mollusks of the northern mariana islands, with special reference to the
DESCRIPTION
selectivity of oceanic dispersal barriers - Micronesica Vol. 19 No. 1-2 Dec., 1983 - By: Vermeij, G.J., Kay, E.A., and Eldredge, L.G.TRANSCRIPT
Molluscs of the Northern Mariana Islands, With Special Reference to the Selectivity
of Oceanic Dispersal Barriers1
GEERAT J. VERMEIJ
Department of Zoology, University of Maryland, College Park, Maryland 20742
E. ALISON KAY
Department of Zoology, University of Hawaii, Honolulu, Hawaii 96822
LUCIUS G. ELDREDGE
University of Guam Marine Laboratory, UOG Station, Mangilao, Guam 96913
Abstract- The shelled molluscan fauna of the Northern Marianas, a chain of volcanic islands in the tropical western Pacific, consists of at least 300 species. Of these, 18 are unknown from or are very rare in the biologically better known southern Marianas. These northern-restricted species are over-represented among limpets and in the middle to high intertidal zones of the northern Marianas. At least 22 gastropods which are common in the intertidal zone and on reef flats of the southern Marianas are absent in the northern Marianas.
The northern Marianas lie within the presumed source area of the planktonically derived part of the Hawaiian marine fauna. The ocean barrier between the northern Marianas and the Hawaiian chain appears to select against archaeogastropods and against intertidal species but is unselective with respect to adult size and to other aspects of gastropod shell architecture. These findings are consistent with those for other dispersal barriers.
Introduction
The Mariana Islands form the southern part of an island chain which extends northward through the Bonin, Volcano, and lzu Islands to central Honshu, Japan. Whereas the marine biota of the southern Marianas (Guam, Rota, Tinian, and Saipan) is becoming relatively well known, that of the northern Marianas (Fig. 1) is largely unstudied. Besides a few scattered records, the only published marine biological accounts are those of Tsuda and Tobias (1977a, b), who provided a checklist of algae.
The taxonomic composition and biogeographical affinities of the marine fauna of the northern Marianas are interesting for at least three reasons. First, the shores of the northern islands are almost entirely volcanic and might be expected to yield species that are absent from the raised limestone and associated reef-flat shorelines of the southern Marianas. Second, the Marianas lie in what has traditionally been
1 Contribution No. 196, University of Guam Marine Laboratory. Micronesica 19(1- 2): 27- 55, 1983 (December).
Vol. 19. December 1983 29
plate on which most of the islands of the Pacific lie, by the deep Marianas Trench, and thus are a test area for some of the biogeographical hypotheses recently proposed by Kay (1980a, 1984) and Springer (1982). ,
The molluscan fauna of the northern Marianas especially invites comparison with that of the Hawaiian Islands which, like the northern Marianas, are largely volcanic. The Hawaiian chain consists of a series of 26 islands extending from Hawaii in the southeast, the youngest at - 1 my, to Kure, an atoll resting on an ancient volcanic pedestal to the northwest. Dating of basalt at Midway near Kure indicates that the island was above the sea in the Miocene, 30 my ago (Ladd et al. 1970). The age sequence in the Mariana Islands is opposite that in Hawaii; Guam and Saipan in the south are the oldest islands and much of their volcanic base is capped with Pliocene and Pleistocene limestone. The northern arc of islands, from Anatahan to Uracas, is a series of young volcanic islands, and Pagan, Asuncion, and Uracas are all active volcanoes.
The subtropical countercurrent, which arises in the west as an offshoot from the Kuroshio Current, flows eastward through the northern Marianas and continues erratically eastward to Johnston Island and thence to the northwestern Hawaiian islands. (For discussions of the biogeographical affinities of the Hawaiian Islands see Kay, 1967, 1980a; and Grigg, 1981.) The fauna of the northern Marianas thus affords the opportunity to examine the selectivity of the dispersal filter between the Hawaiian biota and its presumed source in the northwestern Pacific.
