Marine Bivalvia (Mollusca) of Guam

Micronesica 35-36:218-243. 2003

Marine Bivalvia (Mollusca) of Guam

GUSTAV PAULAY1

Marine Laboratory University of Guam

Mangilao, Guam 96923 USA

AbstractThe marine bivalves of Guam are documented from newcollections and 339 species recorded. Nevertheless Guam’s faunaremains imperfectly known and is estimated to be considerably morediverse. Such high diversity on a local scale in the west-central Pacificsuggests that bivalve diversity in the Indo-west Pacific is much higherthan previously recorded. Pteriomorphs and heterodonts dominateGuam’s fauna, while protobranchs and anomalodesmatans comprise onlyfour species. Galeommatids represent the most species rich family, eventhough they are uncommon in samples.

IntroductionREGIONAL SETTING

Guam (13oN, 144oW), the southernmost of the Mariana Islands, is the largest,most populous, and developed island in Micronesia. As part of the island arcbordering the northwestern margin of the Pacific plate, Guam has had a complexgeological history since initiation around 42 Ma. Submarine and subaerialvolcanic episodes interspersed with periods of sedimentation and reef formation,together with alternating subsidence and uplift have led to complex physiographyboth above and below water. Northern Guam is a relatively flat, upliftedlimestone plateau, bordered by karstic cliffs and limited reef development. Incontrast southern Guam is dominated by volcanic hills, locally with limestoneveneers, with same valleys terminating in small bays. Barrier reefs are limited toshallow Cocos lagoon at the southern tip of the island, and Apra Harbor at thejunction of the volcanic south and karstic north (Fig. 1). Other than these twosmall lagoons, Guam’s shore is fronted by fringing reefs or lacks reef protection.

Guam has a diversity of marine habitats, even if it lacks the large andcomplex lagoons that characterize many Micronesian islands to the south. Majorhabitats include exposed limestone shores without reef protection, supratidalbenches, narrow, intertidal reef flats, fringing reefs with well developed (1-3 mdeep) moats, and shallow (<12 m) Cocos and deep (60 m) Apra Harbor lagoons.A gradient of southwardly increasing fringing reef development appears to bepartly the result of greater tectonic uplift toward the north. Reef development is 1Current address: Florida Museum of Natural History, Gainesville FL 32611-7800 USA, email:

[email protected]

Paulay: Bivalves of Guam 219

under further local geological control, with large embayments, created byfaulting and groundwater discharge in Tumon, Hagåtña (Agaña) and Pago bays,having developed the widest fringing reefs, and with a barrier reef (Apra Harbor)developed north of the projecting Orote peninsula. Additional variation in reefhabitats result from differences in wind, wave, and current exposure, with easternreefs considerably more exposed than western reefs to both tradewinds andtyphoons. Finally, terrigenous sediments are limited to the south, most prevalentalong the southwest coast, and this is reflected in the localization of severalinfaunal bivalve species. Fore reefs exhibit moderate variation around the islandin width, terracing, and in the abundance and nature of mobile sediments.Mangroves are limited to very small stands in stream estuaries, and to larger, butstill narrow bands in inner Apra Harbor and at the southeast end of the island inMerizo. Seagrass beds are developed along the larger fringing reef systems aswell as in Cocos lagoon.

Figure 1. Map of Guam.

Apra Harbor, the largest and busiest port in Micronesia, is the only deeplagoon in the Marianas. Its unique habitats host many species not found

220 Micronesica 35-36, 2003

elsewhere on Guam or in the Marianas. The harbor is also the main entry pointfor species introduced by shipping traffic (Paulay et al. 2002). Apra Harbor hasbeen substantially altered following World War II, as a large breakwater builtover the barrier reef and bank that previously bounded the lagoon along the northhas substantially reduced circulation with the surrounding ocean. Despite suchmajor alterations, Apra Harbor has a vibrant and thriving marine biota, with well-developed reefs, some of the highest coral cover (albeit dominated by Poritesrus) on Guam, and a diverse biota. Physical, chemical, and biotic conditions allshow pronounced east to west gradients in the harbor, related to theunidirectional exchange of water through the western entrance. Benthic habitatsrange from fairly exposed and oceanic reefs along the northwestern end of theOrote Peninsula, to silt-choked, lagoonal patch reefs, mud and silty sand bottoms,and mangroves in Sasa Bay and the Inner Harbor, with corresponding variationin the biota.

BIVALVESMollusks are among the taxonomically better known phyla, partly because

shells have long fascinated naturalists, are often sufficient in themselves forspecies level identification, and persist and accumulate after the animal’s death.Shells provide more abundant material for study, and a better cross section ofwhat lives in an area, than is available for most organisms, which can only bestudied alive. The study of Indo-West Pacific bivalves is nevertheless hamperedby the diversity of the fauna and paucity of revisionary studies. The publicationof several regional faunistic monographs (Oliver 1992, 1995, Lamprell &Whitehead 1992, Lamprell & Healy 1998, Okutani 2000) and taxonomicrevisions (e.g., Willan 1993, Vidal 1999, Glover & Taylor 2001), have greatlyfacilitated taxonomic work on Indo-West Pacific bivalves in the past decade.Nevertheless this diverse class remains poorly known. Although many, especiallysmaller bivalves remain undescribed, a perhaps greater limitation for faunisticstudies is the profusion of inadequately described taxa and consequent abundanceof junior synonyms that have resulted from the ready availability and attractionof shells. Revisionary studies are badly needed for most families to stabilize bothspecies level nomenclature and generic and higher classification.

The bivalves of Guam have received little attention in the past. Species fromthe island have been included in some revisions (e.g., Stenzel 1971, Rosewater1961, 1965, Waller 1972, Willan 1993, Vidal 1993, 1997, 1999), and one specieswas described from Guam (Raines 1996). Several M.S. theses at the Universityof Guam focused on bivalves (Jameson 1975, Day 1977, Braley 1981, Persselin1998, Peshut 2000). The mollusks of the Northern Mariana Islands, but notGuam, were checklisted by Vermeij et al. (1983), while Cloud (1959) recorded48 species from Saipan. Checklist attempts for Guam began with Smith (1986),who listed 65 species, while Paulay (1996a) recorded 201 species, and Peshut(2000) encountered 223 taxa in a quantitative survey. I have been incidentallycollecting and identifying bivalves on Guam since 1991, but attention paid to the


comprehensive marine biodiversity survey featured in this volume has limitedeffort spent specifically on bivalves. The present checklist is thus more a reportof progress to date than an authoritative list of the fauna. Nevertheless it alreadydemonstrates the substantial local diversity of the group. Other than the 20sediment samples (500+ specimens each) picked by Peshut (2000) and ca. 20dredge hauls from 50-150 m depths fully sorted for bivalves, bivalve collectinghas been incidental to other work. Taxonomic determinations remainpreliminary, with many species identified to generic level only. Accurate specieslevel identifications in bivalves often require mini revisions of relevant taxa.

MethodsBivalves were collected from beaches, reef flats, moats, lagoons, fore reefs,

and the lower reef talus to ca. 200m by walking, snorkeling, SCUBA, anddredging. Dredging was largely confined to 50-150m depths, focused on a well-developed terrace at that depth around the island. Epifaunal and endolithicspecies were collected from reef substrata, while dead shells and infaunalbivalves were taken by fanning sand during dives and from sediment samplesretained on 0.8mm mesh sieves (see Peshut 2000). I have examined ~15,000specimens, although over 80% of these came from fewer than 40 sediment anddredge samples.

The extent of coverage varies with macrohabitat as well as microhabitat.Coverage is relatively good on the leeward deep slope and on fringing reefs. It ismoderate on the fore reef at diving depths, in Apra Harbor, and on the windwarddeep slope. Endolithic taxa have received little attention, as have cemented,epibenthic species. Unidentified taxa are given consecutive numbers in my notes,these are used here as they apply to Guamanian taxa. Literature records areincluded where available. Records from other Mariana Islands were notsystematically searched, although are included when they are based on specimensseen or from reliable literature.

Results & DiscussionAt present 339 species of bivalves are known from Guam (Appendix 1).

Because considerable effort was placed on picking shells from sediment retainedon a 0.8mm mesh, microbivalves are at least as well documented as largerspecies. Nevertheless new records are still commonly encountered: almost everysediment sample processed included species previously unrecorded from Guam.Given that sampling was virtually confined to <200m and remains modest there,the entire bivalve fauna of the island is likely considerably greater than 500species.

Such richness underscores Bouchet et al.’s (2002) observation that the Indo-West Pacific mollusk fauna is considerably more diverse than previouslyrecognized. The highest currently documented regional bivalve diversity in the


world is around 1200 species (Crame 2000), in the Southeast Asian – Indo-Malayan region. Species richness rapidly decreases eastward from this diversitycenter in most marine organisms (Paulay 1997). In comparison to these regionaldiversities, the local species richness of bivalves on Guam is remarkable,considering the location and small size of the island, limited habitat diversity, andlimited extent of coverage, especially with regard to depth. Bouchet et al. (2002)documented very high mollusk diversity in a 5 X 5 km area in Koumac, NewCaledonia, with 519 species of bivalves recorded. Their results were based on asample size (N = 45,480) ca. three times greater than in the present study, froman island with substantially greater habitat diversity than Guam. An interestingcomparison can be made between the opisthobranch fauna of Koumac (271species) and Guam (485 species; Carlson & Hoff 2003). Opisthobranchs havebeen more exhaustively sampled on Guam than any other mollusk group. As agroup they are also more difficult to sample in rapid biodiversity surveys thanshelled taxa like bivalves, because most species can only be collected alive.These comparisons suggest that Guam’s bivalve biota remains substantiallyundersampled. It further implies that the species richness of bivalves in the stillpoorly-explored Indo-Malayan diversity center is likely much higher thanheretofore documented. Preliminary samples from the area indeed suggest veryhigh species richness, especially when microbivalves are considered.

