Micronesica 35-36100-120 2003

Taxonomic inventory of the sponges (Porifera) of the Mariana Islands

MICHELLE KELLY

National Institute of Water amp Atmospheric Research (NIWA) LtdPrivate Bag 109-695

Newmarket Auckland New Zealand

JOHN HOOPER

Queensland MuseumPO Box 3300

South Brisbane Queensland 4101 Australia

VALERIE PAUL1 AND GUSTAV PAULAY2

Marine LaboratoryUniversity of Guam

Mangilao Guam 96923 USA

ROB VAN SOEST AND WALLIE DE WEERDT

Institute for Biodiversity and Ecosystem DynamicsZoologisch Museum

University of AmsterdamP O Box 94766 1090 GT Amsterdam The Netherlands

AbstractmdashWe review the sponge fauna of the Mariana Islands based onnew and existing collections and literature records 124 species ofsiliceous sponges (Class Demospongiae) and 4 species of calcareoussponges (Class Calcarea) have been identified to date representing 73genera 44 families within 16 orders Several species are adventiveApproximately 30 (40) of the species encountered are undescribed butnot all are endemics as the authors know them from other locationsApproximately 30 (38) of the species are known from diverse locationswithin the Indo West Pacific but several well-known widespreadspecies are absent The actual diversity of sponge fauna of the Marianasis considerably higher as many species especially cryptic and encrustingtaxa remain to be collected and studied

IntroductionOur knowledge of the sponge fauna of the tropical Pacific has increased

substantially in recent years as a result of enhanced collecting effort driven in

1 current address Smithsonian Marine Station at Fort Pierce Fort Pierce FL 349492 corresponding author current address Florida Museum of Natural History University of Florida

Gainesville FL 32611-7800 USA email paulayflmnhufledu

Kelly et al Sponges of the Marianas 101

part by pharmaceutical interests and by the attention of a larger number ofsystematists working on Pacific sponges than ever before Nevertheless with theexception of Micronesia Hawaii Papua New Guinea and New Caledonia thesponge faunas of most tropical island groups in Oceania are poorly known(Kelly-Borges amp Valentine 1995) Prior to the 1950rsquos records of Pacific spongeswere fragmentary and confined to the reports of major scientific expeditions suchas that of the H M S Alert (Ridley 1884) the H M S Challenger (Ridley ampDendy 1886 1887) and the Albatross (Agassiz 1906 Lendenfeld 1910a b1915 Wilson 1925) Our basic knowledge of Micronesian sponges is basedlargely on the studies of de Laubenfels (1954) and Bergquist (1965) with theformer including material from the Mariana Islands De Laubenfels (1954)recorded 9 species from Guam and 10 from Saipan with one species in commonbased on his own collecting on both islands as well as on material gathered byPreston Cloud on Saipan Few taxonomic papers have appeared on sponges fromthe Marianas since Hartman amp Goreau (1975 1976) described two corallinesponges Acanthochaetetes wellsi and Stromatospongia micronesica from caveson Guam and Saipan Smyth (1990) recorded 7 species of boring sponges fromgastropod shells on Guam Terpios hoshinota received much attention on Guamdue to the threat it poses to reefs because of its rapid enveloping growth (Bryan1973 Plucer-Rosario 1987) it was described by Ruumltzler amp Muzik in 1993 Quinnamp Kojis (1999) described the community structure of coralline sponges from theldquoGrottordquo a well-known marine cavern on Saipan

The Coral Reef Research Foundation (CRRF) has contributed significantlyto our knowledge of the sponge fauna of Saipan Tinian and Rota Islands in theCommonwealth of the Northern Mariana Islands (CNMI) CRRF is a non-profitorganization based in Koror Palau and engaged in a long-term sponge collectionand inventory program in Micronesia and elsewhere in the Indo-West PacificKelly-Borges amp Valentine (1995) reviewed knowledge of sponges of the tropicalisland region of Oceania including published and unpublished species identifiedthrough CRRF research up to that time Unfortunately numerous records wereinadvertently listed from the Marianas in that paper that did not originate fromthere All CRRF records from the Marianas were therefore reevaluated for thepresent publication and records that we do not list from the Marianas but appearin Kelly-Borges amp Valentine (1995) should be disregarded

Our knowledge of the marine sponges of Guam has increased markedlysince the mid-1980rsquos first as a result of a research program on marine naturalproducts and chemical ecology established by Valerie Paul at the University ofGuam She and her collaborators have published extensively on the ecology andchemistry of local sponges and solicited numerous identifications fromspecialists As a result marine natural product reports form the bulk ofpublications on sponges from Guam and the CNMI in recent years (Appendix 1)Much of this research relates to the general use of secondary metabolites insponges as chemical deterrents to fish and invertebrate predation (Rogers amp Paul1991 Pennings et al 1994 Becerro et al 1998) and as allelopathic compounds

102 Micronesica 35-36 2003

in spatial competition in coral reef habitats (Thacker et al 1998) In particularthe sponges Hyrtios erecta (Rogers amp Paul 1991) Luffariella variabilis andDysidea sp 4 (Duffy amp Paul 1992 Pennings et al 1994) Dysidea sp 2 (Paul etal 1997 Thacker et al 1998) and Cacospongia sp 1 (Becerro et al 1998) havereceived considerable attention Some specialist consumers of sponges have alsoreceived attention including cephalaspideans (Carlson amp Hoff 1973 1974) andnudibranchs (Rogers amp Paul 1991 Avila amp Paul 1997 Becerro et al 1998) Ashort course conducted on Guam in sponge taxonomy in 1996 by Michelle Kellyfostered further interest in sponges During the 1990rsquos Gustav Paulayrsquos lab wasinvolved in structured as well as unstructured surveys of the marine biodiversityof Guam New material was collected during three biodiversity surveys carriedout for COMNAVMARIANAS (US Department of Defense) at Apra Harbor(Paulay et al 1997) at the southern Orote ndash North Agat Bay area (Paulay et al2001) and at the Pugua Patchreef ndash Haputo area (Amesbury et al 2001) The firstsurvey was partly focused on sponges while the latter two were generalinvertebrate surveys although exposed macro-sponges were routinelydocumented A non-indigenous species survey funded by Sea Grant providedadditional coverage (Paulay et al 2002)

The aim of this paper is to provide an inventory of the marine spongesknown from the coastal reefs of Guam and the CNMI The taxa listed representwell known and partially identified sponge species many of which remainundescribed The specimens and photo vouchers associated with most of thesemorphospecies should assist in future identifications by allowing theirrecognition through inevitable future nomenclatural and systematic changes Thepresent list constitutes a working list from which taxonomic descriptions of newand known taxa may arise This taxonomic inventory also aims to achievemaximal biological accuracy through examination of actual material by experttaxonomists All too frequently checklists of this nature are uncriticalcompilations of species names directly from the literature resulting in somespecies being covered under multiple names because of misidentifications orsynonymies and others included erroneously from poor identifications or localityrecords Most of the taxonomic identifications have been checked and updatedThe exceptions are a few literature records (see Appendix 1) for which specimenswere not reexamined

MethodsThis paper is based on collections from Guam made by staff of the

University of Guam Marine Laboratory collections from CNMI made by CRRFand records from the literature Several of the species recorded in the literaturewere also recollected but some of de Laubenfelsrsquo (1954) records and the smallboring sponges recorded by Smyth (1990) were not (see Appendix 1) Spongeswere collected using SCUBA and snorkeling All newly collected species weredistinguished in the field characterized by field characters and photographed in

Kelly et al Sponges of the Marianas 103

situ Most species encountered were documented by both vouchers and photoswhile a few were documented by only one of these methods (Appendix 1) Aftercollection material was fixed in 80 ethanol and prepared for identification bythin sectioning and through preparation of spicule slides (see Kelly-Borges ampVacelet 1995) Vouchers (see Appendix 1) are deposited at the Natural HistoryMuseum London (NHM) University of Guam Mangilao (UGI) QueenslandMuseum Brisbane (QM) National Institute of Water amp Atmospheric ResearchInstitute New Zealand (NIWA) Smithsonian Institution (USNM) ZoologischMuseum University of Amsterdam (ZMA) and the collections of the Coral ReefResearch Foundation Republic of Belau (CRRF) Photo vouchers are by Paulay(GP numbers housed at the Florida Museum of Natural History) and CRRF(0CDN numbers housed at CRRF) Cited photographs (Appendix 1) are on theWWW at httpwwwflmnhufledureefs they are also available on the MarineBiodiversity of Guam CD-ROM co-publication (see website for availability)

Results and DiscussionWe have documented 128 sponge species from intertidal to ca 50m depths

on coastal reefs and caves of Guam and the CNMI (Appendix 1) These comprise124 siliceous (Class Demospongiae) and 4 calcareous (Class Calcarea) spongesrepresenting 73 genera 44 families within 16 orders During the process of thisreview several systematic changes were made (see comments to Appendix 1)

The 128 species recorded here represent only part of the sponge diversity ofthe Marianas as both collecting and taxonomic efforts remain limited Over 30additional species are known to the authors from Guam but have not receivedsufficient taxonomic attention to be included here Although there have beennumerous collections made in the Marianas over the years none have beencomprehensive Rather sponges have been collected opportunistically fromdiverse locations on Guam and mostly from cave habitats on Saipan Rota andTinian No sponge records are available from the ecologically distinctivevolcanically active islands of the northern arc of the Marianas

The rich sponge fauna of Apra Harbor on Guam has received the mostattention to date and the exposed macrosponges of that area are relatively wellknown The exposed macrosponges of outer reef slope habitats on Guam are nextbest known but new taxa are regularly encountered on every new reef areasurveyed The rich sponge fauna of caves on the outer reef slope have been onlyincidentally sampled on Guam and only the larger species preferred forpharmaceutical analysis were sampled in this habitat in the CNMI Although thesponge fauna of shallow fringing reefs received the first taxonomic attention (deLaubenfels 1954) this habitat remains poorly known as little recent attention hasbeen devoted to it in the Marianas in recent years Sponge faunas are renownedfor their spatial heterogeneity (Hooper amp Leacutevi 1994 Hooper amp Kennedy 2002)There is considerable variation in species composition even among neighboringreefs on an island with additional differences among habitats and among islands

104 Micronesica 35-36 2003

(eg Hooper 1994) This phenomenon is also clearly evident on Guam wheremany sponge species are known from restricted biotopes or reef areas Weestimate the diversity of exposed macrosponges in the Marianas to be at leasttwice that recorded here The unstudied encrusting and cryptic sponges are likelyat least as diverse as macrosponges making the total poriferan diversity of theMarianas at least 500 species

In contrast to the documented local (alpha scale) heterogeneity in spongedistributions sponge faunas become more predictable at regional (gamma) scales(Hooper et al 2002) Consequently the Mariana Islandsrsquo sponge fauna may beexpected to contain predictable elements of a broader Micronesian or westcentral Pacific fauna However numerous conspicuous and abundant spongespecies that are nearly ubiquitous on neighboring archipelagoes as well as inmuch of the west central Pacific appear to be absent in the Marianas Theseinclude the haplosclerids Haliclona (Reniera) poseidon (de Laubenfels)Gelliodes fibulatus (Carter) Oceanapia sagittaria (Sollas) Callyspongiaaerizusa Desqueyroux-Faundez and Cribrochalina olemda de Laubenfels thedictyoceratids Spongia matamata de Laubenfels Carteriospongia foliascens(Pallas) and Phyllospongia papyracea (Esper) the poecilosclerids Biemnatubulata (Dendy) the axinellid Axinella carteri (Dendy) species of DiacarnusTheonella and the calcareous sponges Pericharax heteroraphis Poleacutejaeff andLeucetta avocada de Laubenfels Many of these species have been recordedconsistently within various Indo-west Pacific reef systems from Vanuatu toThailand the Great Barrier Reef to Chuuk and represent some of the 5 of theregional fauna that is hypothesized to be widely distributed (eg Hooper et al2000) There is a growing body data however to suggest that at least somewidespread morphological species are comprised of genetically-divergentallopatric species complexes (eg Astrosclera willeyana ndash Woerheide et al2003)

Nevertheless part of the regional fauna is represented in the Marianas thusapproximately 30 (38) of the species recorded are regionally widespread (seeHooper et al 2000) The absence of numerous regional species is partly areflection of the lower diversity of sponges in the Marianas than on neighboringisland groups Thus CRRF has recorded 218 species of sponges from Chuuk inthe central Carolines and 295 species from Palau (L Colin pers comm 2001) Asurvey of 6 of the 80 known marine lakes in Palau resulted in more than 140sponge species (Kelly-Borges amp Valentine 1995) Kelly-Borges amp Valentine(1995) indicated that there are over 1000 published and undescribed species inthe tropical region of Oceania including Guam but this is a vast underestimateof the known fauna (JNH MK CRRF unpublished information) In comparisonHooper amp Leacutevi (1994) estimated that there are about 5000 published andundescribed species in the Australian insular and continental faunas alone

The relatively low diversity of sponges in the Marianas relative to Palau andthe Federated States of Micronesia fits general patterns seen in other marineorganisms (eg Randall 1995 Myers 1999) and reflects the isolation location

Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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Kelly et al Sponges of the Marianas 101

part by pharmaceutical interests and by the attention of a larger number ofsystematists working on Pacific sponges than ever before Nevertheless with theexception of Micronesia Hawaii Papua New Guinea and New Caledonia thesponge faunas of most tropical island groups in Oceania are poorly known(Kelly-Borges amp Valentine 1995) Prior to the 1950rsquos records of Pacific spongeswere fragmentary and confined to the reports of major scientific expeditions suchas that of the H M S Alert (Ridley 1884) the H M S Challenger (Ridley ampDendy 1886 1887) and the Albatross (Agassiz 1906 Lendenfeld 1910a b1915 Wilson 1925) Our basic knowledge of Micronesian sponges is basedlargely on the studies of de Laubenfels (1954) and Bergquist (1965) with theformer including material from the Mariana Islands De Laubenfels (1954)recorded 9 species from Guam and 10 from Saipan with one species in commonbased on his own collecting on both islands as well as on material gathered byPreston Cloud on Saipan Few taxonomic papers have appeared on sponges fromthe Marianas since Hartman amp Goreau (1975 1976) described two corallinesponges Acanthochaetetes wellsi and Stromatospongia micronesica from caveson Guam and Saipan Smyth (1990) recorded 7 species of boring sponges fromgastropod shells on Guam Terpios hoshinota received much attention on Guamdue to the threat it poses to reefs because of its rapid enveloping growth (Bryan1973 Plucer-Rosario 1987) it was described by Ruumltzler amp Muzik in 1993 Quinnamp Kojis (1999) described the community structure of coralline sponges from theldquoGrottordquo a well-known marine cavern on Saipan