Very little is currently known about the ecological and architectural differences between widely dispersing and poorly dispersing species. The topic is important because wide dispersal is assoCiated with long-term resistance to extinction (Stanley, 1979). If statistical associations exist between dispersability and biotic competence, long-term evolutionary trends toward an increase in biotic competence might be tied to the evolution of large-scale dispersal by planktonic stages.
Materials and Methods
The material reported on results from several expeditions to the northern Marianas organized by the University of Guam Marine Laboratory between 1972 and 1981. All three of us have made field observations in the northern Marianas and all nine islands have been visited by one or another ofus.Shallow-water habitats were sampled by wading, snorkeling, and scuba diving. Sediment samples collected from tidepools and offshore to depths of 30m were examined for micromolluscs (molluscs less than 7 mm in greatest dimension) and quantitative analyses were performed following the method of Kay (1980b). We have also examined the collections made by C. Carlson and P. J. Hoff aboard the WANDERER and those of the National Museum of Natural History, Smithsonian Institution (USNMNH).
The shelled molluscs reported on are deposited in the University of Guam Marine Laboratory collection, in the Vermeij collections, and in the Kay collections at the University of Hawaii.
30 Micronesica
Only species with shells greater than about I 0 mm in greatest dimension are used for the analyses of architectural differences among shells. Thick -lipped shells are those which have an expanded or thickened rim of the aperture in the adult stage; thinlipped shells are those which lack this modification. Species referred to the intertidal are those found in the intertidal and include species which occur both in the intertidal and the subtidal.
Results
Northern Mariana-Southern Mariana Comparison. A list of species and their occurrence on the nine northern Mariana Islands is
presented in Appendix I. We have excluded small alga-dwellers such as the Eatoniellidae and Cingulopsidae and the sessile Vermetidae (except for Serpulorbis variabilis) from our list, and our inventory of the fauna is certain to be incomplete in other respects. However, it shows clearly that several species which are abundant on the shores of Guam and others of the southern Marianas are conspicuously rare or absent in the northern Marianas and vice versa.
Twenty-two common shallow-water, hard-bottom molluscs from Guam which are absent from the northern islands are listed in Appendix 2. Eight (36%) are confined to the middle or high intertidal zones, and I4 (64%) are found chiefly or exclusively on reef flats . It is interesting that Trochus niloticus has not been found in the northern Marianas. This species was introduced from Belau (formerly Palau) to Saipan in the late I930s and to Guam about two decades later (Eldredge, in press). It is now abundant on reef-flats and subtidally to a depth of at least I2 m (B. Smith, pers. comm.) on both Saipan and Guam.
Several other species which are abundant on Guam are found only sporadically in the northern Marianas; others may have been erroneously recorded from these islands. Only one specimen of Cerithium columna was found in the northern islands (at Pagan in I98I), and it differs from the abundant intertidal form at Guam by having much finer sculpture. Seven species in the USNMNH collections made by U.S. Armed Forces in I945 (Appendix I) have not been found during the University of Guam Marine Laboratory expeditions, and the locality data accompanying the specimens may be erroneous.1 We have identified I8 species from the northern Marianas which are absent or very rare in the biologically better known southern
1 A. H. Banner (pers. comm. to GJV, 28 June 1984) tells us that molluscs which he collected at Agrihan and Maug in 1945 were very likely intermixed with material taken at Saipan when a large military truck drove accidentally over the collected specimens. Although he attempted to salvage the remains, . Banner cannot guarantee the accuracy of the locality data of the molluscs, which were subsequently deposited at the USNMNH. Species which we believe are erroneously recorded from the northern islands in this way include Cerithium nodulosum, Cymatium nicobaricum, C. muricinum, Chicoreus brunneus, Cantharus fumosus, Nebularia chrysalis, and Vasum turbinellus. All these species are common on the reef flats of the southern Marianas including Saipan. We also doubt the record of Trochus maculatus, a species probably absent from the southern Marianas.
