Loisclte Marsh iWAMk C. O'Riordan many Raspailiidae (Raspailia puuiatifidar,
(INM), Shane Parker (SAM). David Parry (NTU Hooper, 1991; Fig.2b) also occurred in C
Darwin). Urs Rahm (NMB), Martin Riddle {Axociella) (Figs 7G, 119A). It ts also probable
(EMU), Frank Rowc (AM). Klaus Rui/lcr and thatskeletalstructureis influencedtosomcextent
Kathleen Smith (USNM), Shirley Stone by environmental conditions, and there is & -n»;
(BMNH'i. B.R. Sluekcnberg (NM), 0k> Tcndal evidence to xuggesi that flexible, compressed
(ZMC;, Jean Vacelet (SMH). Clare Valentine axial skeletons are produced in response io high
fBMNH). Clivc Wilkinson (AIMS), and energy environments (e.g., PalUtttbi. 1984),
WoUgangZeidterfSAM). Similarly, skeletal characters suchaschose found
in encrustingspecies have obviouslyevolved in-
DEFINITION OFCHARACTERS dependent]] in many (otherwise unrelated) tax*
Review of microcionid skriciji siiuclurcs
MINERALSKELETON.Theform,composition showedthat specieswhichweresimilarinspicule
and division of the skeleton remains the most geometry had different skeletal architectures
important character for classification of (Hooper." <H
I
Dcmospongiae. Recent attempts at higher sys- Architectural typesamongsttheMkrocicmuhie
tematicsofDetno\pongiacbased on non-skeletal include: Lhymcdesnnml (with thin layerofba.^il
&
characters leg., Simpson, 1968a; Bergquist .p-i'igin lying on the substrate containing erect
Mailman, 1969: Bcrgquist, 1980a; Lee & megasdere; IRg. 7H); 2 microcionid (with a
Gilchrist 19S.S; Hooper ct at., 1992) have had compressed basal spongin,t producing ascending
only limited success because in many instances fibre nodes and plumose spicule columns) (Fig.
they axe unable to corroborate all skeletal and 100F); 3. renieruid reticulate (consisting or a
iion-skeletal evidence into a single systemaiKs. rectangular reticulation of uni- or
In some cases amongst Demo.spongiae (e.g., paucispiculartractsfullyenclosedwithinspongin
Verongida), non-skeletal evidence has heen fibres or cemented at their nodes by loose col-
decisive and to some extern well correlated with lagen) (Fig. 7C); 4. isotropic reticulation (a dis-
othercharacters. In other cases (e.g., Axinellida) orientated, random uni-, pauci- or multispicular
that evidence has merely highlighted inade- reticulation in creel or massive forms, in which
qmuoarcpihesoliongsyys(tSemiamtpiscoan,bas1e9d68sao;lelByerognqusikeslteta&l there is nodistinction between primary orseCOri
dary tracts (not figured; seen only in nominal
Hariman. 1969;Bergquist, 1980a; Vacelet. 1985; genus Qasimellu), 5, isodictyal reticulation
Hooperctal., 1992). (reticulation with triangular meshes formed by
'POSITION OF THE SKELETON. All uni-orpaucispiculartracts ofspicules,cemented
micrucionids arc siliceous with discrete, free attheirnodesbycollagen0| fully enclosed within
Spicules* So far no desma-bcaring species orhy- spongin fibresi (Fig. 131A); 6, regularly or ir-
pcrcalcificd 'relict' species arc known. Many regularly reticulate (with large, multispicular
species undergo secondary acquisition, loss or tracts and/or fibres forming irregularoval orrec-
reduction Of spicule mineralisation, particularly tangular meshes(Fig. 7A-B); 7, plumo-reticulale
when displaced hy arenaceous particles (e.g., (producing ascending and consecutively diverg-
N&lopsomrno), C IWftswella) is partly defined ing tracts and fibres, forming pauci- or multi-
by this feature, with various degrees, of spicular primary lines, and interconnected by
arenaceous development among species; this Irunsversc uni- orpaucispiculartracts and fibres)
trend is widespread throughout the (Fig. 7D); 8, dendro-reticulate (similar to the
Poccilosclcrida. preceding,but where ascendingtractsaresinuous
andmoreobviouslydivergingandbranchingthan
OrganisationoftiieSk^i.i.iok.Grossorganic the less conspicuous transverse elements) (Fig.