Pteriomorphs and heterodonts dominate Guam’s fauna (Table 1), whileprotobranchs and anomalodesmatans are represented by just 4 species.Dominance of these two subclasses is typical for shallow-water, tropical Pacificisland bivalve faunas (Paulay 1990). 46 families were encountered, 9 with >10species, while 23 families were represented by 1 or 2 species each (Table 2). Incomparison, Bouchet et al. (2002) encountered 62 families at Koumac, NewCaledonia, 14 with >10 species, and 27 with 1 or 2 species. The rankings offamilies in terms of species diversity are similar between Guam and NewCaledonia (Tables 2). The main overall difference is that infaunal taxa areproportionately more diverse than epifaunal taxa in New Caledonia relative toGuam. Thus the ratio between the predominantly infaunal heterodonts andpredominantly epifaunal pteriomorphs is 1.99 for New Caledonia and 1.56 forGuam. This difference reflects the different nature of reef habitats at theselocations, with a very wide, soft-bottom-dominated lagoon enclosed by thebarrier reef at Koumac compared with limited fringing reef development givingway to a hard-bottom-dominated fore reef environment on Guam.

The Galeommatidae sensu lato (cf. Ponder 1998) stand out in both samplesas a megadiverse group. Their actual diversity must be substantially higher thanindicated in Table 3, and could be several times higher than that of any otherfamily. Galeommatid shells are relatively uncommon in samples, and manyspecies are currently known from one or a few valves on Guam. Similarly, atKoumac galeommatids accounted for only 1.6% of the shells, but 12% of thespecies diversity (Bouchet et al. 2002). Their rarity is likely in part apreservational artifact, as shells are often exceptionally fragile, and in part an


ecological / collecting issue, as many species live in specialized microhabitats,often in commensal association with varied invertebrate hosts. Such acombination of rarity and diversity implies that the known diversity of this groupwill grow rapidly and substantially when larger samples become available.Galeommatids are poorly known taxonomically, with generic and even highertaxonomic limits poorly understood, and much if not most of their indicateddiversity may be in undescribed species (Morton & Scott 1989).

Table 1. Species richness of bivalve subclasses on Guam and New Caledonia

Clade Guam Koumac, NC

Protobranchia 1 2Pteriomorphia 131 165Heterodonta 204 329Anomalodesmata 3 21

Data for Koumac, New Caledonia from Bouchet et al. (2002)

At least 10 of the bivalve species documented appear no longer to survive onGuam. Locally extirpated taxa include 3 intentionally (Crassostrea gigas,Saccostrea tuberculata, Tridacna gigas) and 3 accidentally introduced species(Anomia nobilis, Chama fibula, Chama macerophylla), 3 indigenous speciesfrom Apra Harbor (Lopha cristagalli, Timoclea sp. 9, Spondylus varius) and thepreviously widespread Hippopus hippopus. The last two species were probablycollected to death, S. varius for curios following WWII, while Hippopusprehistorically, presumably for food. The other two indigenous species may havesuccumbed to environmental change in Apra Harbor.

Table 2. Most species rich bivalve families on Guam and New Caledonia

Family Guam Guam Koumac, NC Koumac, NC# of species % of fauna # of species % of fauna

Galeommatidae s. l. 39 11% 61 12%Tellinidae 38 11% 51 10%Cardiidae1 29 9% 37 7%Veneridae 28 8% 53 10%Limidae 21 6% 15 3%Pectinidae 21 6% 27 5%Arcidae 19 6% 24 5%Mytilidae 18 5% 27 5%Lucinidae 17 4% 20 4%

Species richness of families with >10 species on Guam compared with Koumac, New Caledonia(Bouchet et al. 2002). The other families in the New Caledonian sample that had >10 speciesare: Semelidae (19), Spondylidae (16), Psammobiidae (13), Pteriidae (12), and Mactridae (11).

1includes Tridacnidae of Bouchet et al. 2002.


AcknowledgementsI thank the students, faculty, and staff of the University of Guam Marine Lab

for help with documenting the bivalves of Guam. Special thanks to HarryConley, Vince Diego, Lisa Kirkendale, Ismael Perez, Sara Persselin, JohnStarmer, Linda Ward and especially Pete Peshut for collecting bivalves, BarrySmith for setting the stage, Frank Cushing for dredging, and Rob Myers for useof his photos. Thanks also to Joe Carter, Aki Matsukuma, Paul Scott, JohnTaylor, Jacques Vidal, and Tom Waller, for help with identifications. Funding byNOAA (Insular Pacific Marine Research Program), U.S. Dept. of Defense atCOMNAVMARIANAS, and SeaGrant (SG-NIS-35) are gratefully acknow-ledged. Contribution 492 of the University of Guam Marine Laboratory.

ReferencesAmesbury, J. R. 1999. Changes in species composition of archaeological marine

shell assemblages on Guam: human preference or environmental change?Micronesica 31: 347-366.

Bouchet, P., P. Lozouet, P. Maestrati & V. Heros. 2002. Assessing the magnitudeof species richness in tropical marine environments: exceptionally highnumbers of molluscs at a New Caledonia site. Biological Journal of theLinnean Society 75: 421-436.

Braley, R. D. 1981. Reproductive periodicity in the indigenous oyster Saccostreacucullata in Sasa Bay, Apra Harbor, Guam, with reference to cultivation.M.S. Thesis in Biology, University of Guam, 44 pp.

Braley, R. D. 1982. Reproductive periodicity in the indigenous oyster Saccostreacucullata in Sasa Bay, Apra Harbor, Guam. Marine Biology 69: 165-173.

Braley, R. D. 1984. Mariculture potential of introduced oysters Saccostreacucullata tuberculata and Crassostrea echinata in Guam and a histologicalstudy of reproduction in C. echinata. Australian Journal of Marine andFreshwater Research 185: 129-141.

Carlson, C. & P. J. Hoff. 2003. The opisthobranchs of the Mariana Islands.Micronesica 35-36: 271-293.

Cloud, P. E. Jr. 1959. Geology of Saipan Mariana Islands Part 4. Submarinetopography and shoal-water ecology. Geological Survey Professional Paper280-K.

Crame, J. A. 2000. Evolution of taxonomic diversity gradients in the marinerealm: evidence from the composition of Recent bivalve faunas.Paleobiology 26: 188-214.

Day, J. E. 1977. Growth rates of, and predation on, different size classes ofQuidnipagus palatum Iredale (Bivalvia) on two reef-flats of Guam. M.S.Thesis in Biology, University of Guam, 51 pp.


Dijkstra, H. H. & B. K. Raines. 1999. Pascahinnites n. gen. for Pecten (Chalmys)[sic] pasca Dall, 1908, a cemented Easter Island scallop (Bivalvia:Pectinidae). Basteria 63: 199-203.

Edmondson, C. H. 1946. Dispersal of shipworms in the Pacific. Nautilus 60: 53-54.

Eldredge, L. G. 1994. Perspectives in aquatic exotic species management in thePacific islands. Volume 1. Introductions of commercially significant aquaticorganisms to the Pacific islands. South Pacific Commission Inshore FisheriesResearch Project Technical Document 7: 1-127.

Glover, E. A. & J. D. Taylor. 2001. Systematic revision of Australian and Indo-Pacific Lucinidae (Mollusca: Bivalvia): Pillucina, Wallucina anddescriptions of two new genera and four new species. Records of theAustralian Museum 53: 263-292.

Hayami, I. & T. Kase. 1993. Submarine cave Bivalvia from the Ryukyu Islands:systematics and evolutionary significance. The University Museum, TheUniversity of Tokyo, Bulletin. 35:1-133

Hayami, I. & T. Kase. 1996. Characteristics of submarine cave bivalves in theNorthwest Pacific. American Malacological Bulletin 12: 59-65.

Hidalgo, J. G. 1904-1905. Catalogo de los moluscos testáceos de las IslasFilipinas, Joló y Marianas. I. Moluscos Marinos. Gaceta de Madrid, Madrid.

Hutsell, K. C. 1999. Semipallium marybellae (Raines, 1996) new rangeextensions. Festivus 31(4):48-49.

Jameson, S. C. 1975. Early life history of the giant clams Tridacna croceaLamarck, Tridacna maxima Röding, and Hippopus hippopus Linnaeus. M.S.Thesis in Biology, University of Guam, 47 pp.

Kay, E. A. 1979. Hawaiian marine shells. Reef and shore fauna of Hawaii.Section 4: Mollusca. Bishop Museum Press, Honolulu.

Kesling, R. V. 1958. Fossil crabs from Guam. Journal of Paleontology 14: 207-263.

Kilburn, R. N. & E. Rippey. 1982. Sea shells of southern Africa. MacMillan,Johannesburg.

Lamprell, K. & J. Healy. 1998. Bivalves of Australia. Volume 2. Backhuys ,Leiden.

Lamprell, K. & T. Whitehead. 1992. Bivalves of Australia. Volume 1. CrawfordHouse Press, Bathurst, NSW.

Morton, B. & P. H. Scott. 1989. The Hong Kong Galeommatacea (Mollusca:Bivalvia) and their hosts, with descriptions of new species. Asian MarineBiology 6: 129-160.

Nicol, D. 1950. Recent species of the lucinoid pelecypod Fimbria. Journal of theWashington Academy of Sciences 40: 82-87.

Okutani, T., ed. 2000. Marine mollusks in Japan.). Tokai University Press,Tokyo.

Oliver, P. G. 1992. Bivalved seashells of the Red Sea. Wiesbaden, Germany,Verlag Christa Hemmen.


Oliver, P. G. 1995. Bivalves (Bivalvia). In: Seashells of Eastern Arabia. D. T.Bosch, S. P. Dance, R. G. Moolenbeek & P. G. Oliver. pp. 194-281.Motivate Publishing, Dubai.

Paulay, G. 1987. Biology of Cook Islands' bivalves, Part I. Heterodont families.Atoll Research Bulletin 298: 1-31.

Paulay, G. 1989. Marine invertebrates of the Pitcairn Islands: species composi-tion and biogeography of corals, molluscs, and echinoderms. Atoll ResearchBulletin 326: 1-28.

Paulay, G. 1990. Effects of Late Cenozoic sea-level fluctuations on the bivalvefaunas of tropical oceanic islands. Paleobiology 16: 415-434.

Paulay, G. 1996a. Working list of the bivalves of Guam. In: Baseline biodiversityassessment of natural harbors at Guam and Hawaii. L. G. Eldredge & G.Paulay. pp. 17-22. Report for Insular Pacific Regional Marine ResearchProgram, SEAGRANT, University of Hawaii.