Vol. 19. December 1983 31
Marianas (Appendix 1). Although Pusia sp. may be easily overlooked because their shells are less than
about 10 mm in greatest dimension, the other species have prominent shells, and we do not believe their absence or rarity on Guam is an artifact of co!J'ection. Additionally, several northern Marianas species are reported from Guam but have not been found in recent years. Haliotis ovina and H . varia varia were recorded by Talmadge (1963). H. ovina is represented in the USNMNH collections by one specimen from Saipan. Nodilittorina pyramidalis is recorded by Rosewater (1970) from "Apra Bay" (Guam) and from Saipan, but we have not found it in the southern Marianas. Planaxis niger, abundant in the northern Marianas, was collected from northern Saipan (Kropp, pers. comm.), but no other southern Marianas record of this species is known.
Most (66 %) of the northern-restricted species were found on two or more of the nine islands (Appendix 1). They are particularly well represented in the intertidal zone. Of the 35 gastropod species found in or above the middle intertidal barnacle (Tetraclita) zone, 10 (29 %) belong to the northern-restricted group. An additional species, Nerita guamensis, occurs at two of the northern islands as a dark form which is unknown from the southern Marianas. Species with a limpet-like form comprise 8% of the northern Marianas gastropod fauna as a whole but 29%of the northernrestricted species.
Northern-restricted species are not confined to the Mollusca. Among corals, Leptastrea sp. and Millepora foveolata are known from Pagan but not from the southern Marianas (R. H. Randall, pers. comm.). At least two porcellanid crabs are northern-restricted (R. K. Kropp, in prep.). The high intertidal hermit crab Calcinus seurati has been found at Pagan and Saipan but is unknown from Guam (D. S. Wooster, pers. comm.). The fishes Labroides pectoralis and Pseudodax molluccanus are also absent from Guam but have been seen at Pagan (R. Myers, pers. comm.). Randall (1982) reports that the surgeon fish Acanthurus leucopareius, rare at Guam, is common in the northern Marianas. Algae found in the northern but not the southern Marianas include the phaeophytes Dictyota hamifera, Hormophyta triquetra, and Stipopodium hawaiiense (Tsuda and Tobias, 1977a; Eldredge et a!., 1977) and the rhodophytes Dermonema frappieri, Polysiphonia sacchorhiza, Laurencia succisa, L . surculifera, Galaxaura filamentosa, G. hystrix, and G. veprecula (Tsuda and Tobias, 1977b; Itono, 1980).
Northern Marianas-Hawaiian Comparison. Data comparing the molluscan faunas of the northern Marianas and the
Hawaiian Islands are set out in Tables 1-3. The northern Marianas Archaeogastropoda (superfamilies Pleurotomariacea, Fissurellacea, Patellacea, and Trochacea) are less well represented in the Hawaiian Islands than are the mesogastropods and neogastropods (Table 1). Indeed, two families of archaeogastropods, the Haliotidae and Acmaeidae, which contribute 17% of the archaeogastropod species to the northern Marianas fauna, are not present at all in Hawaii. The effect of this
32 Micronesica
Table I. Hawaiian affinities of the northern Marianas fauna.
Taxonomic groups Archaeogastropoda Neritacea Mesogastropoda Neogastropoda Euthyneura Polyplacophora Bivalvia
Architectural groups * Gastropods, narrow
aperture * Gastropods, thick
or expanded lip * Gastropods, thin lip * Gastropods, large
(>50mm) * Gastropods, small
(> !Omm) Habitat groups
Gastropods, middle to high intertidal
Gastropods, lower shore, hard bottom
Gastropods, soft bottom Bivalves, soft bottom
No. of species in northern Marianas
41 8
87 Ill 21
2 39
57
72
107
37
53
35
122
56 13
*Comparison excluding Archaeogastropoda
%Occurring in Hawaii
24 50 70 59 33 0
46
58
68
55
51
53
51
60
52 54
Table 2. Numbers of intertidal mollusc species recorded in the northern Marianas and in Hawaii (Hawaiian data from Kay, 1979).
