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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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Phot
o p
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ble
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in th
e M
arin
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ity o
f Gua
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cop
ublic
atio
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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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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
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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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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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
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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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
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
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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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
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
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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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
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
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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nges
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imen
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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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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ITO
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Acic
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illat
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i 19
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DN
101
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RldquoL
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anch
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clat
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arte
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6809
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GPO
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CR
ELLI
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rella
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artm
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1993
NIW
A 9
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lath
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evi
1961
6190
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36-1
110
2G
Cla
thri
a (M
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a) m
ima
(de
Laub
enfe
ls 1
954)
6710
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7343
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16-9
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Cla
thri
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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
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) cf
rein
war
dti V
osm
aer
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6696
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7353
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77-3
57
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thri
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haly
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954)
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H re
f 24
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6810
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3149
13
76
GC
lath
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(Wils
onel
la) s
p 2
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p J
NA
H re
f 17
66)
6711
QM
G31
7365
GP7
46-1
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GPO
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CA
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Myc
ale
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mia
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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
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7358
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36-1
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GU
losa
spon
gia
de L
aube
nfel
s 19
54
NIW
A 9
910
GP6
51-3
61
GPO
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OSC
LER
IDA
MY
XIL
LID
AE
Iotr
ocho
ta b
acul
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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
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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
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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
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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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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
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
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
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talo
gue
num
bers
of v
ouch
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ouse
d in
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v o
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verte
brat
e co
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ions
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imen
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Prov
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Gua
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Ref
ref
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follo
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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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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
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
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Kelly et al Sponges of the Marianas 111
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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
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Springer V G 1982 Pacific plate biogeography with special reference toshorefishes Smithsonian Contributions to Zoology 367 iv + 182 pp
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Received 18 October 2001
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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
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
d in
Uni
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f Gua
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ions
all
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imen
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note
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nd o
f App
endi
xID
ide
ntifi
er 1
Mic
helle
Kel
ly 2
Joh
n H
oope
r 3
Rob
van
Soe
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nd W
allie
de
Wee
rdt
4 P
atric
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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
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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
ucet
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)
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N 0
980-
X (S
)1
5G
S
CLA
SS D
EMO
SPO
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AG
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SID
A A
GEL
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sp 1
60
820C
DN
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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
era
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
(con
tinue
d)
Taxo
nU
GI
Mus
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R
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oto
Not
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nare
s sol
lasi
Thi
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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
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rand
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llata
Thi
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190
0U
SNM
Ac
1837
33 (S
)1
16
SC
hond
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rtic
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Thie
le 1
900
6707
NIW
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15 Q
M G
3173
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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
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(pur
ple)
59
78B
MN
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995
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28
GP5
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0 1
GD
END
RO
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DEN
DR
ILLI
DA
ED
icty
oden
drill
a ni
gra
(de
Laub
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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
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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
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QM
GP8
62-2
84
GIr
cini
a sp
266
14G
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2G
Sarc
otra
gus s
p 1
(n s
p)
0CD
N 0
971-
O(R
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DN
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1-O
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Sarc
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p 2
(n s
p)
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N 1
014-
M(R
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DN
101
4-M
(R)
1R
DIC
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CER
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SPO
NG
IID
AE
Cos
cino
derm
a cf
mat
hew
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ende
nfel
d 1
886)
NIW
A 9
24 amp
925
GP7
63-1
31
GH
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spon
gia
met
achr
omia
de
Laub
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ls 1
954
6619
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21 0
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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
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
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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Uni
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note
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ide
ntifi
er 1
Mic
helle
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Joh
n H
oope
r 3
Rob
van
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allie
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cite
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G =
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m R
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T =
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ian
S =
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191
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191
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PM 9
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Y
PM 9
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N 0
983-
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RO
PHO
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OR
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Mel
ophl
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rasi
noru
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899
5943
BM
NH
199
56
225
11G
P524
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1G
Pena
res n
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e La
uben
fels
195
4)59
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995
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App
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Spo
nges
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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
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
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
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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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
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
ucet
ta sp
1 (n
sp
) 0C
DN
100
3-(R
)0C
DN
100
3-Y
(R)
1R
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
NG
IAE
AG
ELA
SID
A A
GEL
ASI
IDA
EAg
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
era