Paulay, G. 1996b. Dynamic clams: historical changes in the bivalve fauna ofPacific islands as a result of sea level fluctuations. American MalacologicalBulletin 12: 45-57.

Paulay, G. 1997. Diversity and distribution of reef organisms. In: C. E. Birkeland(ed). Life and death of coral reefs. pp. 298-353. Chapman & Hall, New York.

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

Persselin, S. L. 1998. The evolution of shell windows within the Fraginae(Bivalvia: Cardiidae) and the origin of algal symbiosis in cardiids. M.S.Thesis in Biology, University of Guam, 49 pp.

Peshut, P. J. 2000. Diversity of the tropical marine benthos: Bivalves of coralreef and near-reef habitats. M.S. Thesis in Biology, University of Guam, 49pp.

Ponder, W. F. 1998. Superfamily Galeommatoidea. In: P. L. Beesley, G. J. B.Ross & A. Wells (eds), Mollusca: the Southern Synthesis. Fauna ofAustralia. Vol. 5. Part A. pp. 316-318 . CSIRO Publishing, Melbourne.

Raines, B. K. 1996. Description of Semipallium marybellae n. sp. (Bivalvia:Pectinidae) from Guam, with notes and bivariate statistical comparison toSemipallium tigris (Lamarck, 1819). Conchiglia 28: 20-30.

Roch, F. 1976. Die terediniden polynesiens. Bolletino Museo Civico di StoriaNaturale di Venezia 28: 33-55, 1 fig.

Rosewater, J. 1961. The family Pinnidae in the Indo-Pacific. Indo-PacificMollusca 1: 175-226.

Rosewater, J. 1965. The family Tridacnidae in the Indo-Pacific. Indo-PacificMollusca 1: 347-396.

Smith, B. D. 1986. Working list of the bivalves of Guam. University of GuamMarine Laboratory, unpublished manuscript.

Solem, A. 1954. Living species of the pelecypod family Trapeziidae. Proceedingsof the Malacological Society of London. 31: 64-84, pls. 6-7.


Stenzel, H. B.1972. Oysters. In R. C. Moore & C. Teichert (eds), Treatise oninvertebrate paleontology. Mollusca 6. Bivalvia Volume 3. University ofKansas, Lawrence KS.

Vermeij, G. J., E. A. Kay & L. G. Eldredge. 1983. Mollusks of the northernMariana Islands, with special references to the selectivity of oceanicdispersal barrier. Micronesica 19: 27-55.

Vidal, J. 1993. Variability of Acrosterigma elongatum, a polytypic species(Mollusca, Cardiidae). Journal of the Malacological Society of Australia 14:41-58.

Vidal, J. 1997. Large Trachycardiinae from the Indo-West Pacific: The group ofVasticardium orbita (Broderip & Sowerby, 1833)(Mollusca, Cardiidae).Molluscan Research 18: 11-32.

Vidal, J. 1999. Taxonomic review of the elongated cockles: generaTrachycardium, Vasticardium and Acrosterigma (Mollusca, Cardiidae).Zoosystema 21: 259-335.

Waller, T. R. 1972. The Pectinidae (Mollusca: Bivalvia) of Eniwetok Atoll,Marshall Islands. The Veliger 14: 221-264.

Willan, R. C. 1993. Taxonomic revision of the family Psammobiidae (Bivalvia:Tellinoidea) in the Australian and New Zealand region. Records of theAustralian Museum, Supplement 18: 1-132.

Received 1 October 2001

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cula

ris (

Linn

aeus

, 175

8)U

F 28

7437

, UG

I 191

622

1Se

ptife

r cf.

cum

ingi

i Réc

luz,

184

9U

F 28

7330

, UG

I 465

9(P)

P1

P

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsSe

ptife

r exc

isus

(Wie

gman

n, 1

837)

UF

2971

92, U

GI 6

255U

SNM

487

060,

U

SNM

617

675

(P),

USN

M 6

2011

6 (R

)P

GP7

85-7

1P,

R

Sept

ifer c

f. fu

rcill

ata

Gou

ld, 1

861

UF

2879

64, U

GI 6

552

P2

PTER

IOM

OR

PHIA

: AR

CID

AE

Acar

plic

ata

(Dill

wyn

, 181

7)U

F 28

6758

P

1Ac

ar sp

. 2 (c

onge

nita

of H

abe,

not

Sm

ith, 1

885)

U

F 28

4805

P

1An

adar

a an

tiqua

ta (L

inna

eus,

1758

)U

F 28

7431

, UG

I 625

92

11

Arca

ave

llana

Lam

arck

, 181

9U

F 28

7376

22

561

Arca

ven

tric

osa

Lam

arck

, 181

9U

F 29

6972

, UF

2969

44(M

), U

GI 9

97P

1M

Arca

sp. 2

(cav

e)

UF

2969

49

P2

Arca

sp. 3

(aff

. acu

min

ata)

U

F 28

7843

, UG

I 652

9P

2Ba

rbat

ia a

myg

dalu

mto

stum

(Röd

ing,

179

8)U

F 28

7457

, UG

I 626

1P

1Ba

rbat

ia fo

liata

(For

sskå

l, 17

75)

UF

2847

94, U

GI 6

375

P1

Barb

atia

lim

a (R

eeve

, 184

4)U

F 28

4967

, UG

I 653

7P

1Ba

rbat

ia p

arva

(Sow

erby

, 183

3)U

F 29

6967

, UG

I 626

3P

1Ba

rbat

ia te

nella

(Ree

ve, 1

844)

UF

2849

20, U

GI 6

271

P1

Barb

atia

s. l.

sp. 1

0 (A

rcop

sis-

like)

U

F 28

4837

, UG

I 634

4P

22

Barb

atia

sp. 1

1 U

F 28

7824

P

22

Bath

yarc

a sp

. 1

UF

2969

45

P2

2Be

ntha

rca

sp. 1

(cav

e)

UF

2969

46

32,

31

Bent

harc

a sp

. 2 (a

ff. x

enop

hori

cola

(Kur

oda,

193

0))

UF

2969

47, U

GI 6

290

P2

2Be

ntha

rca

sp. 3

(dee

p ca

ve)

UF

2969

84

3?2,

42

Haw

aiar

ca re

ctan

gula

Dal

l, B

arts

ch &

Reh

der,

1938

UF

2873

25, U

GI 6

269

P1,

2PT

ERIO

MO

RPH

IA: N

OET

IID

AE

Arco

psis

orn

ata

(Via

der,

1951

)U

F 29

6976

, UG

I 652

6P

1,2

Arco

psis

sp. 1

U

F 28

7824

P

2Ar

cops

is sp

. 2

UF

2875

63

P2

Arco

psis

sp. 3

U

F 28

7557

P

2St

riar

ca sp

. 1

UF

2969

77, U

SNM

594

151,

UG

I 628

2P

1PT

ERIO

MO

RPH

IA: G

LYC

YM

ERID

IDA

EG

lycy

mer

is re

evei

(May

er, 1

868)

UF

2877

46, U

GI 5

358

P1,

2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsTu

ceto

na p

ectu

ncul

us (L

inna

eus,

1758

)U

F 28

7635

, UG

I 653

9P

2PT

ERIO

MO

RPH

IA: P

HIL

OB

RY

IDA

EC

ratis

cf.

kane

koi H

ayam

i & K

ase,

199

3U

F 29

6966

, UG

I 630

83

1,2

PTER

IOM

OR

PHIA

: AR

CO

IDA

INC

ERTA

E SE

DIS

Lim

opso

id sp

. 1

UF

2969

51

35

1,2

PTER

IOM

OR

PHIA

: PIN

NID

AE

Pinn

a m

uric

ata

Linn

aeus

, 175

8U

F 29

8503

, UG

I 620

84

GP6

54-1

1St

rept

opin

na sa

ccat

a (L

inna

eus,

1758

)U

F 28

4812

, UG

I 621

2P

GP5

81-1

11

PTER

IOM

OR

PHIA

: PTE

RII

DA

EEl

ectr

oma

sp. 1

U

F 28

4809

P

1Pi

ncta

da a

nom

ioid

es?

(Ree

ve, 1

857)

UF

2847

92, U

GI 1

923

P1

Pinc

tada

mac

ulat

a (G

ould

, 185

0)U

F 28

9027

, UG

I 226

4P

1Pi

ncta

da m

arga

ritif

era

(Lin

naeu

s, 17

58)

UF

2407

8, U

GI 1

342

PG

P762

-21

Pter

ia c

hine

nsis

(Lea

ch, 1

814)

UF

2849

72

P1

Pter

ia c

f. ho

wen

sis L

ampr

ell &

Hea

ly, 1

997

UF

2835

34, U

F 28

7515

(M),

UF

2824

09(S

) P

1,2

M,S

PTER

IOM

OR

PHIA

: ISO

GN

OM

ON

IDA

EIs

ogno

mon

eph

ippi

um (L

inna

eus,

1758

)U

F 28

8860

, UG

I 655

9P

1Is

ogno

mon

isog

nom

um (L

inna

eus,

1758

)U

F 28

7324

5

1Is

ogno

mon

legu

men

? (G

mel

in, 1

791)

UF

2824

15

22?