Northern Marianas Hawaii
Archaeogastropods 7 (4 families, 3 (I family, 6 genera) I genus
all endemic) Neritidae 6 4 Littorinidae 6 4 (I endemic) Planaxidae 2 2 Muricacea 8 9 (2 endemic) Mitracea I 2 Smaragdinellidae I I Melampidae 2 6 (2 possibly
endemic) Siphonariidae 2 2 Bivalvia 0 3
Vol. 19. December 1983
Table 3. Numbers of samples and percent species composition of major components of micromolluscan assemblages in the Northern Marianas
and in the Hawaiian Islands.
N. Marianas Hawaii 1. 1 Oahu J.2 Niihau3
No. of samples 19 9 11 6
Number shells 3,053 3,032 3,978 2,705 Percent Composition
Archaeogastropoda5 18 7 3 9
Rissoidae 4 23 17 27 Cerithiidae 19 3 4 7 Dialidae >I 4 31 9
' Samples from 3- lOm depth transects on subtidal reef, Kona, Hawaii 2 Samples from !Om depth on transects at Waianae, Oahu 3 Samples from 3- 10m depth on transects on the lee shore of Niihau 4 French Frigate Shoals. Samples from 8 m depth s Archaeogastropoda do not include Tricolia variabilis.
33
F. F. S.4
11 2,334
II 18 13 15
relative lack of archaeogastropods is reflected in two comparisons (Tables 2 and 3). When the numbers of species found in the intertidal in the northern Marianas and the Hawaiian Islands are compared, about the same numbers are found in both areas, but whereas seven species of archaeogastropods occur in the intertidal of the northern Marianas only three archaeogastropod species occur in that zone in Hawaii, all in the same genus (Cellana) and all endemic. The numbers of species in other groups are similar for the two island groups, i.e. , there are four or five species of nerites, five or six species of littorines, and eight or nine species of muricaceans.
When the relative proportions of micromolluscs from subtidal sediment samples (Table 3) are compared, 18 % of the micromolluscs in the samples from the northern Marianas are archaeogastropods (exclusive of Tricolia variabilis which is highly variable in numbers both in the northern Mariana and Hawaiian Islands) while in the Hawaiian Islands the figure ranges from 3 to 11 %. Rissoids comprise a low proportion of the shells in the northern Marianas but a higher proportion in the Hawaiian Islands, and cerithiids which are also prominent in the northern Marianas (they comprise about 19% of the fauna) are less prominent in Hawaii.
The northern Marianas and Hawaiian marine molluscan faunas are similar in gastropod: bivalve proportions-88: 12 and 88 : 17, respectively, and the figures for both island groups are consistent with those reported for other islands (Kay, 1967). Soft-bottom and hard-bottom species from the lower shore do not differ in their relative proportions (Table 1). The Hawaiian marine molluscan fauna differs from that of the northern Marianas, however, in its relatively high percentage of endemism at the species level (about 15%, Kay, 1979). There is apparently little or no endemism at this level in the northern Marianas. Several species in the Hawaiian fauna are widespread in the Pacific Ocean and have been termed Pacific plate species (cf. Springer, 1981, 1982; Kay, 1980a, 1984) because they appear to be restricted to the
34 Micronesica
Pacific plate. In molluscan groups for which there is good distributional information, such as the cowries and strombids, more than 25% of the species in Hawaii are Pacific plate species. We have detected only one cowrie-Cypraea maculifera which is predominantly Pacific plate species in the northern Marianas.
With respect to gastropod antipredator shell architecture, mesogastropods, neogastropods, and euthyneura with a narrowly elongated aperture are found in about the same proportion in the northern Mariana Islands and the Hawaiian Islands as are member of these groups with a broad aperture. Furthermore, there are no differences in the proportion of thick-lipped and thin-lipped gastropods exclusive of the Archaeogastropoda (that is, the Neritacea, Mesogastropoda, Neogastropoda, and Euthyneura) in the northern Mariana and Hawaiian Islands. Had we compared the distribution of thick-lipped members of these groups to all thin-lipped gastropods rather than excluding the archaeogastropods, then more thick-lipped, narrowapertured species would appear. Gastropods of large size (more than 5 em maximum dimension) are found in Hawaii in the same proportion as are smaller-shelled species (Table 1).