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
(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
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
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
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188
466
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H 1
995
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e La
uben
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195
4)G
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-28
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ILO
SCLE
RID
A R
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AIL
IID
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Cer
atop
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NH
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56
224
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Spo
nges
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aria
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Taxo
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Mus
eum
R
efPh
oto
Not
esID
IsEc
hino
dict
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ant
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s (de
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benf
els
1954
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481
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IID
AE
Teda
nia
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gnis
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hass
aing
amp M
iche
lotti
186
4)61
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M G
3173
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-10
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Teda
nia
mea
ndri
ca T
hiel
e 1
903
5985
BM
NH
199
56
223
5
ZMA
164
25G
P361
-29
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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
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inac
hyra
por
osa
(Len
denf
eld
188
8)U
SNM
231
38 (S
)1
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inac
hyra
sp(p
) 59
93B
MN
H 1
995
622
6G
P578
-27
91
GV
ERO
NG
IDA
APL
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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
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IDA
PSE
UD
OC
ERA
TIN
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eudo
cera
tina
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eis (
de L
aube
nfel
s 19
54)
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Pseu
doce
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arte
r 18
80)
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-14
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eudo
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tina
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er th
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ello
w)
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DN
102
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(T)
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ST
G
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GV
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IDA
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NTH
ELLI
DA
E Ia
nthe
lla b
asta
(Pal
las
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H 1
995
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32
19G
P530
-25
1G
VER
TIC
ILLI
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VER
TIC
ILLI
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Vace
letia
cry
pta
(Vac
elet
197
7)66
10G
P417
-33
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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
) 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
App
endi
x 1
Spo
nges
of t
he M
aria
nas
(con
tinue
d)
Taxo
nU
GI
Mus
eum
R
efPh
oto
Not
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IsD
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OC
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TID
A T
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REC
TID
AE
Aply
sino
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cf
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ans L
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nfel
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888
6706
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11
GP6
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Cac
ospo
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6140
BM
NH
199
56
222
39
10G
P530
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GFa
scap
lysi
nops
is re
ticul
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(Hen
tsch
el 1
912)
6609
NIW
A 9
19 0
CD
N 0
984-
C(S
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P407
-13
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DN
098
4-C
(S)
1G
S
Hyr
tios a
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(Pol
eacutejae
ff 1
884)
6004
BM
NH
199
56
222
4 G
P532
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4G
Hyr
tios e
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a (K
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r 18
89)
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1754
13
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264
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yrtio
s sp
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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
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A A
CA
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ells
i Har
tman
amp G
orea
u 1
975
6615
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otyp
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PM 9
077
422
(S)
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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
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IDA
HEM
IAST
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LID
AE
Hem
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arte
r 18
7167
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LID
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Sphe
cios
pong
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lobu
lari
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Sphe
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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
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
App
endi
x 1
Spo
nges
of t
he M
aria
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(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
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11
G
Axin
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(n s
p)
6163
60
15B
MN
H 1
995
622
40
Q
M G
3173
55G
P752
-12
G
Dra
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QM
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7357
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DN
095
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G S
Styl
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arte
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89)
5942
BM
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199
56
221
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A (S
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GS
HA
LIC
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RID
A D
ESM
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YID
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Myr
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rma
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sper
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0)
6138
BM
NH
199
75
133
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K (S
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elly
Bor
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Ber
gqui
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6042
BM
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199
56
221
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IID
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Axin
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ilson
192
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91Q
M G
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HA
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SCLE
RID
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E C
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M 2
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Cal
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ongi
a sp
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aren
s Pul
itzer
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1982
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A 1
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GC
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fib
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Rid
ley
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endy
188
7)66
88ZM
A 1
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GP5
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GC
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Rid
ley
188
466
95ZM
A 1
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ra D
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188
960
46
6045
B
MN
H 1
995
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19
ZM
A 1
6404
GP5
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G
HA
PLO
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RID
A C
HA
LIN
IDA
EAd
ocia
nee
ns (T
opse
nt 1
918)
USN
M A
c18
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11
6S
Hal
iclo
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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
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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
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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
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RID
A P
HLO
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EAk
a sp
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ello
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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(
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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
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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(
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098
9-J (
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MB
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AE
Mon
anch
ora
clat
hrat
a (C
arte
r 18
83)
6809
QM
G31
7344
GP8
70-1
72
GPO
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OSC
LER
IDA
CR
ELLI
DA
EC
rella
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hora
(Car
ter
1880
) 67
93Q
M G
3173
54
ZMA
164
18G
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RID
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DA
EN
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artm
ani H
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r amp L
eacutevi
1993
NIW
A 9
22G
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1G
POEC
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RID
A M
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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
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) cf
rein
war
dti V
osm
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1880
6696
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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
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p J
NA
H re
f 17
66)
6711
QM
G31
7365
GP7
46-1
22
GPO
ECIL
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LER
IDA
MY
CA
LID
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Myc
ale
(Car
mia
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ilia
de L
aube
nfel
s 19
540C
DN
096
4-H
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)1
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ycal
e (C
arm
ia) s
p 1
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p J
NA
H re
f 21
99)
6796
QM
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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
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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
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TED
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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
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(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
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
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
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
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
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