531

Isog

nom

on n

ucle

us (L

amar

ck, 1

819)

P6

1Is

ogno

mon

per

na (L

inna

eus,

1767

)U

F 29

2278

, UG

I 169

5P

GP7

85-6

1PT

ERIO

MO

RPH

IA: M

ALL

EID

AE

Mal

leus

dec

urta

tus L

amar

ck, 1

819

UF

2847

90, U

GI 6

215

PG

P525

-31

Mal

leus

cf.

nutta

lli (C

onra

d, 1

837)

UF

2847

89, U

GI 6

311

P7

1PT

ERIO

MO

RPH

IA: V

ULS

ELLI

DA

EVu

lsel

la sp

. 1

UF

2848

57, U

GI 5

449

P1

PTER

IOM

OR

PHIA

: EN

TOLI

IDA

EPe

ctin

ella

aeq

uori

s Dijk

stra

, 199

1U

F 28

7556

, UG

I 631

2H

2PT

ERIO

MO

RPH

IA: P

RO

PEA

MU

SSII

DA

EC

hlam

ydel

la sp

. 1 (a

ff. i

ncub

ata

Hay

ami &

Kas

e, 1

993)

U

F 29

7253

, UG

I 629

73

1,2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsPa

rvam

ussi

um c

rypt

icum

Hay

ami &

Kas

e, 1

993

UF

2972

55

P2

Parv

amus

sium

sp. 1

U

F 28

7583

, UG

I 634

9W

2PT

ERIO

MO

RPH

IA: P

ECTI

NID

AE

Angu

ipec

ten

pict

urat

us D

ijkst

ra, 1

998

UF

2889

30, U

GI 5

880

W2

“Chl

amys

” pe

rfec

ta (M

elvi

ll, 1

909)

UF

2985

35, U

GI 5

878

W2

1,2

Cry

ptop

ecte

n nu

x nu

x (R

eeve

, 185

3)U

F 28

6762

H

2D

ecat

opec

ten

radu

la (L

inné

, 175

8)U

SNM

763

576

W1

Exce

llich

lam

ys sp

ecta

bilis

(Ree

ve, 1

853)

UF

2823

99, U

GI 1

449(

S)W

1S

Glo

rich

lam

ys e

lega

ntis

sim

a (D

esha

yes,

1863

)U

F 29

8518

, UG

I 588

5W

1,2

Glo

ripa

llium

pal

lium

(Lin

naeu

s, 17

58)

USN

M 6

0429

3 6,

2254

1G

lori

palli

um sp

ecio

sum

(Sow

erby

, 184

2)U

F 29

8529

, UG

I 589

5W

2H

emip

ecte

n fo

rbes

ianu

s Ada

ms &

Ree

ve, 1

850

UF

2987

89

W1

Juxt

amus

sium

cou

dein

i (B

avay

, 190

2)U

F 29

8520

W1

Juxt

amus

sium

sp. 1

U

F 29

8538

, UG

I 588

6W

2La

evic

hlam

ys ir

regu

lari

s (So

wer

by, 1

842)

UF

2985

25, U

SNM

763

919,

UG

I 123

5W

1La

evic

hlam

ys le

mni

scat

a (R

eeve

, 185

3)U

F 29

8528

, UG

I 226

5, U

GI 5

25 (G

u)W

1G

uLa

evic

hlam

ys sq

uam

osa

(Gm

elin

, 179

1)U

F 29

8521

, USN

M 6

0429

7, U

GI 5

377

61

Mim

achl

amys

sp. 1

(aff

. kau

aien

sis (

Dal

l, B

arts

ch &

Reh

der,

1938

))

UF

2985

48

W2

Mir

apec

ten

mir

ificu

s (R

eeve

, 185

3)U

F 29

8522

, UG

I 545

1, U

GI 5

882

W2

Mir

apec

ten

spic

eri (

Reh

der,

1944

)U

F 28

2407

, UG

I 447

4W

GP7

81-3

21,

2Pa

scah

inni

tes c

orus

cans

cor

usca

ns (H

inds

, 184

5)U

F 28

4891

, USN

M 6

2034

2 6

391

Pedu

m sp

ondy

loid

eum

(Gm

elin

, 179

1)

61

Sem

ipal

lium

dia

nae

(Cra

ndal

l, 19

79)

UF

2985

24, U

GI 5

885

W2

Sem

ipal

lium

mar

ybel

lae

Rai

nes,

1996

SBM

NH

143

017

(hol

otyp

e), U

F 28

7522

,U

GI 5

363

7,8

581

PTER

IOM

OR

PHIA

: SPO

ND

YLI

DA

ESp

ondy

lus i

mbu

tus R

eeve

, 185

6U

F 28

7863

P

2Sp

ondy

lus l

ingu

afel

is S

ower

by, 1

847

UF

2875

28

P2

Spon

dylu

s mul

timur

icat

us R

eeve

, 185

6U

GI 5

379

P1

Spon

dylu

s nic

obar

icus

Sch

reib

ers,

1793

UF

2921

24, U

GI 1

723

P1

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsSp

ondy

lus o

ccid

ens?

Sow

erby

, 190

3U

F 28

7979

P

2Sp

ondy

lus s

quam

osus

Sch

reib

ers,

1793

UF

2849

70, U

GI 5

378

P1

Spon

dylu

s var

ius S

ower

by, 1

827

P

81

Spon

dylu

s vio

lasc

ens L

amar

ck, 1

819

UF

2971

95

PG

P831

-71

Spon

dylu

s sp.

4 (o

rang

e, d

eep)

U

F 28

6474

P

2PT

ERIO

MO

RPH

IA: A

NO

MII

DA

EAn

omia

nob

ilis R

eeve

185

6U

F 29

6979

5

91

Ano

miid

ae sp

p.

UF

2824

22, U

GI 6

557

P10

1,2

PTER

IOM

OR

PHIA

: PLI

CA

TULI

DA

EPl

icat

ula

sp. 1

U

F 28

7918

, UG

I 625

8P

1PT

ERIO

MO

RPH

IA: D

IMY

IDA

ED

imye

lla sp

. 1 (n

. sp.

cav

e)

UF

2888

44

32,

11,

121

Dim

yella

sp. 2

(n. s

p. d

eep)

U

F 28

8013

P

2,11

2PT

ERIO

MO

RPH

IA: L

IMID

AE

Cte

noid

es a

les (

Finl

ay, 1

927)

UF

2985

07, U

GI 5

674

P1

Cte

noid

es a

nnul

ata

(Lam

arck

, 181

9)U

F 29

2762

, UG

I 624

1P

1,2

Cte

noid

es o

shim

ensi

s (So

wer

by, 1

914)

UF

2881

39

P13

1,2

Cte

noid

es sp

. 2

UF

2972

38

P2

Div

arili

ma

sp. 1

(cor

ruga

ted)

U

F 28

7574

P

2D

ivar

ilim

a sp

. 2 (a

ff. e

lega

ns H

ayam

i & K

ase,

199

3)

UF

2880

96

P2

Div

arili

ma

sp. 3

(coa

rse)

U

F 28

7544

P

2Is

olim

ea li

mop

sis (

Nom

ura

& Z

inbo

, 193

4)U

F 29

7241

P

2Li

ma

ogas

awar

aens

is H

abe,

199

3U

F 28

7648

P

2Li

ma

vulg

aris

(Lin

k, 1

807)

U

F 29

6288

, UF

2957

78(S

), U

GI 6

236

P1

Lim

ea sp

. 1

UF

2875

97

P2

Lim

aria

bas

ilani

ca (A

. Ada

ms &

Ree

ve, 1

850)

UF

2972

45, U

GI 6

224

P2

Lim

aria

frag

ilis (

Gm

elin

, 179

1)U

F 28

2396

, UG

I 622

6P

GP5

88-1

01

Lim

aria

sp. 2

U

F 29

7248

P

2Li

mar

ia sp

. 3

UF

2881

16

P2

Lim

aria

sp. 4

U

F 28

7337

P

2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsLi

mar

ia sp

. 5

UF

2871

79

P2

Lim

aria

sp. 6

U

F 29

7244

P

1Li

mat

ula

stra

ngei

(Sow

erby

, 187

2)U

F 28

7274

, UG

I 635

1P

2Li

mat

ula

sp. 1

U

F 29

7244

P

2Li

mat

ula

sp. 2

U

F 29

7235

P

1PT

ERIO

MO

RPH

IA: O

STR

EID

AE

Cra

ssos

trea

gig

as (T

hunb

erg,

179

3)

5,9

141

Den

dost

rea

sand

vich

ensi

s (So

wer

by in

Ree

ve, 1

871)

1

151

Loph

a cr

ista

galli

(Lin

naeu

s, 17

58)

USN

M 6

0808

410

161

Plan

ostr

ea p

estig

ris (

Han

ley,

184

6)U

SNM

301

779b

P

1Sa

ccos

trea

cuc

culla

ta (B

orn,

177

8)U

F 29

8651

, USN

M 2

5451

3, U

GI 1

902

1,5,

121

Sacc

ostr

ea e

chin

ata

(Quo

y &

Gai

mar

d, 1

835)

UF

2985

13

1,5,

917

1Sa

ccos

trea

tube

rcul

ata

(Lam

arck

, 180

4)

118

1PT

ERIO

MO

RPH

IA: G

RY

PHA

EID

AE

Hyo

tissa

hyo

tis (L

inna

eus,

1758

)U

F 29

8650

, UG

I 548

0P

1Pa

rahy

otis

sa n

umis

ma

(Lam

arck

, 181

9)U

F 28

9046

P

1H

ETER

OD

ON

TA: L

UC

INID

AE

Anod

ontia

ovu

m (R

eeve

, 185

0)U

F 29

6843

T1

Anod

ontia

ves

icul

a (G

ould

, 185

0)U

F 28

7236

T

2An

odon

tia sp

. 1 (n

. sp.

?)U

F 30

0527

T2

Anod

ontia

sp. 2

(n. s

p.?)

UF

2877

25T

2“A

nodo

ntia

” sp

. 3 (n

. sp.

?)U

F 29

6756

T2

Cod

akia

pay

teno

rum

(Ire

dale

, 193

7)U

F 28

7703

, UG

I 642

8P

1C

odak

ia p

unct

ata

(Lin

naeu

s, 17

58)

UF

2968

39, U

GI 3

484

2252

1C

odak

ia ti

geri

na (L

inna

eus,

1758

)U

SNM

851

147,

UG

I 642

2P

1“E

pico

daki

a” b

ella

(Con

rad,

183

7)U

F 28

7414

, UG

I 144

222

2,51

1Ep

icod

akia

swee

ti (H

edle

y, 1

899)

UF

2848

87, U

GI 6

425

P1

Funa

futia

levu

kana

(Sm

ith, 1

885)

UF

2880

26, U

GI 6

483

P1,

2M

yrte

a sp

. 1

UF

2876

94, U

GI 6

431

T1,

2Pa

rvilu

cina

s.l.

sp. 4

U

F 28

7737

, UG

I 647

2P

1,2

Parv

iluci

na s.

l. sp

. 5

UF

2871

83, U

GI 6

480

P2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsPa

rvilu

cina

s.l.

sp. 6

U

F 28

7650

, UG

I 647

0P

2W

allu

cina

fijie

nsis

(Sm

ith, 1

885)

UF

2884

40, U

GI 6

484

131

Wal

luci

na sp

. 3

UF

2972

03, U

GI 5

899

P2

HET

ERO

DO

NTA

: FIM

BR

IID

AE

Fim

bria

fim

bria

ta (L

inna

eus,

1758

)U

F 28

2434

, UG

I 172

414

(MS)

,22

501

HET

ERO

DO

NTA

: TH

YA

SIR

IDA

ETh

yasi

ridae

sp. 1

U

F 28

8400

S

2H

ETER

OD

ON

TA: U

NG

ULI

NID

AE

Dip

lodo

nta

sp. 1

U

F 28

7784

P

2D

iplo

dont

a sp

. 2

UF

2877

52

P2

HET

ERO

DO

NTA

: CH

AM

IDA

EAm

phic

ham

a sp

. 1 (n

. sp.