Discussion
The presence/absence and the distribution of molluscs within the northern Marianas are affected by at least three factors: collecting effort, types of habitat, and an island effect, as shown by patchy distributions. More visits were made to Pagan for which the largest number of species was recorded (230) than to other islands of the northern Marianas, and fewer visits were made to Uracas. Pagan is also the most geologically diverse of the islands in the chain, with black sand beaches, raised limestone benches, and subtidal coral communities. Uracas is characterized by a foreshore of basalt boulders and is surrounded by warm sulphurous water. Maug, composed of three weathered volcanic peaks which surround the old crater may have the most extensive coral reef development among the islands. Finally, species presence or absence on all the islands may be the result of an island effect in that the presence or absence of species can be the result of chance. Patchy distributions on islands have been documented for the Line Islands (Kay, 1971) and the Hawaiian Islands (Kay, 1979; 1980a).
Of the occurrences of several species which are abundant on the shores of Guam and others of the southern Marianas but which are absent or rare in the northern Marianas, some are explicable. The absence of species that inhabit seagrass meadows and mangroves is expected because these habitats are not represented in the northern Marianas. For other shallow-water species, however, absence in the north is more puzzling. We rather expected the southern-restricted species to be confined to limestone surfaces and to be absent from volcanic substrates. With the possible exception of the limestone-restricted limpet Patelloida sp., however, the southernrestricted species are known from both limestone and volcanic shores. The 22 common shallaw-water hard-bottom molluscs from Guam which are absent from the
Vol. 19. December 1983 35
northern islands are ecologically and biogeo graphically unrepresentative of the fauna as a whole, with over a third of the species confined to the middle and high intertidal zones, and the other two-thirds found on reef flats.
The northern-restricted species are also ecologically, morphologically, and biogeographically atypical of the northern Marianas fauna as a whole, and a high proportion of intertidal and limpet-like forms also characterizes the northern Marianas forms. Additionally, many of the northern-restricted species appear to be confined to high islands and to be absent from atolls. Usillafusconigra, for example, is known from Okinawa, Samoa, the Marquesas, and the Hawaiian Islands, but has not been collected from atolls in the Marshalls, Societies, or Tuamotus (data from USNMNH, British Museum (Natural History), B. P. Bishop Museum). Other examples of high island species include Cellana toreuma, the three acmaeids, Nodilittorina pyramidalis, and Planaxis niger. Some or all of these species may be restricted to volcanic substrates.
The most obvious difference between the Hawaiian fauna and that of the northern Marianas is in the proportion of archaeogastropods, both in terms of species composition and in terms of absolute numbers. The lesser proportion of archaeogastropods in the Hawaiian fauna as compared with that of the northern Marianas is consistent with what is known of the larval life history of these gastropods: they tend to lack widely dispersing teleplanic larval stages (Kay, 1984) which may prevent them from reaching the Hawaiian chain from the west. Mesogastropods and neogastropods have teleplanic larvae which are taxonomically widespread and therefore are expected in larger proportion. There are, however, some anomalies in the dispersal pattern: for example, only three of the southernrestricted species-Hipponix foliaceus, Cymatium gemmatum, and Mitra cucumerina - - 14% of the southern-restricted species are also found in Hawaii and only one of the common northern-restricted species ( Usilla fusconigra) is also known from Hawaii.
Other studies of selectivity of dispersal barriers in the ocean have yielded results similar to those we report here. The barrier between the Line Islands in the west and the offshore islands of the eastern Pacific in the east is unselective with respect to apertural shape and shell size (Vermeij , 1978), but there is a selective diminution of numbers of species with apertural modification between the eastern Pacific islands and the mainland coast of western tropical America (Vermeij, 1978). This latter conclusion, however, rests on a small sample, because most of the Indo-west Pacific elements which extend as far east as the offshore islands do not also occur on the mainland (see also Emerson, 1978; Zinsmeister and Emerson, 1979). The selectivity of the Suez Canal as a barrier remains a matter of dispute, but Vermeij's (1978) analysis suggests that there is no strong reduction of any one architectural group of species between the Red Sea and the Mediterranean through the Canal. On the other hand, upper shore species have perhaps been less prone to migration through the Suez Canal from the Red Sea to the Mediterranean than have lower shore species (Safriel and Lipkin, 1975; Vermeij, 1978).