The Coral Reef Research Foundation (CRRF) has contributed significantlyto our knowledge of the sponge fauna of Saipan Tinian and Rota Islands in theCommonwealth of the Northern Mariana Islands (CNMI) CRRF is a non-profitorganization based in Koror Palau and engaged in a long-term sponge collectionand inventory program in Micronesia and elsewhere in the Indo-West PacificKelly-Borges amp Valentine (1995) reviewed knowledge of sponges of the tropicalisland region of Oceania including published and unpublished species identifiedthrough CRRF research up to that time Unfortunately numerous records wereinadvertently listed from the Marianas in that paper that did not originate fromthere All CRRF records from the Marianas were therefore reevaluated for thepresent publication and records that we do not list from the Marianas but appearin Kelly-Borges amp Valentine (1995) should be disregarded

Our knowledge of the marine sponges of Guam has increased markedlysince the mid-1980rsquos first as a result of a research program on marine naturalproducts and chemical ecology established by Valerie Paul at the University ofGuam She and her collaborators have published extensively on the ecology andchemistry of local sponges and solicited numerous identifications fromspecialists As a result marine natural product reports form the bulk ofpublications on sponges from Guam and the CNMI in recent years (Appendix 1)Much of this research relates to the general use of secondary metabolites insponges as chemical deterrents to fish and invertebrate predation (Rogers amp Paul1991 Pennings et al 1994 Becerro et al 1998) and as allelopathic compounds

102 Micronesica 35-36 2003

in spatial competition in coral reef habitats (Thacker et al 1998) In particularthe sponges Hyrtios erecta (Rogers amp Paul 1991) Luffariella variabilis andDysidea sp 4 (Duffy amp Paul 1992 Pennings et al 1994) Dysidea sp 2 (Paul etal 1997 Thacker et al 1998) and Cacospongia sp 1 (Becerro et al 1998) havereceived considerable attention Some specialist consumers of sponges have alsoreceived attention including cephalaspideans (Carlson amp Hoff 1973 1974) andnudibranchs (Rogers amp Paul 1991 Avila amp Paul 1997 Becerro et al 1998) Ashort course conducted on Guam in sponge taxonomy in 1996 by Michelle Kellyfostered further interest in sponges During the 1990rsquos Gustav Paulayrsquos lab wasinvolved in structured as well as unstructured surveys of the marine biodiversityof Guam New material was collected during three biodiversity surveys carriedout for COMNAVMARIANAS (US Department of Defense) at Apra Harbor(Paulay et al 1997) at the southern Orote ndash North Agat Bay area (Paulay et al2001) and at the Pugua Patchreef ndash Haputo area (Amesbury et al 2001) The firstsurvey was partly focused on sponges while the latter two were generalinvertebrate surveys although exposed macro-sponges were routinelydocumented A non-indigenous species survey funded by Sea Grant providedadditional coverage (Paulay et al 2002)

The aim of this paper is to provide an inventory of the marine spongesknown from the coastal reefs of Guam and the CNMI The taxa listed representwell known and partially identified sponge species many of which remainundescribed The specimens and photo vouchers associated with most of thesemorphospecies should assist in future identifications by allowing theirrecognition through inevitable future nomenclatural and systematic changes Thepresent list constitutes a working list from which taxonomic descriptions of newand known taxa may arise This taxonomic inventory also aims to achievemaximal biological accuracy through examination of actual material by experttaxonomists All too frequently checklists of this nature are uncriticalcompilations of species names directly from the literature resulting in somespecies being covered under multiple names because of misidentifications orsynonymies and others included erroneously from poor identifications or localityrecords Most of the taxonomic identifications have been checked and updatedThe exceptions are a few literature records (see Appendix 1) for which specimenswere not reexamined

MethodsThis paper is based on collections from Guam made by staff of the

University of Guam Marine Laboratory collections from CNMI made by CRRFand records from the literature Several of the species recorded in the literaturewere also recollected but some of de Laubenfelsrsquo (1954) records and the smallboring sponges recorded by Smyth (1990) were not (see Appendix 1) Spongeswere collected using SCUBA and snorkeling All newly collected species weredistinguished in the field characterized by field characters and photographed in

Kelly et al Sponges of the Marianas 103

situ Most species encountered were documented by both vouchers and photoswhile a few were documented by only one of these methods (Appendix 1) Aftercollection material was fixed in 80 ethanol and prepared for identification bythin sectioning and through preparation of spicule slides (see Kelly-Borges ampVacelet 1995) Vouchers (see Appendix 1) are deposited at the Natural HistoryMuseum London (NHM) University of Guam Mangilao (UGI) QueenslandMuseum Brisbane (QM) National Institute of Water amp Atmospheric ResearchInstitute New Zealand (NIWA) Smithsonian Institution (USNM) ZoologischMuseum University of Amsterdam (ZMA) and the collections of the Coral ReefResearch Foundation Republic of Belau (CRRF) Photo vouchers are by Paulay(GP numbers housed at the Florida Museum of Natural History) and CRRF(0CDN numbers housed at CRRF) Cited photographs (Appendix 1) are on theWWW at httpwwwflmnhufledureefs they are also available on the MarineBiodiversity of Guam CD-ROM co-publication (see website for availability)

Results and DiscussionWe have documented 128 sponge species from intertidal to ca 50m depths

on coastal reefs and caves of Guam and the CNMI (Appendix 1) These comprise124 siliceous (Class Demospongiae) and 4 calcareous (Class Calcarea) spongesrepresenting 73 genera 44 families within 16 orders During the process of thisreview several systematic changes were made (see comments to Appendix 1)

The 128 species recorded here represent only part of the sponge diversity ofthe Marianas as both collecting and taxonomic efforts remain limited Over 30additional species are known to the authors from Guam but have not receivedsufficient taxonomic attention to be included here Although there have beennumerous collections made in the Marianas over the years none have beencomprehensive Rather sponges have been collected opportunistically fromdiverse locations on Guam and mostly from cave habitats on Saipan Rota andTinian No sponge records are available from the ecologically distinctivevolcanically active islands of the northern arc of the Marianas

The rich sponge fauna of Apra Harbor on Guam has received the mostattention to date and the exposed macrosponges of that area are relatively wellknown The exposed macrosponges of outer reef slope habitats on Guam are nextbest known but new taxa are regularly encountered on every new reef areasurveyed The rich sponge fauna of caves on the outer reef slope have been onlyincidentally sampled on Guam and only the larger species preferred forpharmaceutical analysis were sampled in this habitat in the CNMI Although thesponge fauna of shallow fringing reefs received the first taxonomic attention (deLaubenfels 1954) this habitat remains poorly known as little recent attention hasbeen devoted to it in the Marianas in recent years Sponge faunas are renownedfor their spatial heterogeneity (Hooper amp Leacutevi 1994 Hooper amp Kennedy 2002)There is considerable variation in species composition even among neighboringreefs on an island with additional differences among habitats and among islands

104 Micronesica 35-36 2003

(eg Hooper 1994) This phenomenon is also clearly evident on Guam wheremany sponge species are known from restricted biotopes or reef areas Weestimate the diversity of exposed macrosponges in the Marianas to be at leasttwice that recorded here The unstudied encrusting and cryptic sponges are likelyat least as diverse as macrosponges making the total poriferan diversity of theMarianas at least 500 species

In contrast to the documented local (alpha scale) heterogeneity in spongedistributions sponge faunas become more predictable at regional (gamma) scales(Hooper et al 2002) Consequently the Mariana Islandsrsquo sponge fauna may beexpected to contain predictable elements of a broader Micronesian or westcentral Pacific fauna However numerous conspicuous and abundant spongespecies that are nearly ubiquitous on neighboring archipelagoes as well as inmuch of the west central Pacific appear to be absent in the Marianas Theseinclude the haplosclerids Haliclona (Reniera) poseidon (de Laubenfels)Gelliodes fibulatus (Carter) Oceanapia sagittaria (Sollas) Callyspongiaaerizusa Desqueyroux-Faundez and Cribrochalina olemda de Laubenfels thedictyoceratids Spongia matamata de Laubenfels Carteriospongia foliascens(Pallas) and Phyllospongia papyracea (Esper) the poecilosclerids Biemnatubulata (Dendy) the axinellid Axinella carteri (Dendy) species of DiacarnusTheonella and the calcareous sponges Pericharax heteroraphis Poleacutejaeff andLeucetta avocada de Laubenfels Many of these species have been recordedconsistently within various Indo-west Pacific reef systems from Vanuatu toThailand the Great Barrier Reef to Chuuk and represent some of the 5 of theregional fauna that is hypothesized to be widely distributed (eg Hooper et al2000) There is a growing body data however to suggest that at least somewidespread morphological species are comprised of genetically-divergentallopatric species complexes (eg Astrosclera willeyana ndash Woerheide et al2003)

Nevertheless part of the regional fauna is represented in the Marianas thusapproximately 30 (38) of the species recorded are regionally widespread (seeHooper et al 2000) The absence of numerous regional species is partly areflection of the lower diversity of sponges in the Marianas than on neighboringisland groups Thus CRRF has recorded 218 species of sponges from Chuuk inthe central Carolines and 295 species from Palau (L Colin pers comm 2001) Asurvey of 6 of the 80 known marine lakes in Palau resulted in more than 140sponge species (Kelly-Borges amp Valentine 1995) Kelly-Borges amp Valentine(1995) indicated that there are over 1000 published and undescribed species inthe tropical region of Oceania including Guam but this is a vast underestimateof the known fauna (JNH MK CRRF unpublished information) In comparisonHooper amp Leacutevi (1994) estimated that there are about 5000 published andundescribed species in the Australian insular and continental faunas alone

The relatively low diversity of sponges in the Marianas relative to Palau andthe Federated States of Micronesia fits general patterns seen in other marineorganisms (eg Randall 1995 Myers 1999) and reflects the isolation location

Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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102 Micronesica 35-36 2003

in spatial competition in coral reef habitats (Thacker et al 1998) In particularthe sponges Hyrtios erecta (Rogers amp Paul 1991) Luffariella variabilis andDysidea sp 4 (Duffy amp Paul 1992 Pennings et al 1994) Dysidea sp 2 (Paul etal 1997 Thacker et al 1998) and Cacospongia sp 1 (Becerro et al 1998) havereceived considerable attention Some specialist consumers of sponges have alsoreceived attention including cephalaspideans (Carlson amp Hoff 1973 1974) andnudibranchs (Rogers amp Paul 1991 Avila amp Paul 1997 Becerro et al 1998) Ashort course conducted on Guam in sponge taxonomy in 1996 by Michelle Kellyfostered further interest in sponges During the 1990rsquos Gustav Paulayrsquos lab wasinvolved in structured as well as unstructured surveys of the marine biodiversityof Guam New material was collected during three biodiversity surveys carriedout for COMNAVMARIANAS (US Department of Defense) at Apra Harbor(Paulay et al 1997) at the southern Orote ndash North Agat Bay area (Paulay et al2001) and at the Pugua Patchreef ndash Haputo area (Amesbury et al 2001) The firstsurvey was partly focused on sponges while the latter two were generalinvertebrate surveys although exposed macro-sponges were routinelydocumented A non-indigenous species survey funded by Sea Grant providedadditional coverage (Paulay et al 2002)

The aim of this paper is to provide an inventory of the marine spongesknown from the coastal reefs of Guam and the CNMI The taxa listed representwell known and partially identified sponge species many of which remainundescribed The specimens and photo vouchers associated with most of thesemorphospecies should assist in future identifications by allowing theirrecognition through inevitable future nomenclatural and systematic changes Thepresent list constitutes a working list from which taxonomic descriptions of newand known taxa may arise This taxonomic inventory also aims to achievemaximal biological accuracy through examination of actual material by experttaxonomists All too frequently checklists of this nature are uncriticalcompilations of species names directly from the literature resulting in somespecies being covered under multiple names because of misidentifications orsynonymies and others included erroneously from poor identifications or localityrecords Most of the taxonomic identifications have been checked and updatedThe exceptions are a few literature records (see Appendix 1) for which specimenswere not reexamined

MethodsThis paper is based on collections from Guam made by staff of the

University of Guam Marine Laboratory collections from CNMI made by CRRFand records from the literature Several of the species recorded in the literaturewere also recollected but some of de Laubenfelsrsquo (1954) records and the smallboring sponges recorded by Smyth (1990) were not (see Appendix 1) Spongeswere collected using SCUBA and snorkeling All newly collected species weredistinguished in the field characterized by field characters and photographed in

Kelly et al Sponges of the Marianas 103

situ Most species encountered were documented by both vouchers and photoswhile a few were documented by only one of these methods (Appendix 1) Aftercollection material was fixed in 80 ethanol and prepared for identification bythin sectioning and through preparation of spicule slides (see Kelly-Borges ampVacelet 1995) Vouchers (see Appendix 1) are deposited at the Natural HistoryMuseum London (NHM) University of Guam Mangilao (UGI) QueenslandMuseum Brisbane (QM) National Institute of Water amp Atmospheric ResearchInstitute New Zealand (NIWA) Smithsonian Institution (USNM) ZoologischMuseum University of Amsterdam (ZMA) and the collections of the Coral ReefResearch Foundation Republic of Belau (CRRF) Photo vouchers are by Paulay(GP numbers housed at the Florida Museum of Natural History) and CRRF(0CDN numbers housed at CRRF) Cited photographs (Appendix 1) are on theWWW at httpwwwflmnhufledureefs they are also available on the MarineBiodiversity of Guam CD-ROM co-publication (see website for availability)

Results and DiscussionWe have documented 128 sponge species from intertidal to ca 50m depths

on coastal reefs and caves of Guam and the CNMI (Appendix 1) These comprise124 siliceous (Class Demospongiae) and 4 calcareous (Class Calcarea) spongesrepresenting 73 genera 44 families within 16 orders During the process of thisreview several systematic changes were made (see comments to Appendix 1)

The 128 species recorded here represent only part of the sponge diversity ofthe Marianas as both collecting and taxonomic efforts remain limited Over 30additional species are known to the authors from Guam but have not receivedsufficient taxonomic attention to be included here Although there have beennumerous collections made in the Marianas over the years none have beencomprehensive Rather sponges have been collected opportunistically fromdiverse locations on Guam and mostly from cave habitats on Saipan Rota andTinian No sponge records are available from the ecologically distinctivevolcanically active islands of the northern arc of the Marianas