aodinorganicskeletal architecture,Structural dif- 231C);9, plumose(withascendinganddiverging
ferentiation of the inorganic skeleton, and dis- primary lines ih;it are noi connectedbytransverse
tribution ofmineralcomponents in that structure elements) (Fig. 7E); 10, axially or basally com-
arc primary diagnostics (Levi, 1960a, 1973; pressed (having a skeleton clearly divided into a
Bergquist, 1978a; Hartman, 1982). However. compressed central orbasal coreoffibres and/or
REVISIONOFMICROCIONIMAt-
of types: 1. principal spicules
ECTOSOME (Fig. 2A). robust, straight or
slightly curved primary styles
subtylostyles, tylostylcs or
SirBECTOSOME quasi-monactina! fo
generally (but not invariably'
confined within fibres
CHOANOSOME auxiliary spicules (Fig. 2B),
more slender, curved, sinuous
orstraightslyles.subtylostyles,
tylostyles or quasi-diacrinal
9 roost commonly located
ouLside fibres in theectosoma!
or subectosomaJ skeletons, or
BASAL dispersed throughout the
SPONGIN FIBRE mesohyl; 3, accessory spicules
CRg. 4), styles, acanthostyles
or modified quasi-diactinal
forms generally eehinating
fibres.Thus,mostspecieshave
FIG. I.Idealisedmicrocionidskeletalstructure. 1,Echin3tingac«irHhostyle5. principalspiculescuringfibres
2, Reticulate fibre skeleton; 3, Isotropic extra-fibre skeleton. 4. Detntal i=chnanosomal principals',
cmcipping fibres. 5, Renieroid reticulate secondary fibre skeleton. 6, some (e.g., EmnochaUtui}
'Microcionid' radial fibre skeleton. 7, 'Spicate* spicule skeleton. 8,
Hymedesmoid spicule skeleton. 9, Plumose/dendritic Fibre skeleton. 10. have coring spicules derived
Coring principal spicules 11, Suhectnsoma) auxiliary spicules 12. fromauxiliaries(=chconosomal
Eclosomalauxiliaryspicules. auxiliaries), and some (e.g.,
Clathria (Dendrvcia)) IaGk
spicules, forming tight anastomosing meshes, principalspicules altogether.
and fromwhichariseplumoseorplumoreticulate
Ectosomal Skeleton. Within Microciomdac the
extra-axial(subectosomal)fibresand/orspicule-s) ectosomal skeleton has been attributed greater
Fig 7G), Some species have combinations of
I importance V..u Soest, 19S4b) than over ele-
these skeletal structures with different structure]
ments such as choanosomal architecture and
typesfound indifferentpartsoftheskeleton(e.g., growth form (e,g.> Levi. 1960a; Bcrquisl &
axis and periphery).