36 Micronesica
The biogeographical distinctiveness of the gastropods of the upper shore in the northern Marianas recalls the upper shore endemism in the Hawaiian Islands, although endemism per se is not pronounced in the northern Marianas. Similar situations have been reported in the southern tropical Atlantic, western Indian Ocean, and Red Sea (Vermeij, 1972, 1978). The data from the present study suggest that poor dispersability and effective adaptation to locally distinctive environments contribute to the endemism of the high intertidal biota.
Finally, we wish to emphasize that, although faunal surveys have often been maligned as being superficial descriptive studies, they are essential for the understanding of biogeographical affinities, selectivity of barriers, and patterns of diversity. We believe that a comparison of the tropical East Asian mainland biota with that of the northern Marianas will shed further light on the nature of the open-ocean dispersal barrier.
ACKNOWLEDGMENTS
We thank the Office of Coastal Resources Management, Commonwealth of the Northern Mariana Islands (especially J. Elemato, Manager, and B. Aldan, Coastal Coordinator for Pagan) for support to visit Pagan in March and July, 1981. The Governor's Office further cleared the way for our visits. Mary and John Laverny, owners of the SJ-TI-SJ, enabled us to visit several of the islands in July 1981. The Marvin J. Coles Memorial Fund of the Guam Oil and Refining Company, the National Park Service, the National Sea Grant program, and the National Science Foundation, Institutional Grant for Science at the University of Guam, and the Program of Biological Oceanography grant to G. J . Vermeij, are gratefully acknowledged for funding the work reported here. Field assistance was provided by B. Best, R. E. Dickinson, M. J . Gawel, and R. K. Kropp. We thank W. K. Emerson, R. S. Houbrick, D. Lindberg, J. Rosewater, and R. Salisbury for confirming several identifications and B. D. Smith for critical review of the manuscript. We are also grateful to the United States National Museum (Natural History), the British Museum (Natural History), and the Bernice P. Bishop Museum for use of their collections.
References Cited
Abbott, R. T . 1960. The genus Strombus in the Indo-Pacific. Indo-Pacific Mollusca 1(2): 33- 146. Cernohorsky, W. 0 . 1976. The Mitridae of the world. Part I. The subfamily Mitrinae. Indo-Pacific
Mollusca 3(17): 273- 528. Ekman, S. 1953. Zoogeography of the sea. Sidgwick & Jackson, London. 417 p. Eldredge, L. G. 1983. Summary of environmental and fishing information on Guam and the
Commonwealth of the Northern Mariana Islands: historical background, description of the islands, and review of the climate, oceanography, and submarine topography. NOAA Tech. Memo. NMFS 40. 181 p.
- - -. in press/ Coral reef alien species. In Human activities causing damage to coral reefs: knowledge and recommendations. UNESCO.
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aii
Ast
raea
rho
dost
oma
X
X
X
(Lam
arck
) T
urbo
arg
yros
tom
us
X
X
X
X
Sar
igan
Lin
ne
Ala
mag
an
Agr
ihan
T. p
etho
latu
s L
inne
X
X
X
T. s
etos
us G
mel
in
X
X
X
X
X
Agr
ihan
NE
RIT
AC
EA
N
erit
a al
bici
lla L
inne
X
In
tert
idal
; al
so
in H
awai
i
N.
guam
ensi
s Q
uoy
X
Sar
igan
In
tert
idal
; da
rk
<
and
Gai
mar
d fo
rm o
nly
~
N.
insc
u/pt
a R
eclu
z X
In
tert
idal
;o
N.
max
ima
Gm
elin
X
In
tert
idal
I:
)
N.