The rich sponge fauna of Apra Harbor on Guam has received the mostattention to date and the exposed macrosponges of that area are relatively wellknown The exposed macrosponges of outer reef slope habitats on Guam are nextbest known but new taxa are regularly encountered on every new reef areasurveyed The rich sponge fauna of caves on the outer reef slope have been onlyincidentally sampled on Guam and only the larger species preferred forpharmaceutical analysis were sampled in this habitat in the CNMI Although thesponge fauna of shallow fringing reefs received the first taxonomic attention (deLaubenfels 1954) this habitat remains poorly known as little recent attention hasbeen devoted to it in the Marianas in recent years Sponge faunas are renownedfor their spatial heterogeneity (Hooper amp Leacutevi 1994 Hooper amp Kennedy 2002)There is considerable variation in species composition even among neighboringreefs on an island with additional differences among habitats and among islands

104 Micronesica 35-36 2003

(eg Hooper 1994) This phenomenon is also clearly evident on Guam wheremany sponge species are known from restricted biotopes or reef areas Weestimate the diversity of exposed macrosponges in the Marianas to be at leasttwice that recorded here The unstudied encrusting and cryptic sponges are likelyat least as diverse as macrosponges making the total poriferan diversity of theMarianas at least 500 species

In contrast to the documented local (alpha scale) heterogeneity in spongedistributions sponge faunas become more predictable at regional (gamma) scales(Hooper et al 2002) Consequently the Mariana Islandsrsquo sponge fauna may beexpected to contain predictable elements of a broader Micronesian or westcentral Pacific fauna However numerous conspicuous and abundant spongespecies that are nearly ubiquitous on neighboring archipelagoes as well as inmuch of the west central Pacific appear to be absent in the Marianas Theseinclude the haplosclerids Haliclona (Reniera) poseidon (de Laubenfels)Gelliodes fibulatus (Carter) Oceanapia sagittaria (Sollas) Callyspongiaaerizusa Desqueyroux-Faundez and Cribrochalina olemda de Laubenfels thedictyoceratids Spongia matamata de Laubenfels Carteriospongia foliascens(Pallas) and Phyllospongia papyracea (Esper) the poecilosclerids Biemnatubulata (Dendy) the axinellid Axinella carteri (Dendy) species of DiacarnusTheonella and the calcareous sponges Pericharax heteroraphis Poleacutejaeff andLeucetta avocada de Laubenfels Many of these species have been recordedconsistently within various Indo-west Pacific reef systems from Vanuatu toThailand the Great Barrier Reef to Chuuk and represent some of the 5 of theregional fauna that is hypothesized to be widely distributed (eg Hooper et al2000) There is a growing body data however to suggest that at least somewidespread morphological species are comprised of genetically-divergentallopatric species complexes (eg Astrosclera willeyana ndash Woerheide et al2003)

Nevertheless part of the regional fauna is represented in the Marianas thusapproximately 30 (38) of the species recorded are regionally widespread (seeHooper et al 2000) The absence of numerous regional species is partly areflection of the lower diversity of sponges in the Marianas than on neighboringisland groups Thus CRRF has recorded 218 species of sponges from Chuuk inthe central Carolines and 295 species from Palau (L Colin pers comm 2001) Asurvey of 6 of the 80 known marine lakes in Palau resulted in more than 140sponge species (Kelly-Borges amp Valentine 1995) Kelly-Borges amp Valentine(1995) indicated that there are over 1000 published and undescribed species inthe tropical region of Oceania including Guam but this is a vast underestimateof the known fauna (JNH MK CRRF unpublished information) In comparisonHooper amp Leacutevi (1994) estimated that there are about 5000 published andundescribed species in the Australian insular and continental faunas alone

The relatively low diversity of sponges in the Marianas relative to Palau andthe Federated States of Micronesia fits general patterns seen in other marineorganisms (eg Randall 1995 Myers 1999) and reflects the isolation location

Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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Kelly et al Sponges of the Marianas 103

situ Most species encountered were documented by both vouchers and photoswhile a few were documented by only one of these methods (Appendix 1) Aftercollection material was fixed in 80 ethanol and prepared for identification bythin sectioning and through preparation of spicule slides (see Kelly-Borges ampVacelet 1995) Vouchers (see Appendix 1) are deposited at the Natural HistoryMuseum London (NHM) University of Guam Mangilao (UGI) QueenslandMuseum Brisbane (QM) National Institute of Water amp Atmospheric ResearchInstitute New Zealand (NIWA) Smithsonian Institution (USNM) ZoologischMuseum University of Amsterdam (ZMA) and the collections of the Coral ReefResearch Foundation Republic of Belau (CRRF) Photo vouchers are by Paulay(GP numbers housed at the Florida Museum of Natural History) and CRRF(0CDN numbers housed at CRRF) Cited photographs (Appendix 1) are on theWWW at httpwwwflmnhufledureefs they are also available on the MarineBiodiversity of Guam CD-ROM co-publication (see website for availability)

Results and DiscussionWe have documented 128 sponge species from intertidal to ca 50m depths

on coastal reefs and caves of Guam and the CNMI (Appendix 1) These comprise124 siliceous (Class Demospongiae) and 4 calcareous (Class Calcarea) spongesrepresenting 73 genera 44 families within 16 orders During the process of thisreview several systematic changes were made (see comments to Appendix 1)

The 128 species recorded here represent only part of the sponge diversity ofthe Marianas as both collecting and taxonomic efforts remain limited Over 30additional species are known to the authors from Guam but have not receivedsufficient taxonomic attention to be included here Although there have beennumerous collections made in the Marianas over the years none have beencomprehensive Rather sponges have been collected opportunistically fromdiverse locations on Guam and mostly from cave habitats on Saipan Rota andTinian No sponge records are available from the ecologically distinctivevolcanically active islands of the northern arc of the Marianas

The rich sponge fauna of Apra Harbor on Guam has received the mostattention to date and the exposed macrosponges of that area are relatively wellknown The exposed macrosponges of outer reef slope habitats on Guam are nextbest known but new taxa are regularly encountered on every new reef areasurveyed The rich sponge fauna of caves on the outer reef slope have been onlyincidentally sampled on Guam and only the larger species preferred forpharmaceutical analysis were sampled in this habitat in the CNMI Although thesponge fauna of shallow fringing reefs received the first taxonomic attention (deLaubenfels 1954) this habitat remains poorly known as little recent attention hasbeen devoted to it in the Marianas in recent years Sponge faunas are renownedfor their spatial heterogeneity (Hooper amp Leacutevi 1994 Hooper amp Kennedy 2002)There is considerable variation in species composition even among neighboringreefs on an island with additional differences among habitats and among islands

104 Micronesica 35-36 2003

(eg Hooper 1994) This phenomenon is also clearly evident on Guam wheremany sponge species are known from restricted biotopes or reef areas Weestimate the diversity of exposed macrosponges in the Marianas to be at leasttwice that recorded here The unstudied encrusting and cryptic sponges are likelyat least as diverse as macrosponges making the total poriferan diversity of theMarianas at least 500 species

In contrast to the documented local (alpha scale) heterogeneity in spongedistributions sponge faunas become more predictable at regional (gamma) scales(Hooper et al 2002) Consequently the Mariana Islandsrsquo sponge fauna may beexpected to contain predictable elements of a broader Micronesian or westcentral Pacific fauna However numerous conspicuous and abundant spongespecies that are nearly ubiquitous on neighboring archipelagoes as well as inmuch of the west central Pacific appear to be absent in the Marianas Theseinclude the haplosclerids Haliclona (Reniera) poseidon (de Laubenfels)Gelliodes fibulatus (Carter) Oceanapia sagittaria (Sollas) Callyspongiaaerizusa Desqueyroux-Faundez and Cribrochalina olemda de Laubenfels thedictyoceratids Spongia matamata de Laubenfels Carteriospongia foliascens(Pallas) and Phyllospongia papyracea (Esper) the poecilosclerids Biemnatubulata (Dendy) the axinellid Axinella carteri (Dendy) species of DiacarnusTheonella and the calcareous sponges Pericharax heteroraphis Poleacutejaeff andLeucetta avocada de Laubenfels Many of these species have been recordedconsistently within various Indo-west Pacific reef systems from Vanuatu toThailand the Great Barrier Reef to Chuuk and represent some of the 5 of theregional fauna that is hypothesized to be widely distributed (eg Hooper et al2000) There is a growing body data however to suggest that at least somewidespread morphological species are comprised of genetically-divergentallopatric species complexes (eg Astrosclera willeyana ndash Woerheide et al2003)

Nevertheless part of the regional fauna is represented in the Marianas thusapproximately 30 (38) of the species recorded are regionally widespread (seeHooper et al 2000) The absence of numerous regional species is partly areflection of the lower diversity of sponges in the Marianas than on neighboringisland groups Thus CRRF has recorded 218 species of sponges from Chuuk inthe central Carolines and 295 species from Palau (L Colin pers comm 2001) Asurvey of 6 of the 80 known marine lakes in Palau resulted in more than 140sponge species (Kelly-Borges amp Valentine 1995) Kelly-Borges amp Valentine(1995) indicated that there are over 1000 published and undescribed species inthe tropical region of Oceania including Guam but this is a vast underestimateof the known fauna (JNH MK CRRF unpublished information) In comparisonHooper amp Leacutevi (1994) estimated that there are about 5000 published andundescribed species in the Australian insular and continental faunas alone

The relatively low diversity of sponges in the Marianas relative to Palau andthe Federated States of Micronesia fits general patterns seen in other marineorganisms (eg Randall 1995 Myers 1999) and reflects the isolation location

Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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AG

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rone

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976

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e Y

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S)5

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RID

A A

NC

OR

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AE

Mel

ophl

us sa

rasi

noru

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hiel

e 1

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BM

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199

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225

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ux (d

e La

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195

4)59

40B

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995

622

4G

P274

-18

5G

104 Micronesica 35-36 2003

(eg Hooper 1994) This phenomenon is also clearly evident on Guam wheremany sponge species are known from restricted biotopes or reef areas Weestimate the diversity of exposed macrosponges in the Marianas to be at leasttwice that recorded here The unstudied encrusting and cryptic sponges are likelyat least as diverse as macrosponges making the total poriferan diversity of theMarianas at least 500 species

In contrast to the documented local (alpha scale) heterogeneity in spongedistributions sponge faunas become more predictable at regional (gamma) scales(Hooper et al 2002) Consequently the Mariana Islandsrsquo sponge fauna may beexpected to contain predictable elements of a broader Micronesian or westcentral Pacific fauna However numerous conspicuous and abundant spongespecies that are nearly ubiquitous on neighboring archipelagoes as well as inmuch of the west central Pacific appear to be absent in the Marianas Theseinclude the haplosclerids Haliclona (Reniera) poseidon (de Laubenfels)Gelliodes fibulatus (Carter) Oceanapia sagittaria (Sollas) Callyspongiaaerizusa Desqueyroux-Faundez and Cribrochalina olemda de Laubenfels thedictyoceratids Spongia matamata de Laubenfels Carteriospongia foliascens(Pallas) and Phyllospongia papyracea (Esper) the poecilosclerids Biemnatubulata (Dendy) the axinellid Axinella carteri (Dendy) species of DiacarnusTheonella and the calcareous sponges Pericharax heteroraphis Poleacutejaeff andLeucetta avocada de Laubenfels Many of these species have been recordedconsistently within various Indo-west Pacific reef systems from Vanuatu toThailand the Great Barrier Reef to Chuuk and represent some of the 5 of theregional fauna that is hypothesized to be widely distributed (eg Hooper et al2000) There is a growing body data however to suggest that at least somewidespread morphological species are comprised of genetically-divergentallopatric species complexes (eg Astrosclera willeyana ndash Woerheide et al2003)

Nevertheless part of the regional fauna is represented in the Marianas thusapproximately 30 (38) of the species recorded are regionally widespread (seeHooper et al 2000) The absence of numerous regional species is partly areflection of the lower diversity of sponges in the Marianas than on neighboringisland groups Thus CRRF has recorded 218 species of sponges from Chuuk inthe central Carolines and 295 species from Palau (L Colin pers comm 2001) Asurvey of 6 of the 80 known marine lakes in Palau resulted in more than 140sponge species (Kelly-Borges amp Valentine 1995) Kelly-Borges amp Valentine(1995) indicated that there are over 1000 published and undescribed species inthe tropical region of Oceania including Guam but this is a vast underestimateof the known fauna (JNH MK CRRF unpublished information) In comparisonHooper amp Leacutevi (1994) estimated that there are about 5000 published andundescribed species in the Australian insular and continental faunas alone

The relatively low diversity of sponges in the Marianas relative to Palau andthe Federated States of Micronesia fits general patterns seen in other marineorganisms (eg Randall 1995 Myers 1999) and reflects the isolation location

Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

App

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of t

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num

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of v

ouch

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all

spec

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Prov

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para

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Ref

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Phot

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umbe

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note

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6 l

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cite

dIs

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and

G =

Gua

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= R

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T =

Tin

ian

S =

Sai

pan

Taxo

nU

GI

Mus

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efPh

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LASS

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LCA

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ucet

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is D

endy

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8-I(

S)18

GP4

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N 0

988-

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mig

enia

Hae

ckel

187

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SNM

Ac

1837

33 (S

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ucet

ta sp

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) 0C

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)0C

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IDA

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irkpa

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orea

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otyp

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PM 9

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INID

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ophl

us sa

rasi

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hiel

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Kelly et al Sponges of the Marianas 105

and habitat-diversity of the archipelago First the Marianas lie further removedfrom the Indo-Malayan diversity center than Palau and neighboring islandsSecond the east-flowing Equatorial Counter Current (ECC) creates a biogeo-graphic corridor (the ldquoCaroline Conduitrdquo ndash Springer 1982) that flows through andenriches the biodiversity of the islands of Palau Federated States of Micronesiaand even Marshall Islands In contrast the Marianas are isolated from theinfluence of the ECC and bathed by the west-flowing North Equatorial CurrentFinally lagoons are poorly developed in the Marianas especially in comparisonwith the large barrier reef islands and atolls of the Caroline chain Lagoons hostthe most diverse and striking sponge faunas in the area Only one deep lagoonApra Harbor exists in the Marianas it hosts a specialized sponge fauna manyspecies of which are not known elsewhere in the archipelago

Approximately 30 (40) of the species recorded here are new to sciencebut this is not unusual as sponge faunas are underdescribed worldwide (egKelly-Borges amp Valentine 1995 Hooper et al 2000) Many of the new speciesrecognized within the Guam fauna are known by the authors from other locationsand few of the 40 species are likely to be endemic to the Marianas

