Marine Bivalvia (Mollusca) of Guam
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Transcript of 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:
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
Paulay: Bivalves of Guam 221
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
222 Micronesica 35-36, 2003
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
Paulay: Bivalves of Guam 223
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.
224 Micronesica 35-36, 2003
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.
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Received 1 October 2001
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F 29
7252
P
2M
uscu
lus s
p. 2
U
F 28
7441
P
1M
uscu
lus s
p. 3
U
F 29
7079
P
2Rh
ombo
idel
la m
alac
cana
Ock
elm
ann,
198
3U
F 28
7994
P
1Se
ptife
r bilo
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.