plic
ata
Lin
ne
X
X
X
X
Sar
igan
In
tert
idal
; al
so
§ 3 A
lam
agan
in
Haw
aii
but
r::T <>
rare
exc
ept
in
.... :c th
e N
orth
wes
t 0
0
Isla
nds
...,
N. p
olit
a L
inne
X
In
tert
idal
; al
so
in H
awai
i
Pup
erit
a be
nson
i (R
eclu
z)
X
Ner
itop
sis
radu
la (
Lin
ne)
X
X
Als
o in
Haw
aii
ME
SO
GA
ST
RO
PO
DA
L
itto
rina
coc
cine
a X
X
S
arig
an
Inte
rtid
al;
also
(Gm
elin
) in
Haw
aii
but
rare
L.
pint
ado
Woo
d X
X
X
X
X
X
S
arig
an
Inte
rtid
al;
also
A
grih
an
in H
awai
i (R
osew
ater
19
70)
..., -c
App
endi
x. (
cont
inue
d)
"""
0
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
L.
undu
lata
Gra
y X
In
tert
idal
; al
so
in H
awai
i b
ut
rare
N
.f;di
litto
rina
mil
legr
ana
X
Inte
rtid
al
(Phi
lipp
i)
* N.
pyra
mid
alis
X
X
X
X
In
tert
idal
(Q
uoy
and
Gai
mar
d)
* N.
quad
rici
ncta
feej
eens
is
X
X
X
X
X
Inte
rtid
al;
see
(Ree
ve)
Ros
ewat
er a
nd
Kad
olsk
y 19
81 f
or
nom
encl
atur
e R
isso
ina
ambi
gua
(Gou
ld)
X
X
X
Als
o in
Haw
aii
a:: R
. m
ilto
zona
Tom
lin
X
X
X
X
Als
o in
Haw
aii
;:;·
R.
turr
icul
a P
ease
A
lso
in H
awai
i ...
X
X
X
X
0
Schw
artz
iell
a gr
acili
s 1:1
X
A
lso
in H
awai
i " "'
. (P
ease
) ;:;
· "' Sa
nson
ia s
p.
X
Par
ashi
ela
beet
si L
add
X
Als
o in
Haw
aii
Pyr
amid
ello
ides
sut
a X
A
lso
in H
awai
i (P
ilsb
ry)
Zeb
ina
imbr
icat
a (G
ould
) X
A
lso
in H
awai
i Z
. tr
iden
tata
(M
icha
ud)
X
X
X
Als
o in
Haw
aii
Bro
okul
a cf
. B
. ik
i K
ay
X
Als
o in
Haw
aii
Leu
corh
ynch
ia c
f. L.
cro
ssei
X
(Tyr
on)
Lio
tina
loc
ulos
a (G
ould
) X
Van
ikor
o ac
uta
(Rec
luz)
X
A
lso
in H
awai
i V
. ca
ncel
lata
(L
amar
ck)
X
Als
o in
Haw
aii
V. d
ista
ns (
Rec
luz)
X
Sabi
a co
nica
X
X
X
X
X
A
grih
an
Als
o in
Haw
aii
(Sch
umac
her)
(U
SN
MN
H)
* Pla
naxi
s ni
ger
X
X
X
X
Sar
igan
In
tert
idal
(Q
uoy
and
Gai
mar
d)
Ala
mag
an
P.
zona
tus
A. A
dam
s X
X
X
X
X
A
lam
agan
In
tert
idal
Lop
ohoc
och/
ias
X
X
Als
o in
Haw
aii
min
utis
sim
us (
Pil
sbry
) O
mal
ogyr
a ja
poni
ca
X
X
Als
o in
Haw
aii
(Hab
e)
Cyc
lost
rem
iscu
s em
eryi
X
X
X
X
A
lso
in H
awai
i
(Lad
d)
Orb
ites
tell
a re
gina
Kay
X
X
X
A
lso
in H
awai
i C
aec
um a
rcua
tum
de
Fol
in
X
X
X
Als
o in
Haw
aii
C.
oahu
ense
(P
ilsb
ry)
X
X
Als
o in
Haw
aii
C.
sepi
men
tum
de
Fol
in
X
X
X
Als
o in
Haw
aii
C.
cf.