?)U

F 30

0741

M2

Cha

ma

aspe

rella

Lam

arck

, 181

9U

F 28

2420

, UG

I 573

9P

GP7

78-2

71

Cha

ma

bras

sica

Ree

ve, 1

847

UF

2985

06 U

SNM

248

485a

51

Cha

ma

fibul

a R

eeve

, 184

6U

F 29

8753

5

191

Cha

ma

iost

oma

Con

rad,

183

7D

MN

H 8

4404

, UG

I 544

3P

201

Cha

ma

laza

rus (

Linn

aeus

, 175

8)U

F 29

8505

, UG

I 544

4, U

SNM

248

485,

U

SNM

607

895(

S), D

MN

H 8

5947

, P

GP5

25-7

1S

Cha

ma

mac

erop

hylla

(Gm

elin

, 179

1)U

F 29

8754

5

211

Cha

ma

paci

fica

Bro

derip

, 183

5U

GI 5

744

51

Cha

ma

sp. 4

U

F 30

0742

P1

Cha

ma

spin

osa

Ree

ve, 1

847

sens

u La

my,

190

6 U

F 28

2406

P

1H

ETER

OD

ON

TA: C

AR

DIT

IDA

E57

Car

dita

var

iega

ta B

rugu

ière

, 179

2U

F 28

4886

, USN

M 8

4967

1, U

GI 6

404

P1

Car

dite

lla c

f. ie

jimen

sis H

ayam

i & K

ase,

199

3U

F 28

8460

, UG

I 633

13

1,2

Car

dite

lla sp

. 2

UF

2888

17

P2

Gla

ns sp

. 1

UF

2872

41, U

GI 6

502

P2

HET

ERO

DO

NTA

: GA

LEO

MM

ATI

DA

E S.

L.

Barr

imys

ia c

umin

gi (A

. Ada

ms,

1856

)U

F 28

7322

, UG

I 628

7P

1Ep

hipp

odon

ta sp

. 1

UF

2883

94

P1

Gal

eom

ma

sp. 1

(ora

nge)

U

F 29

8631

S

1

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsG

aleo

mm

a sp

. 2 (b

urgu

ndy)

U

F 28

8266

S

1G

aleo

mm

a s.

l. sp

. 3 (p

itted

) U

F 28

8260

S

21

Gal

eom

ma

s. l.

sp. 4

U

F 28

8219

S

?GP8

52-8

21

Gal

eom

ma

s. l.

sp. 5

U

F 28

8258

S

21

Gal

eom

ma

s. l.

sp. 6

U

F 29

8784

S

21

Gal

eom

ma

s. l.

sp. 7

U

F 28

8368

S

21

Gal

eom

ma

sp. 8

U

F 28

8257

S

1G

aleo

mm

a s.

l. sp

. 9

UF

2876

83

P2

1G

aleo

mm

a sp

. 10

UF

2873

47

P1

Gal

eom

mat

idae

sp. 1

U

F 28

8391

S

1G

aleo

mm

atid

ae sp

. 2

UF

2987

87

S1

Gal

eom

mat

idae

sp. 3

U

F 28

8220

S

21,

2K

ellia

sp. 1

U

F 28

8376

S

1,2

Kel

lia sp

. 2

UF

2987

85

S1

Kel

lia sp

. 3

UF

2987

83

S1

Lasa

eida

e? sp

. 1

UF

2882

43

S1

Lasa

eida

e sp

. 2

UF

2874

47

P1

Lasa

eida

e sp

. 3

UF

2883

67

S2

1M

ysel

la sp

. 1

UF

2882

29

S1

Mys

ella

sp. 2

U

F 28

8382

S

1,2

Plan

ktom

ya sp

. 1

UF

2875

93

P1,

2Ro

chef

ortia

sp. 1

U

F 28

8242

S

2Ro

chef

ortia

sp. 2

U

F 28

8212

, UG

I 644

7S

1Ro

chef

ortia

sp. 3

U

F 28

8265

S

1Ro

chef

ortia

sp. 4

U

F 28

8402

S

1Ro

chef

ortia

s. l.

sp. 5

U

F 28

7927

P

22

Roch

efor

tia sp

. 6

UF

2883

83

S1

Scin

tilla

cuv

ieri

? D

esha

yes,

1856

UF

2986

29, U

GI 6

455

SG

P534

-19

1Sc

intil

la s.

l. sp

. 1 (w

hite

) U

F 28

4926

S

GP5

82-2

02

1Sc

intil

la s.

l. sp

. 2

UF

2875

25

S2

1Sc

intil

la s.

l. sp

. 3

UF

2938

39

S2

1

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsSc

intil

la s.

l. sp

. 4

UF

2883

69

S2

1Sc

intil

la s.

l. sp

. 5

UF

2848

47

P2

1Sc

intil

la?

s. l.

sp. 6

U

F 28

8247

S

21

Scin

tillo

na b

ryan

i (Pi

lsbr

y, 1

921)

UF

2938

37, U

GI 6

448

P1

Scin

tillo

na sp

. 1

UF

2969

33, U

GI 6

452

SG

P541

-11

HET

ERO

DO

NTA

: SPO

RTE

LLID

AE

Anis

odon

ta c

aled

onic

a (F

isch

er, 1

886)

UF

2873

23, U

GI 6

280

P1

HET

ERO

DO

NTA

: CA

RD

IID

AE

Acro

ster

igm

a bi

radi

atum

(Bru

guiè

re, 1

789)

UF

2878

69

V2

Acro

ster

igm

a ho

bbsa

e V

idal

, 199

9U

F 28

7986

(par

atyp

e), U

GI 6

353

151

Acro

ster

igm

a pr

ofun

dum

Vid

al, 1

999

UF

2879

17

V2

Afro

card

ium

exo

chum

(Mel

vill

& S

tand

en, 1

906)

UF2

8882

2 V

1Af

roca

rdiu

m ri

char

di (A

udou

in, 1

827)

UF

2877

54, U

GI 6

278

V2

Cor

culu

m d

iona

eum

(Bro

derip

& S

ower

by, 1

829)

UF

2889

24, U

GI 5

333

P1

Cte

noca

rdia

forn

icat

a (S

ower

by, 1

841)

UF

2872

42, U

GI 5

890

P2

Cte

noca

rdia

vic

tor (

Ang

as, 1

872)

UF

2889

36, U

GI 6

460

PG

P771

-19

2C

teno

card

ia sp

. 1 (n

. sp.

) U

F 28

8938

, UG

I 646

9P

1,2

Frag

um fr

agum

(Lin

naeu

s, 17

58)

UF

2889

22, U

SNM

846

285,

UG

I 421

822

GP5

34-3

348

1Fr

agum

looc

hooa

num

Kira

, 195

9U

F 28

8949

, UG

I 654

9P

1Fr

agum

mun

dum

(Ree

ve, 1

845)

UF

2889

55, U

GI 6

463

P1

Frag

um su

ezie

nse

(Iss

el, 1

869)

UF

2888

02, U

GI 6

465

P1

Frig

idoc

ardi

um k

iran

a Sa

kura

i & H

abe,

196

6U

F 28

7512

V

2Fr

igid

ocar

dium

torr

esi (

Smith

, 188

5)

UF

2873

14

V2

Fulv

ia a

ustr

alis

(Sow

erby

, 183

4)U

F 29

5375

, UG

I 645

8P

GP5

17-3

31

Fulv

ia u

ndat

opic

ta (P

ilsbr

y, 1

904)

UF

2871

51, U

GI 6

366

V1,

2H

ippo

pus h

ippo

pus (

Linn

aeus

, 175

8)U

GI 1

892

P22

1Lu

nulic

ardi

a cf

. ret

usa

(Lin

naeu

s, 17

67)

UF

2872

53

P23

2M

icro

frag

um fe

stiv

um (D

esha

yes,

1855

)U

F 28

7733

P

2N

emoc

ardi

um ly

ratu

m (S

ower

by, 1

834)

UF

2985

09, U

SNM

851

267

P1

Nem

ocar

dium

? sp

. 1

UF

2889

46

P1,

2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsTr

idac

na d

eras

a (R

ödin

g, 1

798)

5,

9R

FM-

213A

241

Trid

acna

gig

as (L

inna

eus,

1758

)

5,9

251

Trid

acna

max

ima

(Röd

ing,

179

8)U

F 29

7124

, USN

M 4

8705

3, U

GI 1

317

16(G

,M,S

),22

GP7

42-1

249

1M

,S

Trid

acna

squa

mos

a La

mar

ck, 1

819

UG

I 306

216

(S,T

)G

P644

-22

1S,

TVa

stic

ardi

um e

long

atum

eno

de (S

ower

by, 1

840)

U

F 28

8838

, UG

I 567

817

1Va

stic

ardi

um o

rbita

phi

lippi

nens

e (H

edle

y, 1

899)

UF

2888

37, U

SNM

851

098,

UG

I 637

118

1Va

stic

ardi

um se

wel

li (P

rash

ad, 1

932)

UF

2875

27, U

SNM

849

694

151

HET

ERO

DO

NTA

: TEL

LIN

IDA

EAe

nigm

otel

lina

sp. 1

U

F 28

7671

P

2Ar

cope

lla is

seli

(H. A

dam

s, 18

70)