ADVENTISMThe Marianas were among the first islands settled in Micronesia with clear

evidence for human occupation dating to 3500BP and indications of humanpresence to 4300BP (Kirch 2000) Magellanrsquos landing on Guam in 1521 was thefirst western contact in Oceania Thus Guam has had an especially prolongedopportunity to accumulate a non-indigenous biota (Paulay et al 2002) Todaycommercial shipping traffic to Guam originates mostly from the US mainland viaHawaii and to a lesser extent from Asia The two largest freight companiesMatson Shipping and CSX Lines route through Hawaii to Guam and return tothe US via Asia Smaller shipping lines however also connect Guam with severalAsian cities as well as with islands in Micronesia The arrival of two dry docksin the 1990rsquos the Adept from Subic Naval Station in the Philippines in 1992 andthe Machinist from Pearl Harbor Hawaii in 1999 have demonstrated theimportance of these structures in transporting fouling organisms Thus about 20species of sponges arrived in abundance from Hawaii on the hull of theMachinist Given what we know about the global transport of marine species(Carlton 1996) adventive sponges are to be expected to occur on Guam althoughtheir recognition is difficult

Recognition of non-indigenous status relies on accurate taxonomic iden-tifications together with a series of criteria established to evaluate evidence foradventism (Chapman amp Carlton 1991) Widely disjunct distributions are one ofthese criteria and two sponges on Guam Clathria mima and Hyrtios altum arerecorded in the literature from the Atlantic as well as the Indo-West PacificHowever Atlantic records of both are poorly documented and likely the artificialresult of poor taxonomy rather than of non-indigenous status Another criterionis restriction to artificial substrata including substrata associated with human

106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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106 Micronesica 35-36 2003

transport At least four sponges (Callyspongia aff fibrosa Mycale (Carmia) sp1 Niphates sp 1 and Tedania cf ignis) in Apra Harbor have been encounteredonly on artificial bottoms (buoys wharfs ship halls) and are not known to usfrom extensive sampling in the region or in the Indo-West Pacific Until thesesponges are encountered in a natural habitat their status remains to be accuratelyestablished thus at present they are best regarded as cryptogenic (cf Carlton1996) The presence of striking sponge Ianthella basta in Guam has led tospeculation about whether its population on Guam represents an isolated naturaloccurrence or whether it is adventive (Birkeland pers comm 1993) Ianthellabasta ranges from the Mascarene Islands to Vanuatu the Philippines and GuamThe absence of the species from all intervening Micronesian Islands includingPalau and the Federated States of Micronesia implies either that it colonizedGuam by jump dispersal or that it became established through anthropogenictransport On Guam the species is restricted to Apra Harbor the busiest port inMicronesia where it appears to be spreading If adventive Ianthella could havearrived from the Philippines an area linked to Guam with frequent shippingtraffic The environmental conditions in which the sponge is found in Guam areno different from that in other locations where it is common (Bergquist amp Kelly-Borges 1995)

In addition to these sponges numerous species previously unknown onGuam arrived on the hull of the dry dock Machinist in 1999 These sponges arecurrently under study by Ralph DeFelice (Bishop Museum) and their fate onGuam remains to be seen

AcknowledgementsWe thank Dame Professor Patricia R Bergquist Emeritus Professor of

Zoology University of Auckland New Zealand for identifications and adviceover the years on Guam and broader Micronesian sponge taxonomy The CoralReef Research Foundation Republic of Belau Micronesia contributed to ourknowledge of the sponge fauna of Guam through their collections in SaipanTinian and Rota Islands NIH Minority Biomedical Research Support grant GM-44796 to Valerie Paul funded Michelle Kelly to conduct a workshop on spongesat the University of Guam Marine Lab at Mangilao in 1996 Funding by the USDept of Defense at COMNAVMARIANAS and Sea Grant (SG-NIS-35) toGustav Paulay for biodiversity surveys is gratefully acknowledged This is acontribution of the Coral Reef Research Foundation Contribution 481 of theUniversity of Guam Marine Laboratory

Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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Kelly et al Sponges of the Marianas 107

ReferencesAgassiz A 1906 General report of the expedition Reports on the scientific

results of the expedition to the Eastern Tropical Pacific in charge of AAgassiz by the U S Fisheries Commission Steamer Albatross 1904-1905and 1888-1904 21 Memoirs of the Museum of Comparative Zoology Har-vard Collection 33 i-xiii 1-75 96 plates amp 8 figs in text

Amesbury S V Bonito R Chang L Kirkendale C Meyer G Paulay RRitson-Williams amp T Rongo 2001 Marine biodiversity resource survey andbaseline reef monitoring survey of the Haputo Ecological Reserve AreaCOMNAVMARIANAS Report and Interactive GIS Document Prepared forUS Dept of Defense COMNAVMARIANAS 111 pp amp CD-ROM

Avila C amp V J Paul 1997 Chemical ecology of the nudibranch Glossodorispallida is the location of diet-derived metabolites important for defenseMarine Ecology Progress Series 150 171-180

Becerro M A V J Paul amp J Starmer 1998 Intracolonial variation in chemicaldefenses of the sponge Cacospongia sp and its consequences on generalistfish predators and the specialist nudibranch predator Glossodoris pallidaMarine Ecology Progress Series 168 187-196

Bergquist P R 1965 The sponges of Micronesia Part I The Palau archipelagoPacific Science 19 123-204

Bergquist P R 1980 A revision of the supraspecific classification of the ordersDictyoceratida Dendroceratida and Verongida (class Demospongiae) NewZealand Journal of Zoology 7 443-503

Bergquist P R amp M Kelly-Borges 1991 An evaluation of the genus Tethya(Porifera Demospongiae Hadromerida) with descriptions of new speciesfrom the Southwest Pacific The Beagle Records of the Northern TerrritoryMuseum of Arts and Sciences 8 37-72

Bergquist P R amp M Kelly-Borges 1995 Systematics and Biogeography of thegenus Ianthella (Demospongiae Verongida Ianthellidae) in the South PacficThe Beagle Records of the Northern Territory Museum of Arts and Sciences12 151-176

Bryan P G 1973 Growth rate toxicity and distribution of the encrusting spongeTerpios sp (Hardomerida Suberitidae) in Guam Mariana Islands Microne-sica 9 237-242

Carlson C H amp P J Hoff 1973 Two new species of Gastropteridae fromGuam Marianas Islands (Opsithobranchia Cephalaspidea) Publications ofthe Seto Marine Biological Laboratory 21 141-151

Carlson C H amp P J Hoff 1974 The Gastropteridae of Guam with descriptionsof four new species (Opisthobranchia Cephalaspidea) Publications of theSeto Marine Biological Laboratory 21 345-363

Carlton J T 1996 Biological invasions and cryptogenic species Ecology 771653-1655

Chapman J W amp J T Carlton 1991 A test of criteria for introduced species

108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

App

endi

x 1

Spo

nges

of t

he M

aria

nas

UG

I ca

talo

gue

num

bers

of v

ouch

ers h

ouse

d in

Uni

v o

f Gua

m In

verte

brat

e co

llect

ions

all

spec

imen

s fro

m G

uam

M

useu

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cata

logu

e nu

mbe

rs o

f vo

uche

rs h

ouse

d at

per

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ent

repo

sito

ries

see

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hods

for

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Prov

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ce i

f ot

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than

Gua

m n

oted

para

nthe

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ly

Ref

ref

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bere

d re

fere

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follo

w a

t the

end

of t

he A

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dix

Phot

o p

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cher

s a

vaila

ble

at h

ttp

ww

wfl

mnh

ufl

edu

reef

s and

in th

e M

arin

e B

iodi

vers

ity o

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108 Micronesica 35-36 2003

the global invasion by the isopod Synidotea laevidorsalis (Miers 1881)Journal of Crustacean Biology 11 386-400

Davidson B S 1993 Variation in the secondary metabolites of the spongeDysidea cf avara MS Thesis in Biology University of Guam

de Laubenfels M W 1954 The sponges of the West Central Pacific OregonState Monographs in Zoology 7 1-306

Duffy J E amp V J Paul 1992 Prey nutritional quality and the effectiveness ofchemical defenses against tropical reef fishes Oecologia 90 333-339

Hartman W D amp T F Goreau 1975 A Pacific tabulate sponge livingrepresentative of a new order of sclerosponges Postilla Peabody MuseumYale University 167 1-14 figs 1-15

Hartman W D amp T F Goreau 1976 A new ceratoporellid sponge (PoriferaSclerospongiae) from the Pacific In F W Harrison amp R R Cowden (eds)Aspects of sponge biology pp 329-347 Academic Press New York

Hooper J N A 1991 Revision of the family Raspailiidae (Porifera Demospon-giae) with description of Australian species Invertebrate Taxonomy 51179-1415

Hooper J N A 1994 Coral reef sponges of the Sahul Shelf - a case for habitatpreservation Memoirs of the Queensland Museum 36 93-106

Hooper J N A amp C Leacutevi 1994 Biogeography of Indo-west Pacific spongesMicrocionidae Raspailiidae Axinellidae In R W M van Soest T M Gvan Kempen amp J C Brakeman (eds) Sponges in time and space biologychemistry paleontology Proceedings of the 4th International Porifera Con-gress Amsterdam Netherlands 19-23 1993 pp 191-212 Rotterdam AABalkema

Hooper J N A amp F Wiedenmayer 1994 Porifera Zoological catalogue ofAustralia Ed A Wells 632 pp CSIRO Melbourne Australia

Hooper J N A J A Kennedy amp R W M van Soest 2000 Annotatedchecklist of sponges (Porifera) of the South China Sea Region The RafflesBulletin of Zoology Supplement 8 125-207

Hooper J N A amp J A Kennedy 2002 Small-scale patterns of biodiversity insponges (Porifera) from the Sunshine Coast southeast QueenslandInvertebrate Systematics 16 637-653

Hooper J N A J A Kennedy amp R J Quinn 2002 Biodiversity hotspotspatterns of richness and endemism and taxonomic affinities of tropicalAustralian sponges (Porifera) Biodiversity and Conservation 11 851-885

Kelly-Borges M amp J Vacelet 1995 A revision of Diacarnus Burton andNegombata de Laubenfels (Demospongiae Latrunculiidae) with descriptionsof new species from the West Central Pacific and the Red Sea Memoirs ofthe Queensland Museum 38 477-503

Kelly-Borges M amp C Valentine 1995 The sponges of the tropical island regionof Oceania a taxonomic review In J E Maragos M N A Peterson L GEldredge J E Bardach and H F Takeuchi (eds) Marine and coastalbiodiversity in the tropical island Pacific region Volume 1 Species

Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

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n H

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ulay

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ture

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and

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m R

= R

ota

T =

Tin

ian

S =

Sai

pan

Taxo

nU

GI

Mus

eum

R

efPh

oto

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is D

endy

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Kelly et al Sponges of the Marianas 109

systematics and information management priorities pp 83-120 East-WestCenter University of Hawaii Honolulu

Kennedy J A 2000 Resolving the Jaspis stellifera complex Memoirs of theQueensland Museum 45 453-76

Kirch P 2000 On the roads of the winds An archaeological history of thePacific Islands Berkeley University of California Press

Lee H-S Y Seo J R Rho J Shin amp V J Paul 2001 New steroidal alkaloidsfrom an undescribed sponge of the genus Corticium Journal of NaturalProducts 64 1474-1476

Lee H-S Y Seo K W Cho J R Rho J Shin amp V J Paul 2000 Newtriterpenoid saponins from the sponge Melophlus isis Journal of NaturalProducts 63 915-919

Lendenfeld R von 1910a The sponges 1 The Geodiidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 1-258 48 pls

Lendenfeld R von 1910b The sponges 2 The Erylidae In Reports on thescientific results of the Expedition to the Eastern Tropical Pacific in chargeof Alexander Agassiz by the U S Fisheries Commission Steamer by theAlbatross 1904-1905 and of other expeditions of the Albatross 1888-190421 Memoirs of the Museum of Comparative Zoology Harvard Collection41 260-324 8 pls

Lendenfeld R von 1915 XXIX The sponges 3 Hexactinellida In Reports onthe scientific results of the Expedition to the Eastern Tropical Pacific incharge of Alexander Agassiz by the U S Fisheries Commission Steamer bythe Albatross 1904-1905 and of other expeditions of the Albatross 1888-1904 21 Memoirs of the Museum of Comparative Zoology HarvardCollection 42 1-397 109 pls

Myers R F 1999 Micronesian reef fishes 3rd revised and expanded editionCoral Graphics Barrigada Guam

Paul V J Y Seo K W Cho J-R Rho J Shin amp P R Bergquist 1997Sesquiterpenoids of the drimane class from a sponge of the genus DysideaJournal of Natural Products 60 1115-1120

Paulay G L Kirkendale G Lambert amp J Starmer 1997 The marineinvertebrate biodiversity of Apra Harbor significant areas and introducedspecies with focus on sponges echinoderms and ascidians Draft ReportPrepared for US Dept of Defense COMNAVMARIANAS 103 pp

Paulay G L Kirkendale C Meyer P Houk T Rongo amp R Chang 2001Marine biodiversity resource survey and baseline reef monitoring survey ofthe Southern Orote Peninsula and North Agat Bay Area COMNAV-MARIANAS Report and Interactive GIS Document Prepared for US Deptof Defense COMNAVMARIANAS 111 pp amp CD-ROM

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

App

endi

x 1

Spo

nges

of t

he M

aria

nas

UG

I ca

talo

gue

num

bers

of v

ouch

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ouse

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Uni

v o

f Gua

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M

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mbe

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ouse

d at

per

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Prov

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m n

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nthe

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Ref

ref

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follo

w a

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Phot

o p

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ww

wfl

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in th

e M

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iodi

vers

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f Gua

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D-R

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cop

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r tha

n G

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not

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Not

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s fol

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nd o

f App

endi

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ntifi

er 1

Mic

helle

Kel

ly 2

Joh

n H

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r 3

Rob

van

Soe

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4 P

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ergq

uist

5 G

usta

v Pa

ulay

6 l

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ture

cite

dIs

isl

and

G =

Gua

m R

= R

ota

T =

Tin

ian

S =

Sai

pan

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsC

LASS

CA

LCA

REA

CLA

THR

INID

A L

EUC

ETTI

DA

ELe

ucet

ta c

hago

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is D

endy

191

361

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DN

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8-I(

S)18

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1G

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Leuc

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pri

mig

enia

Hae

ckel

187

2U

SNM

Ac

1837

33 (S

)1

16

SLe

ucet

ta sp

1 (n

sp

) 0C

DN

100

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)0C

DN

100

3-Y

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RR

AY

ON

IDA

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RR

AY

ON

IDA

EM

urra

yona

pha

nole

pis K

irkpa

trick

191

060

830C

DN

098

0-X

(S)