Fmmont, 19S8) microsciere diversity (e.g., dc
t
Structur/vl regions oi the Skeleton, The Laubenfels. I9?6a), or megasclcres eehinating
skeleton may be divided into three main struc- fibresandchofiflOSoma] spicules(e.g., Halimann,
tures or regions (Fig. I): 1, ectosoma! (outer 1912, 1920). Within the family ectosomal
perforated epithelium comprised ofsingle elon- development ranges frommembraneous,without
gate flattened pinaeoL vies, including an ec- an ectosomal skeleton (Fig. MF); membran*
tosomal skeleton, wherepresent,arisingfromthe with a specialised tangential, reticulate fibre
mesohyl directly below the exopinacodcrrn); 2, skeleton (Fig. 255C); with anectosomal tangen-
choanosomaloraxialcore(centralorbasalregion tial reticulation ofdetritus (Fig 92A); with sub-
containing choanocyte chambers, fibres and ectosomal spicules erect, paratangential or
spicule tracts and the mesohyl ground substance, tangential to surface (Fig. 65F). orforming div
in addition to the diverse and mobile cell types); crete bundles (Fig. 59E); with special ectosomal
3, subectosomal or cortical zone (strictly part of spicules erect orparatangential to surface, form-
thechoanosomesince itlieswithintheboundsof ing acontinuous palisade (Fig. 155G)ordiscTCte
the external epithelial barrier, but is immediately bundles(Fig, 151F)
subdermal.andthedelineationofthisregionfrom Subectosomal Skeleton. The region between
the choanosome properis based on the develop- choanosomal and ectosomal layers may or may
mentofanextra-fibreinsomespecies structurally notbedifferentiatedinloasubectosomal(subder-
differentiated from the deeper choanosomal or mal or txitt axial) structure. This region may
superficial ectosomal skeletons). occupy a small proportion of the peripheral
This system isusedinconjunction withspicule mesohyl (eg > Hohpsamma; Fig. 269D), or it
nomenclature (Kallmann. 1912), based on origin may comprise ihe majority of sponge diameter
MEMOIRS OFTHEQUEENSLANDMUSEUM
FIG.2.Majorstructuralmegascleres.A,Principalspicules(style,subtylostyle,acanthostylesubtylostyle,hastate
style). B, Auxiliary spicules (style, subtylostyle, rhabdose subtylostyle, spined subtylostyle, asymmetrical
styloid,tornostyle,quasistrongyle,mucronatestyloid).C,Basesofprincipalspicules(style,subtylostyle,hastate
style, acanthose subtylostyle, vestigial basal spination). D, Bases ofauxiliary spicules (pointed subtylostyle,
tylostyle, style,acanthosestyle,terminal spines,tornostyle).
REVISIONOFMICROCIONIDAE
FIG. 3. Terminations ofstructural megascleres. A, Fusiform (pointed and rounded). B, Telescoped. C, Hastate
(pointedandrounded).D, Spined(granular,tuberculate,terminalsharp,subterminalsharp,mucronate,bifurcate).
(e.g., C. (Axociella); Fig. 118F). When present, truly diactinal (Fig. 307A-B) as seen in E.
subectosomal extra-fibre development may be (Protophlitaspongia).
completely disorganised or it may be well or- Choanosoinal Skeleton. In Microcionidae skele-
ganised into radial non-plumose bundles of talstructuresarediverse(listedabove),andtradi-
spicules (Fig. 127B). The composition and tionally important in classification. In terms of
geometry of spicules within the subectosomal differentiated regions within the choanosomal
skeleton varies from species lacking any form of skeleton there are species with simple reticulate
subectosomal spicules (e.g., C. (Micrvciona); fibreswithoutanyaxial orextra-axialdifferentia-
Fig. 102F), those with a single form ofauxiliary tion(Fig. 7C),thosewith markedaxial andextra-
spicule throughout the skeleton (e.g., C axial differentiation of the choanosomal and
(Dendrvcia); Fig. 11 2D), those with two subectosomal skeletons, which are structurally
categories of structural megascleres, one in the distinct from the peripheral skeleton (Fig. 7G),
C
ectosome and one in spongin fibres (e.g., and those in which the choanosome is simply
(Clathria); Fig.73F),tothosewiththree formsof reduced to a basal layer ofspongin lying on the
structuralmegascleres,allofwhichoccurtosome substrate (Fig. 7H). Similarly, choanosomal
C
extent in the subdermal region (e.g., skeletal tracts may be diversified into more than
(Thalysias); Fig. 197G), and spicules may be one element, including those with no obvious
monactinal (for most microcionid species; e.g., divisionofprimaryorsecondaryskeletallines(Fig.
Fig. 28A-B), orquasi-diactinal (Fig. 298A-B), or 159F), those with more-or-less well differen-
MEMOIRS OFTHEQUEENSLAND MUSEUM
10
FIG.4.Echinatingandacanthoseprincipalmegascleresandspination.A,Microspinedechinatingacanthostyles
(typical recurvedspines, typical erect,vestigial,poorly silicified, hastate smooth,entirely smooth). B, Myxil-
lid-like echinating acanthostyles (aspinose apex, clavulate apex, entirely spined, bare neck). C, Modified
principal spicules (three forms of intermediate principal echinating spicules, acanthostrongyles). D, Spine
geometry(greatlyrecurvedspines, slightlyrecurvedspines,erectspines, granularspines,tuberculatespines).