UF

2871

86, U

GI 6

475

P2

Cad

ella

sem

en (H

anle

y, 1

845)

UF

2885

44, U

GI 6

534

P1,

2C

lath

rote

llina

tenu

istr

iata

(Sow

erby

, 186

7)U

F 28

8104

, UG

I 589

3P

1,2

Cyc

lote

llina

rem

ies (

Linn

aeus

, 175

8)U

F 30

0524

, UG

I 637

4P

1El

lipto

telli

na e

ugly

pta

(Gou

ld, 1

861)

UF

2873

53, U

GI 6

397

P1,

2El

lipto

telli

na sp

. 1

UF

2875

08

P2

Ellip

tote

llina

sp. 2

U

F 28

7567

P

2Ex

otic

a fij

iens

is (S

ower

by, 1

868)

UF

2885

34, U

GI 6

522

P26

1Ex

otic

a sp

. 1

UF

2877

45, U

GI 6

505

P27

1,2

Exot

ica

sp. 2

(aff

. neb

ulos

us (D

all,

Bar

tsch

& R

ehde

r,19

38))

U

F 28

8506

, UG

I 650

8P

281,

2

Exot

ica

sp. 3

U

F 28

8491

P

2H

emim

etis

ang

ulat

a (L

inna

eus,

1767

)U

SNM

851

056

P1

Jact

ellin

a ob

liqua

ria

(Des

haye

s, 18

55)

UF

2840

12, U

GI 6

379

P1

Loxo

glyp

ta c

lath

rata

(Des

haye

s, 18

35)

UF

2956

87, U

GI 5

737

2242

1Lo

xogl

ypta

sp. 1

U

F 29

7230

, UG

I 171

1P

1“M

acom

inae

” sp

. 1 (p

ink)

U

F 28

9101

, UG

I 640

3P

1“M

acom

inae

” sp

. 2 (t

rans

pare

nt)

UF

2872

18

P1

Obt

ellin

a bo

ugei

(Sow

erby

, 190

9)U

F 28

4843

, UG

I 638

4P

1Ph

arao

nella

cf.

astu

la (H

edle

y, 1

917)

UF

2877

43

P2

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsPi

ngui

telli

na ro

bust

a (H

anle

y, 1

844)

UF

2956

93, U

GI 3

588

2244

1,2

Ping

uite

llina

sp. 1

U

F 28

7427

P

1Q

uadr

ans g

arga

dia

of a

utho

rs, n

ot L

inna

eus,

1758

UF

2985

12

P29

1Q

uadr

ans p

arvi

tas I

reda

le, 1

931

UF

2877

29, U

GI 6

532

P1,

2Q

uidn

ipag

us p

alat

am Ir

edal

e, 1

929

UF

3005

26, U

GI 1

422

19,2

243

1Sc

issu

lina

disp

ar (C

onra

d, 1

837)

UF

2888

11

2241

1Sc

utar

copa

gia

lingu

afel

is (L

inna

eus,

1758

)U

F 29

6943

, UG

I 573

8P

1Sc

utar

copa

gia

scob

inat

a (L

inna

eus,

1758

)U

F 29

6446

, USN

M 8

4970

3, U

GI 1

538

P1

Telli

na c

hari

essa

Sal

isbu

ry, 1

934

UF

2969

42

P30

1Te

llina

s.l.

pata

giat

a Pr

asha

d, 1

932

UF

2871

77, U

GI 6

394

P2

2Te

llina

s.l.

sp. 2

U

F 29

7251

P

21

Telli

na s.

l. sp

. 3

UF

2885

23

P2

1Te

llina

s.l.

sp. 4

U

F 28

8499

P

21

Telli

nella

cru

cige

ra (L

amar

ck, 1

818)

UF

2856

15, U

GI 5

735

2241

1Te

lline

lla sp

. 2 (a

ff. v

erru

cosa

(Han

ley,

184

4))

UF

2881

50, U

GI 6

381

P1,

2Te

lline

lla st

aure

lla (L

amar

ck, 1

818)

UF

2985

08, U

GI 6

383

2241

1Te

lline

lla v

irga

ta (L

inna

eus,

1758

)U

SNM

607

844(

S), U

GI 2

212,

U

GI 1

447(

S)22

411

S

Telli

nide

s ova

lis (S

ower

by, 1

825)

UF

2881

36, U

GI 6

399

P1

HET

ERO

DO

NTA

: SEM

ELID

AE

Abra

seur

ati L

amy,

190

6U

F 28

8519

, UG

I 647

9P

1,2

Abra

sp. 1

U

F 28

7700

P

2Er

vilia

bis

culp

ta G

ould

, 186

1U

F 28

8064

P

1,2

Lept

omya

psi

ttacu

s Han

ley,

188

2U

F 28

8850

, UG

I 639

1P

1Lo

noa

haw

aien

sis D

all,

Bar

tsch

& R

ehde

r, 19

38U

F 28

8525

, UG

I 652

8P

1Ro

chef

ortin

a sa

ndw

iche

nsis

(Sm

ith, 1

885)

UF

2884

49, U

GI 6

294

P1,

2Se

mel

e au

stra

lis (S

ower

by, 1

832)

UF

2840

22, U

GI 6

490

P1

Sem

elid

ae sp

. 1

UF

2884

93

P2

HET

ERO

DO

NTA

: PSA

MM

OB

IID

AE

Asap

his v

iola

scen

s (Fo

rssk

ål, 1

775)

UF

3005

25, U

GI 6

377

2240

1G

ari p

usill

a B

ertin

, 188

0U

F 28

8496

, USN

M 8

5129

8, U

GI 6

400

P1

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsG

ari s

ibog

ai P

rash

ad, 1

932

UF

2881

13, U

GI 6

389

202

HET

ERO

DO

NTA

: SO

LEC

UR

TID

AE

Sole

curt

us sp

. 1

UF

2873

03

P2

HET

ERO

DO

NTA

: MA

CTR

IDA

EM

actri

dae

sp. 1

U

F 28

7759

, UG

I 649

2P

1H

ETER

OD

ON

TA: M

ESO

DES

MA

TID

AE

Atac

tode

a st

riat

a (G

mel

in, 1

791)

UF

2985

11, U

GI 1

705

P31

1H

ETER

OD

ON

TA: T

RA

PEZI

DA

EC

oral

lioph

aga

cora

lliop

haga

? (G

mel

in, 1

791)

UF

2884

75, U

GI 6

500

332

1G

loss

ocar

dia

obes

a (R

eeve

, 184

3)U

F 28

8076

, USN

M 8

5128

4,

UF

2880

75(S

), U

GI 6

207

31

Trap

eziu

m o

blon

gum

(Lin

naeu

s, 17

58)

UF

2880

63, U

GI 6

200

P1

Trap

eziu

m su

blae

viga

tum

(Lam

arck

, 181

9)U

F 28

3530

, UG

I 619

95

1H

ETER

OD

ON

TA: G

LOSS

IDA

EM

eioc

ardi

a m

oltk

iana

(Spe

ngle

r, 17

83)

UF

2867

83

P2

HET

ERO

DO

NTA

: KEL

LIEL

LID

AE

Alve

inus

sp. 1

U

F 28

8410

P

2Al

vein

us s.

l. sp

. 2

UF

2987

81

P2

HET

ERO

DO

NTA

: VEN

ERID

AE

Cal

lista

acc

inct

a (R

ömer

, 186

4)

UF

2872

33

P2

Dor

isca

cf.

cook

ei D

all,

Bar

tsch

& R

ehde

r, 19

38U

F 28

7248

, UG

I 654

6P

2D

osin

ia sp

. 7

UF

2874

69, U

GI 6

495

P1

Dos

inia

sp. 8

U

F 28

7313

P

2G

afra

rium

pec

tinat

um (L

inna

eus,

1758

)U

F 28

7472

, USN

M 2

4741

5, U

GI 1

452,

U

SNM

607

852(

S)22

461

S

Gaf

rari

um tu

mid

um R

ödin

g, 1

798

UF

2874

65, U

SNM

774

709,

UG

I 191

92

11

Glo

bive

nus t

oreu

ma

(Gou

ld, 1

851)

UF

2824

05, U

GI 5

373

P1,

2G

ould

ia s.

l. sp

. 1

UF

2881

69

P1

Gou

ldia

s. l.

sp. 2

(aff

. kilb

urni

Driv

as &

Jay,

199

0)

UF

2881

70

P2

Lioc

onch

a ca

stre

nsis

(Lin

naeu

s, 17

58)

UF

2824

47, U

GI 4

518

P1

Lioc

onch

a or

nata

(Lam

arck

, 181

7)

UF

2874

73, U

GI 5

357

P1

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsLi

ocon

cha

phili

ppin

arum

(Han

ley,

184

4)U

F 28

7714

, UG

I 573

3P

1,2

Lioc

onch

a m

elha

rtae

Lam

prel

l & S

tani

sic,

199

6U

F 28

7304

, UG

I 649

8P

2Ir

us sp

. 1

UF

2881

76

P1,

2Pe

rigl

ypta

pue

rper

a (L

inna

eus,

1771

)U

F 28

5618

, USN

M 2

4774

5, U

GI 1

726,

U

SNM

608

009(

S)P

1S

Peri

glyp

ta re

ticul

ata

(Lin

naeu

s, 17

58)

UF

2887

15, U

SNM

847

262,

UG

I 474

522

471

Peri

glyp

ta c

risp

ata

(Des

haye

s, 18

53)

UF

2824

13, A

NSP

197

714,

UG

I 642

0,

USN

M 6

0798

0(S)

P1

S

Pita

r fes

toui

Har

te, 1

993

UF

2876

64, U

GI 5

889

P2

Pita

r int

rica

ta (D

autz

enbe

rg, 1

907)

UF

2835

35, U

GI 6

414

P1

Pita

r pro

ra (C

onra

d, 1

837)

UF

2874

51, U

GI 1

704,

UG

I 145

9(S)

2245

1Pi

tar s

poor

i Lam

prel

l & W

hite

head

, 199

0U

F 29

3836

, UG

I 640

9P

1Pi

tar s

p. 6

(dee

p)

UF

2872

38, U

GI 6

417

P2

Sam

aran

gia

quad

rang

ular

is (A

dam

s & R

eeve

, 185

0)U

F 28

7623

, UG

I 588

4P

2Ti

moc

lea

mar

ica

(Lin

naeu

s, 17

58)

UF

2848

45, U

SNM

851

223,

UG

I 428

4P

1Ti

moc

lea

sp. 9

U

GI 6

516

P33

1Ti

moc

lea

sp. 1

0 U

F 28

7942

, UG

I 651

0P

2Ti

moc

lea

sp. 1

1 U

F 28

7349

, UG

I 652

4P

1,2

Tim

ocle

a sp

. 12

UF

2880

40, U

GI 6

518

P2

HET

ERO

DO

NTA

: MY

OID

A: C

OR

BU

LID

AE

Cor

bula

tahe

itens

is L

amar

ck, 1

818

UF

2873

43

P1,

2Va

rico

rbul

a ro

talis

(Hin

ds, 1

843)

UF

2875

41

P1,

2H

ETER

OD

ON

TA: M

YO

IDA

: GA

STR

OC

HA

ENID

AE

Roce

llari

a gi

gant

ea (D

esha

yes,

1830

)U

F 29

6963

, UG

I 654

5C

1D

ufoi

chae

na d

entif

era?