GP4

17-2

(S)

0CD

N 0

980-

X (S

)1

5G

S

CLA

SS D

EMO

SPO

NG

IAE

AG

ELA

SID

A A

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ASI

IDA

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elas

sp 1

60

820C

DN

097

0-N

(S)

GP5

87-6

0C

DN

097

0-N

(S)

14

GS

Agel

as sp

2

6800

QM

G31

7348

GP8

63-1

82

GA

GEL

ASI

DA

AST

RO

SCLE

RID

AE

Astr

oscl

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eyan

a Li

ster

190

060

8522

(S)

GP5

72-3

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GS

Stro

mat

ospo

ngia

mic

rone

sica

Har

tman

amp G

orea

u 1

976

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otyp

e Y

PM 9

104

Y

PM 9

110(

S)5

GP6

49-1

0

0CD

N 0

983-

A (S

)6

GS

AST

RO

PHO

RID

A A

NC

OR

INID

AE

Mel

ophl

us sa

rasi

noru

m T

hiel

e 1

899

5943

BM

NH

199

56

225

11G

P524

-36

1G

Pena

res n

ux (d

e La

uben

fels

195

4)59

40B

MN

H 1

995

622

4G

P274

-18

5G

110 Micronesica 35-36 2003

Paulay G L Kirkendale G Lambert amp C Meyer 2002 Anthropogenic bioticinterchange in a coral reef ecosystem a case study from Guam PacificScience 56 403-422

Pennings S C S R Pablo V J Paul amp J E Duffy 1994 Effects of spongesecondary metabolites in different diets on feeding by three groups of con-sumers Journal of Experimental Marine Biology and Ecology 180 137-149

Plubrukarn A D W Smith R E Cramer and B S Davidson 1997 (2E9E)-pyronaamidine 9-(N-methylimine) a new imidazole alkaloid from theNorthern Mariana Islands sponge Leucetta sp cf chagosensis Journal ofNatural Products 60 712-715

Plucer-Rosario G 1987 The effect of substratum on the growth of Terpios anencrusting sponge that kills corals Coral Reefs 5 197-200

Pordesimo E O amp F J Schmitz 1990 New bastadins from the spongeIanthella basta Journal of Organic Chemistry 55 4704-4709

Quinn N J amp B L Kojis 1999 Community structure of the living fossilcoralline sponge populations at the Grotto Saipan Northern Mariana IslandsBulletin of Marine Science 65 227-234

Randall R H 1995 Biogeography of reef-building corals in the Mariana andPalau Islands in relation to back-arc rifting and the formation of the easternPhilippine Sea Natural History Research Natural History Museum andInstitute Chiba 3 193-210

Ridley S O 1884 Spongiida Report on the zoological collection made in theIndo-Pacific Ocean during the Voyage of HMS Alert 1881-2 366-482582-635 British Museum (Natural History) London

Ridley S O amp A Dendy 1886 Preliminary report on the Monaxonida collectedby the HMS Challenger Annals and Magazine of Natural History (5) 18325-352 470-493

Ridley S O amp A Dendy 1887 Report on the Monaxonida collected by theHMS Challenger during the years 1873-76 Report on the scientific resultsof the HMS Challenger during the years 1873-76 20 1-275 Her Majes-tys Stationary Office London

Rogers S D 1989 Feeding preferences and chemical defenses of threeGlossodoris nudibranchs and their diet sponges MS Thesis in BiologyUniversity of Guam

Rogers S D amp V J Paul 1991 Chemical defenses of three Glossodorisnudibranchs and their dietary Hyrtios sponges Marine Ecology ProgressSeries 77 221-232

Ruumltzler K amp K Muzik 1993 Terpios hoshinota a new cyanobacteriospongethreatening Pacific reefs Scientia Marina 57 395-403

Schmitz F J S K Agarwal S P Gunasekera P G Schmidt amp J N Shoolery1983 Amphimedine new aromatic alkaloid from a Pacific spongeAmphimedon sp Carbon connectivity determination from natural abundance13C-13C coupling constants Journal of the American Chemical Society 1054835-4836

Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

App

endi

x 1

Spo

nges

of t

he M

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nas

UG

I ca

talo

gue

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bers

of v

ouch

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d in

Uni

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Prov

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Ref

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Phot

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ttp

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Joh

n H

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r 3

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usta

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ulay

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ture

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and

G =

Gua

m R

= R

ota

T =

Tin

ian

S =

Sai

pan

Taxo

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GI

Mus

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R

efPh

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Not

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ta c

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sens

is D

endy

191

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620C

DN

098

8-I(

S)18

GP4

07-1

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0CD

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988-

I (S)

1G

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Leuc

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187

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830C

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0-X

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980-

X (S

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60

820C

DN

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0-N

(S)

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87-6

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097

0-N

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14

GS

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Astr

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190

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Stro

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Kelly et al Sponges of the Marianas 111

Schmitz F J S P Gunasekera V Lakshmi amp L M V Tillekeratne 1985Marine natural products pyrrololactams from several sponges Journal ofNatural Products 48 47-53

Shin J Y Seo K W Cho J-R Rho amp V J Paul 1998 Osirisynes A-F highlyoxygenated polyacetylenes from the sponge Haliclona osiris Tetrahedron54 8711-8720

Shin J H-S Lee Y Seo J-R Rho K W Cho amp V J Paul 2000 Newbromotyrosine metabolites from the sponge Aplysinella rhax Tetrahedron56 9071-9077

Smyth M J 1990 Incidence of boring organisms in gastropod shells on reefsaround Guam Bulletin of Marine Science 46 432-49

Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp

Thacker R W M A Becerro W A Lumbang amp V J Paul 1998 Allelopathicinteractions between sponges on a tropical reef Ecology 79 1740-1750

Wilson H V 1925 Siliceous and horny sponges collected by the US FisheriesSteamer lsquoAlbatrossrsquo during the Philippine Expedition 1907-1910 InContributions to the biology of the Philippine Archipelago and adjacentregions Bulletin of the United States National Museum 100 273-532

Whitelegge T 1901 Report on Sponges from the Coastal Beaches of New SouthWales Records of the Australian Museum 4 (2) 1-70 [55-118]

Woerheide G B M Degnan J N A Hooper amp J Reitner 2003Biogeography and taxonomy of the Indo-Pacific reef cave dwelling corallinedemosponge Astrosclera lsquowilleyanarsquo new data from nuclear internaltranscribed spacer sequences Proceedings of the 9th International Coral ReefSymposium Bali 1 339-346

Received 18 October 2001

App

endi

x 1

Spo

nges

of t

he M

aria

nas

UG

I ca

talo

gue

num

bers

of v

ouch

ers h

ouse

d in

Uni

v o

f Gua

m In

verte

brat

e co

llect

ions

all

spec

imen

s fro

m G

uam

M

useu

m

cata

logu

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mbe

rs o

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Prov

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Ref

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t the

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Phot

o p

hoto

vou

cher

s a

vaila

ble

at h

ttp

ww

wfl

mnh

ufl

edu

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s and

in th

e M

arin

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f Gua

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D-R

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Not

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low

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f App

endi

xID

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ntifi

er 1

Mic

helle

Kel

ly 2

Joh

n H

oope

r 3

Rob

van

Soe

st a

nd W

allie

de

Wee

rdt

4 P

atric

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uist

5 G

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v Pa

ulay

6 l

itera

ture

cite

dIs

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G =

Gua

m R

= R

ota

T =

Tin

ian

S =

Sai

pan

Taxo

nU

GI

Mus

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R

efPh

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Not

esID

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endy

191

361

620C

DN

098

8-I(

S)18

GP4

07-1

1

0CD

N 0

988-

I (S)

1G

S

Leuc

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pri

mig

enia

Hae

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187

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Ac

1837

33 (S

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16

SLe

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1 (n

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DN

100

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100

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EM

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191

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830C

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0-X

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17-2

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980-

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)1

5G

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CLA

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EMO

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IAE

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ELA

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sp 1

60

820C

DN

097

0-N

(S)

GP5

87-6

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DN

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0-N

(S)

14

GS

Agel

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2

6800

QM

G31

7348

GP8

63-1

82

GA

GEL

ASI

DA

AST

RO

SCLE

RID

AE

Astr

oscl

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will

eyan

a Li

ster

190

060

8522

(S)

GP5

72-3

7 5

GS

Stro

mat

ospo

ngia

mic

rone

sica

Har

tman

amp G

orea

u 1

976

Hol

otyp

e Y

PM 9

104

Y

PM 9

110(

S)5

GP6

49-1

0

0CD

N 0

983-

A (S

)6

GS

AST

RO

PHO

RID

A A

NC

OR

INID

AE

Mel

ophl

us sa

rasi

noru

m T

hiel

e 1

899

5943

BM

NH

199

56

225

11G

P524

-36

1G

Pena

res n

ux (d

e La

uben

fels

195

4)59

40B

MN

H 1

995

622

4G

P274

-18

5G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

UG

I ca

talo

gue

num

bers

of v

ouch

ers h

ouse

d in

Uni

v o

f Gua

m In

verte

brat

e co

llect

ions

all

spec

imen

s fro

m G

uam

M

useu

m

cata

logu

e nu

mbe

rs o

f vo

uche

rs h

ouse

d at

per

man

ent

repo

sito

ries

see

met

hods

for

cod

es

Prov

enan

ce i

f ot

her

than

Gua

m n

oted

para

nthe

tical

ly

Ref

ref

eren

ces

num

bere

d re

fere

nces

follo

w a

t the

end

of t

he A

ppen

dix

Phot

o p

hoto

vou

cher

s a

vaila

ble

at h

ttp

ww

wfl

mnh

ufl

edu

reef

s and

in th

e M

arin

e B

iodi

vers

ity o

f Gua

m C

D-R

OM

cop

ublic

atio

n P

rove

nanc

e if

othe

r tha

n G

uam

not

ed p

aran

thet

ical

ly

Not

es n

umbe

red

note

s fol

low

at e

nd o

f App

endi

xID

ide

ntifi

er 1

Mic

helle

Kel

ly 2

Joh

n H

oope

r 3

Rob

van

Soe

st a

nd W

allie

de

Wee

rdt

4 P

atric

ia B

ergq

uist

5 G

usta

v Pa

ulay

6 l

itera

ture

cite

dIs

isl

and

G =

Gua

m R

= R

ota

T =

Tin

ian

S =

Sai

pan

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsC

LASS

CA

LCA

REA

CLA

THR

INID

A L

EUC

ETTI

DA

ELe

ucet

ta c

hago

sens

is D

endy

191

361

620C

DN

098

8-I(

S)18

GP4

07-1

1

0CD

N 0

988-

I (S)

1G

S

Leuc

etta

pri

mig

enia

Hae

ckel

187

2U

SNM

Ac

1837

33 (S

)1

16

SLe

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ta sp

1 (n

sp

) 0C

DN

100

3-(R

)0C

DN

100

3-Y

(R)

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MU

RR

AY

ON

IDA

MU

RR

AY

ON

IDA

EM

urra

yona

pha

nole

pis K

irkpa

trick

191

060

830C

DN

098

0-X

(S)

GP4

17-2

(S)

0CD

N 0

980-

X (S

)1

5G

S

CLA

SS D

EMO

SPO

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IAE

AG

ELA

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IDA

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sp 1

60

820C

DN

097

0-N

(S)

GP5

87-6

0C

DN

097

0-N

(S)

14

GS

Agel

as sp

2

6800

QM

G31

7348

GP8

63-1

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GA

GEL

ASI

DA

AST

RO

SCLE

RID

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Astr

oscl

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eyan

a Li

ster

190

060

8522

(S)

GP5

72-3

7 5

GS

Stro

mat

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mic

rone

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Har

tman

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976

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PM 9

104

Y

PM 9

110(

S)5

GP6

49-1

0

0CD

N 0

983-

A (S

)6

GS

AST

RO

PHO

RID

A A

NC

OR

INID

AE

Mel

ophl

us sa

rasi

noru

m T

hiel

e 1

899

5943

BM

NH

199

56

225

11G

P524

-36

1G

Pena

res n

ux (d

e La

uben

fels

195

4)59

40B

MN

H 1

995

622

4G

P274

-18

5G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsPe

nare

s sol

lasi

Thi

ele

190

360

95ZM

A 1

6415

GP3

80-2

13

GRh

abda

stre

lla g

lobo

stel

lata

(Car

ter

1883

)60

94B

MN

H 1

995

622

2G

P651

-14

171

5G

CH

ON

DR

OSI

DA

CH

ON

DR

ILLI

DA

EC

hond

rilla

aus

tral

iens

is C

arte

r 18

7360

10B

MN

H 1

995

622

37

U

SNM

Ac

1837

33 (S

)1

16

GS

Cho

ndri

lla g

rand

iste

llata

Thi

ele

190

0U

SNM

Ac

1837

33 (S

)1

16

SC

hond

rosi

a co

rtic

ata

Thie

le 1

900

6707

NIW

A 9

15 Q

M G

3173

46G

P763

-13

12

GD

END

RO

CER

ATI

DA

DA

RW

INEL

LID

AE

Che

lona

plys

illa

sp 1

(gre

y)

5977

BM

NH

199

56

222

7G

P531

-27

1G

Che

lona

plys

illa

sp 2

(pur

ple)

59

78B

MN

H 1

995

622

28

GP5

13-1

0 1

GD

END

RO

CER

ATI

DA

DIC

TYO

DEN

DR

ILLI

DA

ED

icty

oden

drill

a ni

gra

(de

Laub

enfe

ls 1

954)

0CD

N 0

973-

Q0C

DN

097

3-Q

(S)

191

SD

ICTY

OC

ERA

TID

A D

YSI

DEI

DA

ED

ysid

ea a

rena

ria

Ber

gqui

st 1

965

BM

NH

199

56

223

01

GD

ysid

ea sp

1A

(n s

p a

ff h

erba

cea

(Kel

ler

1889

) rid

ged)

G

P741

-29

1G

Dys

idea

sp 1

B (n

sp

aff

her

bace

a (K

elle

r 18

89) s

moo

th)

23G

P573

-6

1G

Dys

idea

sp 2

(n s

p)

BM

NH

199

75

131

12 1

6G

P389

-33

4G

Dys

idea

sp 3

(n s

p a

ff a

vara

(Sch

mid

t 18

62))

60

48B

MN

H 1

995

622

8G

P869

-32

251

GD

ysid

ea sp

4 (n

sp

aff

gra

nulo

sa B

ergq

uist

196

5)60

09

6049

BM

NH

199

56

227

N

IWA

909

142

324

GP7

68-2

5 11

1G

Dys

idea

sp 5

(n s

p c

ave)