(Duf

o, 1

840)

UF

2875

07C

1D

ufoi

chae

na p

exip

hora

? (S

tura

ny, 1

899)

UF

2895

51C

1G

astr

ocha

ena

cune

iform

is S

peng

ler,

1783

UF

2969

74C

1Sp

engl

eria

plic

atili

s (D

esha

yes,

1855

)U

F 28

4769

C

GP8

26-1

21

HET

ERO

DO

NTA

: MY

OID

A: T

ERED

INID

AE

Lyro

dus p

edic

illat

us (Q

uatre

fage

s, 18

49)

11(G

,S),2

134

1S

Not

oter

edo

edax

(Hed

ley,

189

5)

1135

App

endi

x 1.

Biv

alve

s of G

uam

/ (c

ontin

ued)

Taxo

nM

useu

mR

efPh

oto

Not

eD

epth

IsTe

redo

cla

ppi B

arts

ch, 1

923

11

,21

361

Tere

do fu

lleri

Cla

pp, 1

923

21

1Te

redo

furc

ifera

Mar

tens

, 189

4

11,2

137

1Te

redo

thyr

a m

atoc

otan

a B

arts

ch, 1

927

11

38A

NO

MA

LOD

ESM

ATA

: PO

RO

MY

IDA

EPo

rom

ya sp

. 1

UF

2848

83

P2

AN

OM

ALO

DES

MA

TA: V

ERTI

CO

RD

IID

AE

Spin

osip

ella

eri

cia

(Hed

ley,

191

1)U

F 28

7249

, UG

I 628

5P

2A

NO

MA

LOD

ESM

ATA

: CU

SPID

AR

IID

AE

Cus

pida

ria

sp. 1

U

F 28

7919

, UG

I 634

3P

2

Ref

eren

ces:

Lite

ratu

re r

ecor

ds li

sted

by

num

ber,

iden

tifie

r of

spe

cies

not

pre

viou

sly

docu

men

ted

from

the

Mar

iana

s lis

ted

lette

r as

fol

low

s. C

) J.

Car

ter,

H) I

. Hay

ami,

M) A

. Mat

suku

ma,

P) G

. Pau

lay,

S) P

. Val

entit

ch S

cott,

T) J

. Tay

lor,

V) J

. Vid

al, W

) T.R

. Wal

ler,

1) B

rale

y 19

84, 2

)A

mes

bury

199

9, 3

) H

ayam

i & K

ase

1996

, 4)

Ros

ewat

er 1

961,

5)

Paul

ay e

t al.

2002

, 6)

Wal

ler

1972

, 7)

Rai

nes

1996

, 8)

Hut

sell

1999

, 9)

Eldr

edge

199

4, 1

0) S

tenz

el 1

971,

11)

Roc

h 19

76, 1

2) B

rale

y 19

82, 1

3) G

love

r & T

aylo

r 200

1, 1

4) N

icol

195

0, 1

5) V

idal

199

9, 1

6) R

osew

ater

1965

, 17)

Vid

al 1

993,

18)

Vid

al 1

997,

19)

Day

197

7, 2

0) W

illan

199

3, 2

1) E

dmon

dson

194

6, 2

2) H

idal

go 1

904-

1905

Not

es:

1) S

ubfo

ssil

rem

ains

of

Hol

ocen

e ag

e of

thes

e sp

ecie

s ar

e w

ides

prea

d on

Gua

m, h

owev

er e

xtan

t pop

ulat

ions

are

larg

ely

rest

ricte

d to

inne

r A

pra

Har

bor.

This

rest

rictio

n m

ay c

orre

spon

d to

redu

ctio

n of

man

grov

e ha

bita

ts d

urin

g th

e H

oloc

ene

(Am

esbu

ry 1

999)

. 2)

Gen

eric

ass

ignm

ents

tent

ativ

e, n

eeds

revi

sion

.3)

May

cor

resp

ond

to H

ayam

i & K

ase’

s (19

96) r

ecor

d of

Ben

thar

ca sp

. B fr

om G

uam

. 4)

May

cor

resp

ond

to H

ayam

i & K

ase’

s (19

96) r

ecor

d of

Ben

thar

ca te

nuis

? fr

om G

uam

. 5)

A m

inut

e, u

ndes

crib

ed, a

rcoi

d bi

valv

e te

ntat

ivel

y as

sign

ed to

the

Lim

opso

idea

by

Hay

ami &

Kas

e (1

993)

. 6)

Bas

ed o

n si

ght r

ecor

d by

GP,

no

vouc

her c

urre

ntly

ava

ilabl

e fo

r thi

s abu

ndan

t spe

cies

.7)

Indo

-Wes

t Pac

ific

Mal

leus

(Mal

vifu

ndus

) spe

cies

hav

e be

en o

ver-

syno

nym

ized

. The

Gua

m s

peci

es is

clo

sest

to th

e H

awai

ian

M. n

utta

lli, w

hich

,co

ntra

ry to

cur

rent

use

age,

is n

ot a

syno

nym

of M

. reg

ula.

8)

Kno

wn

only

fro

m A

pra

Har

bor,

whe

re th

is s

peci

es in

habi

ted

stee

p re

ef fa

ces.

Alth

ough

dea

d sh

ells

are

stil

l enc

ount

ered

, no

live

anim

als

have

been

seen

in y

ears

, and

this

gia

nt sp

ondy

lid a

ppea

rs to

hav

e be

en e

xtirp

ated

, pro

babl

y by

ove

rcol

lect

ing.

Bas

ed o

n si

ght r

ecor

d by

GP.

9)

Thi

s spe

cies

was

des

crib

ed fr

om H

awai

i, w

here

it is

com

mon

in P

earl

Har

bor a

nd p

oten

tially

intro

duce

d. It

was

bro

ught

to G

uam

on

the

dry

dock

Mac

hini

st in

July

, 199

9. M

ost a

nim

als d

ied

with

in a

mon

th o

f arr

ival

, so

the

spec

ies i

s unl

ikel

y to

bec

ome

esta

blis

hed

on G

uam

.10

) Ano

miid

s rem

ain

unst

udie

d on

Gua

m d

ue to

lim

ited

wel

l-pre

serv

ed m

ater

ial o

n ha

nd.

11) T

hese

spe

cies

are

und

escr

ibed

mem

bers

of a

cla

de o

f min

ute

dim

yids

, with

wel

l dev

elop

ed h

inge

teet

h, th

at o

ccur

in re

ef c

aves

thro

ugho

ut th

eIn

do-W

est P

acifi

c.

12) D

imye

lla sp

. B o

f Hay

ami &

Kas

e (1

996)

. 13

) Ofte

n in

corr

ectly

iden

tifie

d as

C. c

once

ntri

cus (

e.g.

, Oku

tani

200

0).

14) I

ntro

duce

d to

Apr

a H

arbo

r Gua

m (s

ee E

ldre

dge

1994

), bu

t app

ears

not

to h

ave

beco

me

esta

blis

hed.

15

) As O

stre

a no

mad

es in

Bra

ley

(198

4); s

ynon

ymy

afte

r Lam

prel

l & H

ealy

(199

8).

16) K

now

n on

ly fr

om d

ead

shel

ls in

Apr

a H

arbo

r dre

dgin

gs, n

ever

seen

aliv

e on

Gua

m.

17) I

ntro

duce

d to

Apr

a H

arbo

r, G

uam

(see

Bra

ley

1984

and

Eld

redg

e 19

94).

18) A

s Sac

cost

rea

cucu

llata

tube

rcul

ata

in B

rale

y (1

984)

; int

rodu

ced

to G

uam

.19

) Bro

ught

to G

uam

from

Haw

aii o

n th

e dr

y do

ck M

achi

nist

in J

uly,

199

9. M

ost a

nim

als

died

with

in a

mon

th o

f arr

ival

, so

the

spec

ies

is u

nlik

ely

to e

stab

lish

on G

uam

.20

) Kilb

urn

& R

ippe

y (1

982)

and

Pau

lay

(198

7, 1

989)

hav

e pr

evio

usly

refe

rred

to th

is s

peci

es a

s C

ham

a lim

bula

Lam

arck

, 181

9. I

have

sin

ce s

een

Lam

arck

’s ty

pes a

nd th

ey d

o no

t mat

ch th

is c

omm

on In

do-W

est P

acifi

c sp

ecie

s wel

l. C

ham

a io

stom

a is

the

olde

st c

lear

ly v

alid

nam

e.

21) T

his

Wes

t Atla

ntic

spe

cies

was

bro

ught

to G

uam

from

Haw

aii o

n th

e dr

y do

ck M

achi

nist

in J

uly,

199

9 in

gre

at a

bund

ance

. Mos

t ani

mal

s di

edw

ithin

a m

onth

of a

rriv

al, s

o th

e sp

ecie

s is u

nlik

ely

to b

ecom

e es

tabl

ishe

d on

Gua

m.

22) T

his s

peci

es is

ext

irpat

ed o

n G

uam

, alth

ough

Hol

ocen

e sh

ells

are

occ

asio

nally

foun

d. T

he o

nly

livin

g in

divi

dual

reco

rded

(UG

I 189

2) w

as ta

ken

in A

pra

Har

bor n

ear t

he d

ocks

and

is su

spec

ted

to h

ave

been

dro

pped

off

a b

oat a

fter b

eing

bro

ught

from

ano

ther

isla

nd.

23) K

now

n on

ly fr

om w

orn,

mos

tly s

mal

l (<1

cm) s

hells

from

dee

p (~

100m

) wat

er. W

heth

er th

ese

repr

esen

t an

exta

nt d

eep-

wat

er p

opul

atio

n or

ano

w e

xtirp

ated

pop

ulat

ion

from

the

glac

ial l

ow s

tand

is n

ot k

now

n. S

imila

r sm

all L

unul

icar

dia

are

com

mon

ly d

redg

ed d

eep

from

the

lago

ons

of th

e M

arsh

all I

slan

ds (U

SNM

col

lect

ions

). 24

) Int

rodu

ced

from

Pal

au. T

he in

trodu

ced

anim

als a

re st

ill a

live

(see

pho

to c

ited)

, but

no

recr

uitm

ent h

as b

een

obse

rved

. 25

) An

atte

mpt

to in

trodu

ce T

. gig

as to

Gua

m (E

ldre

dge

1994

) was

app

aren

tly n

ot su

cces

sful

. Thi

s spe

cies

has

a P

leis

toce

ne fo

ssil

reco

rd o

n G

uam

,bu

t the

pop

ulat

ion

appe

ars

to h

ave

died

out

bef

ore

the

arriv

al o

f peo

ple,

per

haps

dur

ing

a gl

acia

l low

sta

nd, a

s no

Hol

ocen

e sh

ells

are

kno

wn

(Pau

lay

1996

b).

26) S

emel

angu

lus s

p. 3

of P

aula

y (1

996b

); A

. Mat

suku

ma

(in p

rep.

) con

side

rs S

emel

angu

lus t

o be

a ju

nior

syno

nym

of E

xotic

a.27

) Sem

elan

gulu

s sp.

1 o

f Pau

lay

(199

0); n

ot 2

6.28

) Sem

elan

gulu

s sp.

2 o

f Pau

lay

(199

0) a

nd S

emel

angu

lus c

rebr

imac

ulat

us o

f Pau

lay

(199

6b),

not S

ower

by, 1

868;

see

not 2

6.

29)

Kno

wn

from

a s

ingl

e, d

ead

sam

ple

of w

orn

shel

ls o

nly,

fro

m th

e Pi

ti re

ef f

lat.

This

spe

cies

is c

omm

on in

ato

ll la

goon

s th

roug

h m

uch

of th

ew

este

rn P

acifi

c.

30) K

now

n fr

om a

sing

le, f

resh

val

ve o

n G

uam

col

lect

ed o

n th

e le

ewar

d fo

re re

ef.

31) A

lthou

gh th

is su

rf c

lam

is a

bund

ant o

n m

ost i

slan

ds w

here

it o

ccur

s, it

is ra

re o

n G

uam

and

has

not

bee

n se

en a

live

in th

e pa

st d

ecad

e.

32)

As

C.

hyal

ina

in H

ayam

i &

Kas

e (1

996)

, w

hich

may

be

a va

lid n

ame

for

som

e sp

ecim

ens.

The

cons

ider

able

mor

phol

ogic

al v

aria

tion

inC

oral

lioph

aga

has

been

attr

ibut

ed to

thei

r en

dolit

hic

lifes

tyle

, and

onl

y tw

o In

do-W

est P

acifi

c sp

ecie

s w

ere

reco

gniz

ed in

the

last

rev

isio

n(S

olem

195

4). T

he p

ossi

bilit

y th

at a

dditi

onal

cry

ptic

div

ersi

ty m

ay u

nder

lie th

is v

aria

tion

is w

orth

inve

stig

atin

g.

33) A

sm

all s

peci

es th

at m

ust h

ave

been

ext

rem

ely

abun

dant

in in

ner A

pra

Har

bor s

ome

time

in th

e pa

st, b

ut I

have

nev

er s

een

it al

ive.

Whe

n th

eIn

ner H

arbo

r was

exc

avat

ed a

fter W

WII

, dre

dge

spoi

ls w

ere

depo

site

d ar

ound

Riz

al B

each

in n

orth

ern

Aga

t Bay

. Tim

ocle

a sp

. 1 sh

ells

was

hed

out o

f the

se s

poils

now

form

a s

hell

hash

on

larg

e be

ach

area

s he

re a

nd w

e ha

ve e

ven

been

col

lect

ed o

ffsh

ore

in th

e ar

ea a

s w

ell.

Num

erou

sot

her

mol

lusk

s (e

.g.,

Spon

dylu

s m

ultim

uric

atus

, Lop

ha c

rist

agal

li, C

ham

a br

assi

ca)

as w

ell a

s fo

ssil

crab

s ha

ve a

lso

was

hed

out o

f ha

rbor

dred

ge sp

oils

in th

is a

rea.

See

Kes

ling

(195

8) fo

r a re

view

of t

he o

rigin

of t

hese

dep

osits

. 34

) As T

ered

o di

egen

sis i

n Ed

mon

dson

(194

6); a

s Ter

edo

yats

ui in

Roc

h (1

976)

; syn

onym

y af

ter T

urne

r (19

66).

35) A

s Ter

edo

jutti

ngae

in R

och

(197

6); s

ynon

ymy

afte

r Tur

ner (

1966

).36

) As T

ered

o tr

ullif

orm

is in

Edm

onds

on (1

946)

; syn

onym

y af

ter T

urne

r (19

66).

37) A

s Ter

edo

bens

oni i

n Ed

mon

dson

(194

6) a

nd R

och

(197

6); s

ynon

ymy

afte

r Tur

ner (

1966

).38

) As T

ered

o m

atoc

otan

a in

Roc

h (1

976)

; gen

eric

ass

igne

men

t afte

r Tur

ner (

1966

).39

) Pe

cten

cor

usca

ns H

inds

, 184

5, c

urre

ntly

ass

igne

d to

Sem

ipal

lium

, diff

ers

from

oth

er m

embe

rs o

f th

e ge

nus

in d

evel

opin

g di

rect

ly f

rom

the

prod

isso

conc

h to

the

radi

ally

plic

ate

stag

e, w

ithou

t dep

ositi

ng a

n in

terv

enin

g pr

ism

atic

she

ll la

yer

in th

e ea

rly d

isso

conc

h of

the

right

val

ve(W

alle

r 19

72).

Pasc

ahin

nite

s is

an

avai

labl

e ge

neric

nam

e fo

r th

is li

neag

e, e

rect

ed f

or P

ecte

n pa

sca

Dal

l, 19

08, a

n Ea

ster

Isl

and

ende

mic

offs

hoot

from

cor

usca

ns s

tock

. Alth

ough

Pas

cahi

nnite

s w

as e

rect

ed b

ecau

se o

f the

cem

ente

d lif

esty

le o

f pas

ca (D

ijkst

ra &

Rai

nes

1999

), th

ela

ck o

f an

exte

rnal

pris

mat

ic la

yer i

n th

e ea

rly ri

ght d

isso

conc

h is

the

diag

nost

ic c

hara

cter

of t

he g

enus

(T.R

. Wal

ler p

ers.

com

m. 2

002)

.40

) Mis

iden

tifie

d as

Asa

phis

def

lora

ta in

Hid

algo

(190

4-19

05),

an A

tlant

ic c

onge

ner.

41) A

ll cl

assi

fied

as T

ellin

a in

Hid

algo

(190

4-19

05).

42) A

s Tel

lina

rhom

boid

es in

Hid

algo

(190

4-19

05),

syno

nym

y af

ter L

ampr

ell &

Whi

tehe

ad (1

992)

.43

) As T

ellin

a ru

gosa

in H

idal

go (1

904-

1905

), sy

nony

my

afte

r Kay

(197

9).

44) C

lass

ified

as A

rcop

agia

in H

idal

go (1

904-

1905

).45

) As C

arya

tis o

bliq

uata

in H

idal

go (1

904-

1905

), sy

nony

my

afte

r Lam

prel

l & W

hite

head

(199

2).

46) C

lass

ified

as C

rist

a pe

ctin

ata

in H

idal

go (1

904-

1905

).47

) Cla

ssifi

ed a

s Ven

us in

Hid

algo

(190

4-19

05).

48) C

lass

ified

as H

emic

ardi

um in

Hid

algo

(190

4-19

05).

49) A

s Tri

dacn

a m

utic

a in

Hid

algo

(190

4-19

05),

syno

nym

y af

ter R

osew

ater

(196

5).

50) C

lass

ified

as C

orbi

s in

Hid

algo

(190

4-19

05).

51) A

s Luc

ina

fibul

a in

Hid

algo

(190

4-19

05),

syno

nym

y af

ter L

ampr

ell &

Whi

tehe

ad (1

992)

.52

) Cla

ssifi

ed a

s Luc

ina

in H

idal

go (1

904-

1905

).53

) Hid

algo

’s (1

904-

1905

) rec

ord

of P

erna

ano

mio

ides

may

per

tain

to th

is sp

ecie

s, or

to Is

ogno

mon

per

na.

54) C

lass

ified

as C

hlam

ys in

Hid

algo

(190

4-19

05).

55) C

lass

ified

as M

odio

la st

riat

ula

in H

idal

go (1

904-

1905

).56

) As A

rca

volu

cris

in H

idal

go (1

904-

1905

), sy

nony

my

afte

r Lam

prel

l & H

ealy

(199

8).

57) H

idal

go’s

(190

4-19

05) r

ecor

d of

Ven

eric

ardi

a ca

nalic

ulat

a R

eeve

nee

ds to

be

verif

ied

and

is n

ot in

clud

ed.

58) R

aine

s (19

96) a

lso

reco

rds S

emip

alliu

m ti

gris

from

Gua

m, b

ut I

have

nev

er se

en a

ny sp

ecim

ens o

f tha

t spe

cies

on

the

isla

nd.

Marine Bivalvia (Mollusca) of Guam

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Transcript of Marine Bivalvia (Mollusca) of Guam