N

IWA

917

GP7

40-2

71

GD

ICTY

OC

ERA

TID

A I

RC

INII

DA

EIr

cini

a sp

166

08N

IWA

QM

GP8

62-2

84

GIr

cini

a sp

266

14G

P838

-25

2G

Sarc

otra

gus s

p 1

(n s

p)

0CD

N 0

971-

O(R

)0C

DN

097

1-O

(R)

1R

Sarc

otra

gus s

p 2

(n s

p)

0CD

N 1

014-

M(R

)0C

DN

101

4-M

(R)

1R

DIC

TYO

CER

ATI

DA

SPO

NG

IID

AE

Cos

cino

derm

a cf

mat

hew

si (L

ende

nfel

d 1

886)

NIW

A 9

24 amp

925

GP7

63-1

31

GH

ippo

spon

gia

met

achr

omia

de

Laub

enfe

ls 1

954

6619

NIW

A 9

21 0

CD

N 0

996-

Q(S

)G

P615

-27

0C

DN

099

6-Q

(S)

1G

S

Stre

psic

hord

aia

lend

enfe

ldi B

ergq

uist

et a

l 1

988

0CD

N 0

995-

P(R

)0C

DN

099

5-P

(R)

1R

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsD

ICTY

OC

ERA

TID

A T

HO

REC

TID

AE

Aply

sino

psis

cf

eleg

ans L

ende

nfel

d 1

888

6706

NIW

A 9

11

GP6

49-2

1G

Cac

ospo

ngia

sp 1

(n s

p)

6140

BM

NH

199

56

222

39

10G

P530

-12

261

GFa

scap

lysi

nops

is re

ticul

ata

(Hen

tsch

el 1

912)

6609

NIW

A 9

19 0

CD

N 0

984-

C(S

)G

P407

-13

0C

DN

098

4-C

(S)

1G

S

Hyr

tios a

ltum

(Pol

eacutejae

ff 1

884)

6004

BM

NH

199

56

222

4 G

P532

-9

4G

Hyr

tios e

rect

a (K

elle

r 18

89)

USN

M 3

1754

13

GP6

44-2

264

GH

yrtio

s sp

1

BM

NH

199

56

222

54

GLu

ffari

ella

geo

met

rica

Kirk

patri

ck 1

910

0CD

N 0

982-

Z(S)

0CD

N 0

982-

Z (S

)1

SLu

ffari

ella

var

iabi

lis (P

oleacutej

aeff

188

4)60

81B

MN

H 1

995

622

26

141

5G

P573

-26

1G

HA

DR

OM

ERID

A A

CA

NTH

OC

HA

ETET

IDA

EAc

anth

ocha

etet

es w

ells

i Har

tman

amp G

orea

u 1

975

6615

Hol

otyp

e Y

PM 9

077

422

(S)

GP6

49-7

6G

SH

AD

RO

MER

IDA

CLI

ON

IDA

EAl

ecto

na sp

1

22

6G

Clio

na e

nsife

ra (S

olla

s 18

88)

22

6G

Clio

na sp

1

22

6G

Clio

na v

astif

ica

(Han

cock

184

9)2

26

GC

liona

vir

idis

Sch

mid

t 18

622

26

GTh

oosa

bul

bosa

Han

cock

184

92

26

GH

AD

RO

MER

IDA

HEM

IAST

EREL

LID

AE

Hem

iast

erel

la ty

pus C

arte

r 18

7167

09ZM

A 1

6407

GP2

61B

-23

3G

HA

DR

OM

ERID

A S

PIR

AST

REL

LID

AE

Sphe

cios

pong

ia g

lobu

lari

s Den

dy 1

921

669

0B

MN

H 1

995

622

38

1G

Sphe

cios

pong

ia v

agab

unda

(Rid

ley

188

4)60

08B

MN

H 1

995

622

39

GP2

70-3

61

GSp

iras

trel

la p

otam

ophe

ra d

e La

uben

fels

195

4U

SNM

Ac

1837

33 (S

)1

16

SH

AD

RO

MER

IDA

SU

BER

ITID

AE

Terp

ios a

ploo

s de

Laub

enfe

ls 1

954

6612

Hol

otyp

e U

SNM

231

411

46

GTe

rpio

s hos

hino

ta R

uumltzl

er amp

Mus

ik 1

993

BM

NH

199

56

221

16

78

GP3

62-2

6G

Terp

ios s

p 1

N

IWA

923

GP7

61-5

1G

HA

DR

OM

ERID

A T

ETH

YID

AE

Teth

ya se

yche

llens

is W

right

188

1U

SNM

Ac

1837

33 (S

)1

166

STe

thya

sp(p

) 62

17G

P400

-216

5G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsH

ALI

CH

ON

DR

IDA

AX

INEL

LID

AE

Acan

thel

la c

aver

nosa

Den

dy 1

922

5975

59

32B

MN

H 1

995

622

12

GP5

30-3

11

G

Axin

ella

sp 1

(n s

p)

6163

60

15B

MN

H 1

995

622

40

Q

M G

3173

55G

P752

-12

G

Dra

gmac

idon

sp 1

(n s

p)

6802

QM

G31

7357

0CD

N 0

959-

Z (S

)G

P868

-10

0C

DN

095

9-Z

(S)

12

G S

Styl

issa

mas

sa (C

arte

r 18

89)

5942

BM

NH

199

56

221

3

0CD

N 0

960-

A (S

) 1

27G

P868

-31

51

GS

HA

LIC

HO

ND

RID

A D

ESM

OX

YID

AE

Myr

mek

iode

rma

gran

ulat

a (E

sper

183

0)

6138

BM

NH

199

75

133

0C

DN

099

0-K

(S)

GP5

30-1

5

0CD

N 0

990-

K (S

)1

GS

HA

LIC

HO

ND

RID

A D

ICTY

ON

ELLI

DA

ELi

osin

a cf

gra

nula

ris K

elly

Bor

ges amp

Ber

gqui

st 1

988

6042

BM

NH

199

56

221

0G

P868

-24

1G

HA

LIC

HO

ND

RID

A H

ALI

CH

ON

DR

IID

AE

Amor

phin

opsi

s sp

1

22

6G

Axin

yssa

acu

leat

a W

ilson

192

5ZM

A 1

6419

GP6

51-3

03

GAx

inys

sa sp

1 (n

sp

) 66

91Q

M G

3173

52G

P582

-33

2G

HA

PLO

SCLE

RID

A C

ALL

YSP

ON

GII

DA

E C

ally

spon

gia

diffu

sa (R

idle

y 1

884)

USN

M 2

3143

16

6G

Cal

lysp

ongi

a sp

1 (a

ff c

aren

s Pul

itzer

-Fin

ali

1982

ndashR

vS 4

2)66

87ZM

A 1

6410

GP7

35-2

13

GC

ally

spon

gia

sp 2

(aff

fib

rosa

Rid

ley

amp D

endy

188

7)66

88ZM

A 1

6411

GP5

14-3

03

GC

ally

spon

gia

suba

rmig

era

Rid

ley

188

466

95ZM

A 1

6409

GP4

59-2

83

GEu

plac

ella

sp 1

(n s

p)

0CD

N 0

972-

P(S)

0CD

N 0

972-

P (S

)1

SSi

phon

ocha

lina

cras

sifib

ra D

endy

188

960

46

6045

B

MN

H 1

995

622

19

ZM

A 1

6404

GP5

36-1

53

G

HA

PLO

SCLE

RID

A C

HA

LIN

IDA

EAd

ocia

nee

ns (T

opse

nt 1

918)

USN

M A

c18

3733

(S)

11

6S

Hal

iclo

na (G

elliu

s) g

raci

lis (H

ents

chel

191

2)67

05ZM

A 1

6420

GP4

588

3G

Hal

iclo

na (R

enie

ra) v

iola

(de

Laub

enfe

ls 1

954)

6616

Hol

otyp

e U

SNM

231

42

ZMA

164

031

136

G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsH

alic

lona

(Soe

stel

la) s

p 2

(n s

p b

lue

aqua

rium

Hal

iclo

na)

6043

59

83B

MN

H 1

995

622

18

ZM

A 1

6413

GP3

61-2

63

G

Hal

iclo

na li

gula

ta (W

hite

legg

e 1

901)

U

SNM

231

441

122

6G

Hal

iclo

na o

siri

s (de

Lau

benf

els

1954

)59

74

5929

BM

NH

199

56

221

4

USN

M 3

1763

ZM

A 1

6422

20G

P288

-51

G

Hal

iclo

na sp

1 (o

rang

e)

5972

59

34B

MN

H 1

995

622

17

ZM

A 1

6423

GP3

61-3

01

G

Hal

iclo

na sp

4 -

(n s

p c

oral

line

spon

ge)

6692

NIW

A 9

121

GH

alic

lona

stre

ble

de L

aube

nfel

s 19

54H

olot

ype

USN

M 2

3139

(S)

11

6S

HA

PLO

SCLE

RID

A N

IPH

ATI

DA

EAm

phim

edon

aff

vir

idis

Duc

hass

aing

amp M

iche

lotti

186

460

40ZM

A 1

6406

GP3

88-1

63

GN

ipha

tes s

p 1

(n s

p)

6192

NIW

A 9

13 9

16G

P536

-91

GN

ipha

tes s

p 2

(n s

p)

6807

NIW

A 9

18G

P868

-27

1G

Nip

hate

s sp

3 (w

ith si

gmas

) 61

03ZM

A 1

6421

GP5

77-2

86

3G

HA

PLO

SCLE

RID

A P

ETR

OSI

IDA

EPe

tros

ia sp

1

6698

ZMA

164

08G

P781

-24

3G

Petr

osia

sp 2

(aff

aru

ensi

s Hen

tsch

el 1

912)

6689

ZMA

164

12G

P649

-63

GXe

stos

pong

ia e

xigu

a (K

irkpa

trick

190

0)

6016

BM

NH

199

56

223

6G

P273

-33

1G

Xest

ospo

ngia

sp 1

65

74ZM

A 1

6417

GP8

69-1

33

GXe

stos

pong

ia sp

2 (a

ff c

arbo

nari

a (L

amar

ck 1

814)

)59

98B

MN

H 1

995

622

22

ZM

A 1

6416

28G

P532

-421

3G

HA

PLO

SCLE

RID

A P

HLO

EOD

ICTY

IDA

EAk

a sp

2 (y

ello

w)

6024

GP4

07-3

35

GAk

a tr

achy

s de

Laub

enfe

ls 1

954

Hol

otyp

e U

SNM

231

461

16

GPh

loeo

dict

yida

e N

ew g

enus

sp 1

(n s

p)

GP8

24-7

0C

DN

098

5-F(

S)0C

DN

098

5-F

(S)

141

GS

Oce

anap

ia

sp 1

(cre

vice

stic

k sp

onge

) 67

02G

P261

B-2

21

GldquoP

ellin

ardquo p

ulvi

lla (T

hiel

e 1

903)

USN

M 2

3145

11

6G

HO

MO

SCLE

RO

PHO

RID

A P

LAK

INID

AE

Cor

ticiu

m sp

1 (n

sp

) 68

06N

IWA

914

17G

P868

-51

GPl

akin

a sp

1

6078

BM

NH

199

56

221

GP2

88-6

1G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsldquoL

ITH

ISTI

DA

rdquo A

ZOR

ICID

AE

Leio

derm

atiu

m sp

122

6S

ldquoLIT

HIS

TID

Ardquo

SC

LER

ITO

DER

MID

AE

Acic

ulite

s pap

illat

a Leacute

vi amp

Leacutev

i 19

830C

DN

101

5-N

(R)

0CD

N 1

015-

N (R

)1

RldquoL

ITH

ISTI

DA

rdquo T

HEO

NEL

LID

AE

Theo

nelli

dae

new

gen

us sp

1 (n

sp

) 0C

DN

098

9-J(

S)0C

DN

098

9-J (

S)12

1S

POEC

ILO

SCLE

RID

A C

RA

MB

IID

AE

Mon

anch

ora

clat

hrat

a (C

arte

r 18

83)

6809

QM

G31

7344

GP8

70-1

72

GPO

ECIL

OSC

LER

IDA

CR

ELLI

DA

EC

rella

cya

thop

hora

(Car

ter

1880

) 67

93Q

M G

3173

54

ZMA

164

18G

P253

-20

3G

POEC

ILO

SCLE

RID

A D

ESM

AC

ELLI

DA

EN

eofib

ular

ia h

artm

ani H

oope

r amp L

eacutevi

1993

NIW

A 9

22G

P838

-26

1G

POEC

ILO

SCLE

RID

A M

ICR

OC

ION

IDA

EC

lath

ria

(Cla

thri

a) b

asila

na L

evi

1961

6190

GP5

36-1

110

2G

Cla

thri

a (M

icro

cion

a) m

ima

(de

Laub

enfe

ls 1

954)

6710

QM

G31

7343

GP6

16-9

2G

Cla

thri

a (T

haly

sias

) cf

hirs

uta

Hoo

per amp

Lev

i 19

9359

80B

MN

H 1

995

622

34

GP8

69-3

410

2G

Cla

thri

a (T

haly

sias

) cf

rein

war

dti V

osm

aer

1880

6696

QM

G31

7353

GP5

77-3

57

2G

Cla

thri

a (T

haly

sias

) eur

ypa

(de

Laub

enfe

ls 1

954)

GP8

68-3

01

GC

lath

ria

(Tha

lysi

as) s

p 1

(n s

p J

NA

H re

f 24

31)

6810

QM

G31

7351

GP8

70-1

22

GC

lath

ria

(Tha

lysi

as) v

ulpi

na (L

amar

ck 1

814)

USN

M 2

3149

13

76

GC

lath

ria

(Wils

onel

la) s

p 2

(n s

p J

NA

H re

f 17

66)

6711

QM

G31

7365

GP7

46-1

22

GPO

ECIL

OSC

LER

IDA

MY

CA

LID

AE

Myc

ale

(Car

mia

) cec

ilia

de L

aube

nfel

s 19

540C

DN

096

4-H

0CD

N 0

964-

H (S

)1

SM

ycal

e (C

arm

ia) s

p 1

(n s

p J

NA

H re

f 21

99)

6796

QM

G31

7358

GP5

36-1

32

GU

losa

spon

gia

de L

aube

nfel

s 19

54

NIW

A 9

910

GP6

51-3

61

GPO

ECIL

OSC

LER

IDA

MY

XIL

LID

AE

Iotr

ocho

ta b

acul

ifera

Rid

ley

188

466

17B

MN

H 1

995

622

16

GP8

68-1

71

GIo

troc

hota

ditr

ocho

ta (d

e La

uben

fels

195

4)G

P288

-28

4G

POEC

ILO

SCLE

RID

A R

ASP

AIL

IID

AE

Cer

atop

sion

sp 1

(n s

p)

5938

BM

NH

199

56

224

2

QM

G31

7364

GP6

51-6

12

G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsEc

hino

dict

yum

ant

rode

s (de

Lau

benf

els

1954

)U

SNM

231

481

86

GPO

ECIL

OSC

LER

IDA

TED

AN

IID

AE

Teda

nia

cf i

gnis

(Duc

hass

aing

amp M

iche

lotti

186

4)61

94Q

M G

3173

56G

P536

-10

2G

Teda

nia

mea

ndri

ca T

hiel

e 1

903

5985

BM

NH

199

56

223

5

ZMA

164

25G

P361

-29

3G

Teda

nia

sp 1

(n s

p)

6799

BM

NH

199

56

229

Q

M G

3173

47G

P866

-14

2G

SPIR

OPH

OR

IDA

TET

ILLI

DA

EC

inac

hyra

por

osa

(Len

denf

eld

188

8)U

SNM

231

38 (S

)1

96

SC

inac

hyra

sp(p

) 59

93B

MN

H 1

995

622

6G

P578

-27

91

GV

ERO

NG

IDA

APL

YSI

NEL

LID

AE

Aply

sine

lla rh

ax (d

e La

uben

fels

195

4)60

57B

MN

H 1

995

622

30

21G

P514

-12

201

GV

ERO

NG

IDA

PSE

UD

OC

ERA

TIN

IDA

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7)66

10G

P417

-33

(S)

5G

Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
  • GP768-25
  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

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N 0

959-

Z (S

)G

P868

-10

0C

DN

095

9-Z

(S)

12

G S

Styl

issa

mas

sa (C

arte

r 18

89)

5942

BM

NH

199

56

221

3

0CD

N 0

960-

A (S

) 1

27G

P868

-31

51

GS

HA

LIC

HO

ND

RID

A D

ESM

OX

YID

AE

Myr

mek

iode

rma

gran

ulat

a (E

sper

183

0)

6138

BM

NH

199

75

133

0C

DN

099

0-K

(S)

GP5

30-1

5

0CD

N 0

990-

K (S

)1

GS

HA

LIC

HO

ND

RID

A D

ICTY

ON

ELLI

DA

ELi

osin

a cf

gra

nula

ris K

elly

Bor

ges amp

Ber

gqui

st 1

988

6042

BM

NH

199

56

221

0G

P868

-24

1G

HA

LIC

HO

ND

RID

A H

ALI

CH

ON

DR

IID

AE

Amor

phin

opsi

s sp

1

22

6G

Axin

yssa

acu

leat

a W

ilson

192

5ZM

A 1

6419

GP6

51-3

03

GAx

inys

sa sp

1 (n

sp

) 66

91Q

M G

3173

52G

P582

-33

2G

HA

PLO

SCLE

RID

A C

ALL

YSP

ON

GII

DA

E C

ally

spon

gia

diffu

sa (R

idle

y 1

884)

USN

M 2

3143

16

6G

Cal

lysp

ongi

a sp

1 (a

ff c

aren

s Pul

itzer

-Fin

ali

1982

ndashR

vS 4

2)66

87ZM

A 1

6410

GP7

35-2

13

GC

ally

spon

gia

sp 2

(aff

fib

rosa

Rid

ley

amp D

endy

188

7)66

88ZM

A 1

6411

GP5

14-3

03

GC

ally

spon

gia

suba

rmig

era

Rid

ley

188

466

95ZM

A 1

6409

GP4

59-2

83

GEu

plac

ella

sp 1

(n s

p)

0CD

N 0

972-

P(S)

0CD

N 0

972-

P (S

)1

SSi

phon

ocha

lina

cras

sifib

ra D

endy

188

960

46

6045

B

MN

H 1

995

622

19

ZM

A 1

6404

GP5

36-1

53

G

HA

PLO

SCLE

RID

A C

HA

LIN

IDA

EAd

ocia

nee

ns (T

opse

nt 1

918)

USN

M A

c18

3733

(S)

11

6S

Hal

iclo

na (G

elliu

s) g

raci

lis (H

ents

chel

191

2)67

05ZM

A 1

6420

GP4

588

3G

Hal

iclo

na (R

enie

ra) v

iola

(de

Laub

enfe

ls 1

954)

6616

Hol

otyp

e U

SNM

231

42

ZMA

164

031

136

G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsH

alic

lona

(Soe

stel

la) s

p 2

(n s

p b

lue

aqua

rium

Hal

iclo

na)

6043

59

83B

MN

H 1

995

622

18

ZM

A 1

6413

GP3

61-2

63

G

Hal

iclo

na li

gula

ta (W

hite

legg

e 1

901)

U

SNM

231

441

122

6G

Hal

iclo

na o

siri

s (de

Lau

benf

els

1954

)59

74

5929

BM

NH

199

56

221

4

USN

M 3

1763

ZM

A 1

6422

20G

P288

-51

G

Hal

iclo

na sp

1 (o

rang

e)

5972

59

34B

MN

H 1

995

622

17

ZM

A 1

6423

GP3

61-3

01

G

Hal

iclo

na sp

4 -

(n s

p c

oral

line

spon

ge)

6692

NIW

A 9

121

GH

alic

lona

stre

ble

de L

aube

nfel

s 19

54H

olot

ype

USN

M 2

3139

(S)

11

6S

HA

PLO

SCLE

RID

A N

IPH

ATI

DA

EAm

phim

edon

aff

vir

idis

Duc

hass

aing

amp M

iche

lotti

186

460

40ZM

A 1

6406

GP3

88-1

63

GN

ipha

tes s

p 1

(n s

p)

6192

NIW

A 9

13 9

16G

P536

-91

GN

ipha

tes s

p 2

(n s

p)

6807

NIW

A 9

18G

P868

-27

1G

Nip

hate

s sp

3 (w

ith si

gmas

) 61

03ZM

A 1

6421

GP5

77-2

86

3G

HA

PLO

SCLE

RID

A P

ETR

OSI

IDA

EPe

tros

ia sp

1

6698

ZMA

164

08G

P781

-24

3G

Petr

osia

sp 2

(aff

aru

ensi

s Hen

tsch

el 1

912)

6689

ZMA

164

12G

P649

-63

GXe

stos

pong

ia e

xigu

a (K

irkpa

trick

190

0)

6016

BM

NH

199

56

223

6G

P273

-33

1G

Xest

ospo

ngia

sp 1

65

74ZM

A 1

6417

GP8

69-1

33

GXe

stos

pong

ia sp

2 (a

ff c

arbo

nari

a (L

amar

ck 1

814)

)59

98B

MN

H 1

995

622

22

ZM

A 1

6416

28G

P532

-421

3G

HA

PLO

SCLE

RID

A P

HLO

EOD

ICTY

IDA

EAk

a sp

2 (y

ello

w)

6024

GP4

07-3

35

GAk

a tr

achy

s de

Laub

enfe

ls 1

954

Hol

otyp

e U

SNM

231

461

16

GPh

loeo

dict

yida

e N

ew g

enus

sp 1

(n s

p)

GP8

24-7

0C

DN

098

5-F(

S)0C

DN

098

5-F

(S)

141

GS

Oce

anap

ia

sp 1

(cre

vice

stic

k sp

onge

) 67

02G

P261

B-2

21

GldquoP

ellin

ardquo p

ulvi

lla (T

hiel

e 1

903)

USN

M 2

3145

11

6G

HO

MO

SCLE

RO

PHO

RID

A P

LAK

INID

AE

Cor

ticiu

m sp

1 (n

sp

) 68

06N

IWA

914

17G

P868

-51

GPl

akin

a sp

1

6078

BM

NH

199

56

221

GP2

88-6

1G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsldquoL

ITH

ISTI

DA

rdquo A

ZOR

ICID

AE

Leio

derm

atiu

m sp

122

6S

ldquoLIT

HIS

TID

Ardquo

SC

LER

ITO

DER

MID

AE

Acic

ulite

s pap

illat

a Leacute

vi amp

Leacutev

i 19

830C

DN

101

5-N

(R)

0CD

N 1

015-

N (R

)1

RldquoL

ITH

ISTI

DA

rdquo T

HEO

NEL

LID

AE

Theo

nelli

dae

new

gen

us sp

1 (n

sp

) 0C

DN

098

9-J(

S)0C

DN

098

9-J (

S)12

1S

POEC

ILO

SCLE

RID

A C

RA

MB

IID

AE

Mon

anch

ora

clat

hrat

a (C

arte

r 18

83)

6809

QM

G31

7344

GP8

70-1

72

GPO

ECIL

OSC

LER

IDA

CR

ELLI

DA

EC

rella

cya

thop

hora

(Car

ter

1880

) 67

93Q

M G

3173

54

ZMA

164

18G

P253

-20

3G

POEC

ILO

SCLE

RID

A D

ESM

AC

ELLI

DA

EN

eofib

ular

ia h

artm

ani H

oope

r amp L

eacutevi

1993

NIW

A 9

22G

P838

-26

1G

POEC

ILO

SCLE

RID

A M

ICR

OC

ION

IDA

EC

lath

ria

(Cla

thri

a) b

asila

na L

evi

1961

6190

GP5

36-1

110

2G

Cla

thri

a (M

icro

cion

a) m

ima

(de

Laub

enfe

ls 1

954)

6710

QM

G31

7343

GP6

16-9

2G

Cla

thri

a (T

haly

sias

) cf

hirs

uta

Hoo

per amp

Lev

i 19

9359

80B

MN

H 1

995

622

34

GP8

69-3

410

2G

Cla

thri

a (T

haly

sias

) cf

rein

war

dti V

osm

aer

1880

6696

QM

G31

7353

GP5

77-3

57

2G

Cla

thri

a (T

haly

sias

) eur

ypa

(de

Laub

enfe

ls 1

954)

GP8

68-3

01

GC

lath

ria

(Tha

lysi

as) s

p 1

(n s

p J

NA

H re

f 24

31)

6810

QM

G31

7351

GP8

70-1

22

GC

lath

ria

(Tha

lysi

as) v

ulpi

na (L

amar

ck 1

814)

USN

M 2

3149

13

76

GC

lath

ria

(Wils

onel

la) s

p 2

(n s

p J

NA

H re

f 17

66)

6711

QM

G31

7365

GP7

46-1

22

GPO

ECIL

OSC

LER

IDA

MY

CA

LID

AE

Myc

ale

(Car

mia

) cec

ilia

de L

aube

nfel

s 19

540C

DN

096

4-H

0CD

N 0

964-

H (S

)1

SM

ycal

e (C

arm

ia) s

p 1

(n s

p J

NA

H re

f 21

99)

6796

QM

G31

7358

GP5

36-1

32

GU

losa

spon

gia

de L

aube

nfel

s 19

54

NIW

A 9

910

GP6

51-3

61

GPO

ECIL

OSC

LER

IDA

MY

XIL

LID

AE

Iotr

ocho

ta b

acul

ifera

Rid

ley

188

466

17B

MN

H 1

995

622

16

GP8

68-1

71

GIo

troc

hota

ditr

ocho

ta (d

e La

uben

fels

195

4)G

P288

-28

4G

POEC

ILO

SCLE

RID

A R

ASP

AIL

IID

AE

Cer

atop

sion

sp 1

(n s

p)

5938

BM

NH

199

56

224

2

QM

G31

7364

GP6

51-6

12

G

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

Mus

eum

R

efPh

oto

Not

esID

IsEc

hino

dict

yum

ant

rode

s (de

Lau

benf

els

1954

)U

SNM

231

481

86

GPO

ECIL

OSC

LER

IDA

TED

AN

IID

AE

Teda

nia

cf i

gnis

(Duc

hass

aing

amp M

iche

lotti

186

4)61

94Q

M G

3173

56G

P536

-10

2G

Teda

nia

mea

ndri

ca T

hiel

e 1

903

5985

BM

NH

199

56

223

5

ZMA

164

25G

P361

-29

3G

Teda

nia

sp 1

(n s

p)

6799

BM

NH

199

56

229

Q

M G

3173

47G

P866

-14

2G

SPIR

OPH

OR

IDA

TET

ILLI

DA

EC

inac

hyra

por

osa

(Len

denf

eld

188

8)U

SNM

231

38 (S

)1

96

SC

inac

hyra

sp(p

) 59

93B

MN

H 1

995

622

6G

P578

-27

91

GV

ERO

NG

IDA

APL

YSI

NEL

LID

AE

Aply

sine

lla rh

ax (d

e La

uben

fels

195

4)60

57B

MN

H 1

995

622

30

21G

P514

-12

201

GV

ERO

NG

IDA

PSE

UD

OC

ERA

TIN

IDA

EPs

eudo

cera

tina

tyro

eis (

de L

aube

nfel

s 19

54)

0CD

N 0

987-

H(S

) 0C

DN

101

1-J(

R)

0CD

N 0

987-

H (S

)0C

DN

101

1-J (

R)

1S

R

Pseu

doce

ratin

a pu

rpur

ea (C

arte

r 18

80)

669

4G

P578

-14

185

GPs

eudo

cera

tina

sp 1

(pap

er th

in y

ello

w)

0CD

N 0

981-

Y(S

) 0C

DN

102

3-V

(T)

0CD

N 0

981-

Y (S

)0C

DN

102

3-V

(T)

GP8

69-2

5(G

)

151

ST

G

Pseu

doce

ratin

a sp

2 (n

sp

) 67

08N

IWA

920

GP7

36-2

61

GV

ERO

NG

IDA

IA

NTH

ELLI

DA

E Ia

nthe

lla b

asta

(Pal

las

1766

)59

30B

MN

H 1

995

622

32

19G

P530

-25

1G

VER

TIC

ILLI

DA

VER

TIC

ILLI

TID

AE

Vace

letia

cry

pta

(Vac

elet

197

7)66

10G

P417

-33

(S)

5G

Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
  • GP768-25
  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

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App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

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eum

R

efPh

oto

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alic

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iclo

na)

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59

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622

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122

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74

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NH

199

56

221

4

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e)

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59

34B

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622

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stre

ble

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aube

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s 19

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iche

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App

endi

x 1

Spo

nges

of t

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aria

nas

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tinue

d)

Taxo

nU

GI

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eum

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622

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GP8

68-1

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GIo

troc

hota

ditr

ocho

ta (d

e La

uben

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195

4)G

P288

-28

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POEC

ILO

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RID

A R

ASP

AIL

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AE

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atop

sion

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199

56

224

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App

endi

x 1

Spo

nges

of t

he M

aria

nas

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tinue

d)

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nU

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efPh

oto

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els

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481

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AE

Teda

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gnis

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hass

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iche

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186

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3173

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nia

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hiel

e 1

903

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NH

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56

223

5

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164

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(n s

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6799

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NH

199

56

229

Q

M G

3173

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OPH

OR

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TET

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DA

EC

inac

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osa

(Len

denf

eld

188

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231

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)1

96

SC

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sp(p

) 59

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MN

H 1

995

622

6G

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e La

uben

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195

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H 1

995

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201

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de L

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s 19

54)

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101

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TIC

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letia

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pta

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elet

197

7)66

10G

P417

-33

(S)

5G

Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
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  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

App

endi

x 1

Spo

nges

of t

he M

aria

nas

(con

tinue

d)

Taxo

nU

GI

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eum

R

efPh

oto

Not

esID

IsH

alic

lona

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stel

la) s

p 2

(n s

p b

lue

aqua

rium

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iclo

na)

6043

59

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MN

H 1

995

622

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Hal

iclo

na li

gula

ta (W

hite

legg

e 1

901)

U

SNM

231

441

122

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iclo

na o

siri

s (de

Lau

benf

els

1954

)59

74

5929

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NH

199

56

221

4

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M 3

1763

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Hal

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1 (o

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e)

5972

59

34B

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622

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ge)

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alic

lona

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ble

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s 19

54H

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M 2

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11

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iche

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6192

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13 9

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6807

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) 61

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1

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aru

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s Hen

tsch

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912)

6689

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164

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190

0)

6016

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las

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19G

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VER

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VER

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7)66

10G

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Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

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iclo

na li

gula

ta w

as d

escr

ibed

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ther

n N

ew S

outh

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es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
  • GP768-25
  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

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Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

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f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

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gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

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ge P

sam

map

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lla p

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ter

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ow k

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n as

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erat

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ter

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he re

view

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ergq

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(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

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olle

ctio

n of

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drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

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rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

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t alrsquo

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83) r

ecor

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lmos

t cer

tain

ly th

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outh

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hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

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iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
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App

endi

x 1

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of t

he M

aria

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tinue

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1954

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186

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164

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199

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Ref

eren

ces

1)

de L

aube

nfel

s 19

54 2

) Sm

yth

1990

3)

Hoo

per

amp L

eacutevi 1

993

4)

Har

tman

amp G

orea

u 19

75 5

) H

artm

an amp

Gor

eau

1976

6)

Bry

an19

73 7

) Plu

cer-

Ros

ario

198

7 8

) Ruumlt

zler

amp M

usik

199

3 9

) A

vila

amp P

aul 1

997

10)

Bec

erro

et a

l 19

98 1

1) L

ee e

t al

2000

12)

Pau

l et a

l19

97 1

3) R

oger

s amp

Pau

l 199

1 1

4) D

uffy

amp P

aul 1

992

15)

Pen

ning

s et

al

1994

16)

Tha

cker

et a

l 19

98 1

7) L

ee e

t al

2001

18)

Plu

bruk

arn

et a

l 19

97 1

9) P

orde

sim

o amp

Sch

mitz

199

0 2

0) S

hin

et a

l 19

98 2

1) S

hin

et a

l 20

00 2

2) Q

uinn

amp K

ojis

199

9 2

3) C

arls

on amp

Hof

f 197

3 2

4)C

arls

on amp

Hof

f 197

4 2

5) D

avid

son

1993

26)

Rog

ers 1

989

27)

Sch

mitz

et a

l 19

85 2

8) S

chm

itz e

t al

1983

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

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iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
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  • GP532-9
  • GP270-36
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  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

Not

es

1) d

e La

uben

fels

(19

54)

reco

rds

that

wer

e no

t rec

olle

cted

nor

taxo

nom

ical

ly u

pdat

ed V

ouch

ers

of th

ese

reco

rds

wer

e de

posi

ted

at th

e U

SNM

as

note

d

2) S

myt

h (1

990)

reco

rds

not r

ecol

lect

ed a

nd ta

xono

my

not v

erifi

ed n

or u

pdat

ed S

myt

hrsquos

colle

ctio

ns w

ere

appa

rent

ly n

ot d

epos

ited

anyw

here

and

thes

e re

cord

s exi

st so

lely

as l

iste

d in

her

pap

er (K

Ruumlt

zler

per

s co

mm

200

0)3)

Thi

s spe

cies

can

gro

w in

eith

er a

larg

e e

xpos

ed o

r a sm

all

cryp

tic g

row

th fo

rm t

hese

two

grow

th fo

rms

hint

at c

rypt

ic d

iver

sity

Onl

y th

e la

tter

grow

th fo

rm w

as e

ncou

nter

ed o

n G

uam

The

cry

ptic

mor

ph h

as a

few

sm

all

surf

ace

man

ifest

atio

ns w

ith th

e ty

pica

l app

eara

nce

of th

e sp

ecie

sbu

t its

bul

k is

nes

tled

with

in th

e re

ef m

atrix

lar

gely

con

ceal

ed b

y tu

rf a

nd re

ef se

dim

ent

and

with

pie

ces o

f rub

ble

inco

rpor

ated

into

its m

ass

4) T

erpi

os a

ploo

s was

reco

llect

ed a

t the

type

loca

lity

but

has

not

yet

bee

n ta

xono

mic

ally

reex

amin

ed

5) A

s St

ylot

ella

agm

inat

a in

de

Laub

enfe

ls (1

954)

Sty

lote

lla a

uran

tium

is a

juni

or s

ynon

ym (s

ee a

bove

) M

isid

entif

ied

as H

ymen

iaci

don

aldi

s in

Schm

itz e

t al

(198

5)

6) C

ally

spon

gia

diffu

sa h

as n

ot b

een

seen

on

Gua

m in

rece

nt y

ears

but

de

Laub

enfe

lsrsquo (

1954

pla

te IV

b) il

lust

ratio

n an

d ha

bitu

s de

scrip

tion

of th

atsp

ecie

s fro

m G

uam

is st

rikin

gly

sim

ilar t

o N

ipha

tes

sp 3

a sp

ecie

s tha

t is c

omm

on a

t Dun

gas

Bea

ch w

here

de

Laub

enfe

lsrsquo r

ecor

d ca

me

from

D

emos

tratio

n of

the

pote

ntia

l ide

ntity

of t

hese

two

reco

rds a

wai

ts a

n ex

amin

atio

n of

de

Laub

enfe

lsrsquo s

peci

men

7)

The

rec

ord

of C

lath

ria

vulp

ina

is b

ased

on

de L

aube

nfel

srsquo (

1954

) sp

ecim

en o

f C

lath

ria

fron

dife

ra w

hich

was

ree

xam

ined

by

Hoo

per

amp L

eacutevi

(199

3) a

nd li

sted

und

er th

e sy

nony

my

of C

vul

pina

Whi

le C

vul

pina

was

not

enc

ount

ered

on

Gua

m d

urin

g re

cent

surv

eys

the

very

sim

ilar C

cf

rei

nwar

dti w

as a

nd th

at s

peci

es w

as a

lso

foun

d at

Dun

gas

Bea

ch f

rom

whe

re d

e La

uben

fels

rsquo sp

ecim

en o

f C

fro

ndife

ra o

rigin

ated

It i

sth

eref

ore

poss

ible

that

thes

e tw

o re

cord

s rep

rese

nt th

e sa

me

spec

ies

8) A

s K

iepl

itela

ant

rode

s in

de

Laub

enfe

ls (1

954)

Kie

plite

la d

e La

uben

fels

195

4 is

a ju

nior

syn

onym

of E

chin

odic

tyum

Rid

ley

amp D

unca

n 1

881

(Hoo

per 1

991

Hoo

per amp

Wie

denm

ayer

199

4) T

his s

peci

es w

as n

ot re

colle

cted

nor

taxo

nom

ical

ly re

exam

ined

9)

The

ide

ntity

of

the

one

or m

ore

(pro

babl

y tw

o) s

peci

es o

f C

inac

hyra

on

Gua

m h

as n

ot b

een

esta

blis

hed

lik

ely

one

corr

espo

nds

with

de

Laub

enfe

lsrsquo

(195

4) i

nter

pret

atio

n of

Cin

achy

ra p

oros

a (L

ende

nfel

d 1

888)

fro

m S

aipa

n a

spe

cies

tha

t w

as n

ot r

ecol

lect

ed t

here

nor

taxo

nom

ical

ly re

exam

ined

10

) Id

entif

ied

sam

ples

of

Cla

thri

a cf

hir

suta

hav

e a

nest

ling

grow

th f

orm

with

the

oute

r pa

rts o

f th

e sp

onge

are

flu

sh w

ith t

he c

oral

(us

ually

Pori

tes

rus)

into

whi

ch th

ey n

estle

One

spe

cim

en o

f C b

asila

na a

ppea

red

iden

tical

to C

cf

hirs

uta

in th

e fie

ld t

he o

nly

pote

ntia

l exa

mpl

eam

ong

spon

ges

on G

uam

whe

re f

ield

app

eara

nce

was

not

dia

gnos

tic (

alth

ough

fur

ther

atte

ntio

n m

ay p

rovi

de u

sefu

l fie

ld c

hara

cter

s) O

ther

exam

ples

of C

bas

ilana

had

an

expo

sed

mas

sive

gro

wth

form

on

buoy

s 11

) R

ecor

ded

as D

ysid

ea c

f re

ticul

ata

by C

arls

on amp

Hof

f (1

973

197

4)

who

des

crib

ed t

he c

epha

lasp

id o

pist

hobr

anch

s Sa

gam

inop

tero

nni

grop

unct

atum

and

S p

sych

edel

icum

from

this

hos

t12

) Th

e ne

w g

enus

is

the

sam

e as

tha

t re

pres

ente

d by

de

Laub

enfe

lsrsquo

(195

4) P

laki

nalo

pha

mir

abili

s Th

is s

pong

e is

a t

heon

ellid

lith

istid

with

unus

ual n

on-a

rticu

late

d de

sma

spic

ules

typi

cal o

f lith

istid

spon

ges

and

it ha

s unu

sual

irre

gula

r dis

cotri

aene

s w

ithou

t a rh

abd

(MK

unp

ublis

hed

data

) an

d is

clo

sely

rela

ted

to S

iliqu

ario

spon

gia

Hos

hino

13

) de

Laub

enfe

lsrsquo (

1954

) Ado

cia

viol

a w

as re

colle

cted

and

reex

amin

ed a

nd is

now

con

side

red

to b

elon

g to

Hal

iclo

na (R

enie

ra)

14) T

his s

peci

es h

as m

any

size

cat

egor

ies o

f tox

as G

uam

reco

rd b

ased

on

phot

o on

ly n

ot v

erifi

ed

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
  • GP768-25
  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25

15) T

his v

ery

unus

ual s

peci

es e

ncru

sts c

oral

line

spon

ges i

n ca

ves

and

has b

een

enco

unte

red

in S

aipa

n an

d w

est T

inia

n

16)

The

iden

tity

of th

e Te

thya

spe

cies

on

Gua

m h

ave

not b

een

esta

blis

hed

it is

like

ly th

at o

ne c

orre

spon

ds w

ith d

e La

uben

fels

rsquo (1

954)

Tet

hya

dipl

oder

ma

(now

syn

onym

ized

und

er T

ethy

a se

yche

llens

is

see

Ber

gqui

st amp

Kel

ly-B

orge

s 19

91)

from

Sai

pan

a s

peci

es t

hat

was

not

reco

llect

ed th

ere

nor t

axon

omic

ally

reex

amin

ed

17)

de L

aube

nfel

srsquo (

1954

) re

cord

of

Jasp

is t

uber

cula

ta (

a te

mpe

rate

Aus

tralia

n en

dem

ic)

from

Sai

pan

not

lis

ted

here

sep

arat

ely

has

not

bee

nre

exam

ined

it

mos

t lik

ely

corr

espo

nds

to R

habd

astr

ella

glo

bost

ella

ta (

see

Ken

nedy

200

0 fo

r a

revi

ew o

f th

is c

ompl

ex (

ofte

n re

cord

ed a

slsquoJ

aspi

s ste

llife

rarsquo)

and

pas

t con

fusi

on su

rrou

ndin

g it)

18

) The

com

mon

yel

low

-gre

en v

eron

gid

spon

ge P

sam

map

lysi

lla p

urpu

rea

Car

ter

1880

is n

ow k

now

n as

Pse

udoc

erat

ina

purp

urea

(Car

ter

1880

)af

ter t

he re

view

of B

ergq

uist

(198

0) w

hich

syno

nym

ised

the

form

er g

enus

with

the

late

r19

) Rec

olle

ctio

n of

Den

drill

a ni

gra

de L

aube

nfel

s 19

54 th

roug

hout

Mic

rone

sia

reve

aled

that

the

spec

ies

is a

dic

tyod

endr

illid

and

thus

it is

now

refe

rred

to a

s Dic

tyod

endr

illa

nigr

a (d

e La

uben

fels

195

4)

20)

Dys

idea

rha

x de

Lau

benf

els

1954

is

now

kno

wn

as A

plys

inel

la r

hax

(de

Laub

enfe

ls 1

954)

bec

ause

the

spon

ge e

xper

ienc

es a

n ae

roph

obic

disc

olor

atio

n to

wal

nut b

row

n up

on d

eath

and

exp

osur

e to

air

due

to th

e fle

shy

text

ure

of th

e m

esoh

yal

the

prom

inen

ce o

f ho

llow

det

ritus

-lin

ed g

olde

n fib

res

and

smal

l wel

l spa

ced

choa

nocy

te c

ham

bers

(Kel

ly u

npub

lishe

d)

21) S

chm

itz e

t alrsquo

s (19

83) r

ecor

d of

Am

phim

edon

is a

lmos

t cer

tain

ly th

is sp

ecie

s a

vou

cher

is a

vaila

ble

USN

M 3

1765

22

) Hal

iclo

na li

gula

ta w

as d

escr

ibed

from

sou

ther

n N

ew S

outh

Wal

es (W

hite

legg

e 19

01)

and

thus

is li

kely

diff

eren

t tha

n th

e sp

ecie

s so

iden

tifie

don

Gua

m

• Queensland Museum
• PO Box 3300
• South Brisbane Queensland 4101 Australia
• Marine Laboratory
• University of Guam
• Mangilao Guam 96923 USA
• • GP572-37
  • GP531-27
  • GP573-6
  • GP768-25
  • GP532-9
  • GP270-36
  • GP288-5
  • GP532-4
  • HOMOSCLEROPHORIDA PLAKINIDAE
  • GP288-28
  • GP530-25