C.
glab
el/u
m A
. Ada
ms
X
Als
o in
Haw
aii
< ~
Mod
ulus
tec
tum
(G
mel
in)
X
Als
o in
Haw
aii
-B
itti
um z
ebru
m (
Kie
ner)
X
X
X
X
A
lso
in H
awai
i ~
B. p
arcu
m (
Gou
ld)
X
X
X
X
Als
o in
Haw
aii
0 2 S
erpu
lorb
is v
aria
bilis
X
X
X
In
tert
idal
; al
so
"' 9 H
adfi
eld
and
Kay
in
Haw
aii
cr "'
Cer
ithi
um a
trom
argi
natu
m
X
Als
o in
Haw
aii
.... -D
autz
enbe
rg a
nd B
ouge
"' 00 w
C
. co
lum
na S
ower
by
X
Als
o in
Haw
aii
C.
inte
rstr
iatu
m S
ower
by
X
X
Als
o in
Haw
aii
C.
egen
um G
ould
X
X
X
X
A
lam
agan
A
lso
in h
awai
i A
grih
an
C.
mut
atum
Sow
erby
X
X
X
X
A
lam
agan
A
lso
in H
awai
i A
grih
an
C.
nesi
otic
um P
ilsb
ry a
nd
X
X
X
Als
o in
Haw
aii
Van
atta
C
. no
dulo
sum
Bru
guii:
re
X
(US
NM
NH
)
C.
cf.
C.
rubu
s D
esha
yes
Agr
ihan
Se
e R
ehde
r ( 1
980)
(U
SN
MN
H)
for
disc
ussi
on o
f th
is s
peci
es
:::
App
endi
x. (
cont
inue
d)
"""
N
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
Rhi
nocl
avis
asp
era
X
(Lin
ne)
R.
fasc
iata
(B
rugu
iere
) X
X
X
A
lso
in H
awai
i
R\
sine
nsis
(G
mel
in)
X
X
X
Sar
igan
L
arge
for
m o
nly;
al
so i
n H
awai
i
Cer
ithi
dium
per
parv
ulum
X
X
X
A
lso
in H
awai
i
(Wat
son)
C
erit
hidi
um s
p.
X
X
Dia
la j
lam
mea
(Pe
ase)
X
Fin
ella
pup
oide
s A
. A
dam
s X
X
X
A
lso
in H
awai
i
Scal
iola
sp.
X
A
lso
in H
awai
i
Dia
lida
e cf
. A
/aba
sp.
X
Stro
mbu
s de
ntat
us L
inne
X
~
S.
mic
rour
ceus
Kir
a X
A
grih
an
c;·
.... (A
bbot
t 0 ::s "
1960
) "' c;·
!>
l S
. m
utab
ilis
Sw
ains
on
X
S. p
lica
tus
pulc
hell
us
X
Ree
ve
(Abb
ott
1960
) L
ambi
s ch
irag
ra c
hira
gra
X
(Lin
ne)
L. t
runc
ata
seba
e X
X
(Kie
ner)
T
ereb
el/u
m t
ereb
el/u
m
X
(Lin
ne)
Pol
inic
es m
elan
osto
mus
X
A
lso
in H
awai
i
(Gm
elin
) (U
SN
MN
H)
Nat
icid
ae s
p.
X
Lac
hrym
a su
/cif
era
X
X
(Sow
erby
)
Cyp
raea
cap
utse
rpen
tis
X
X
X
X
X
X
Sar
igan
A
lso
in H
awai
i L
inne
A
lam
agan
, A
grih
an
C.
cam
eo/a
Lin
ne
X
X
X
Agr
ihan
A
lso
in H
awai
i C
. de
pres
sa G
ray
X
C.
eros
a L
inne
X
X
X
E
xtre
mel
y ra
re i
n (U
SN
MN
H)
Haw
aii
C.
fim
bria
ta G
mel
in
X
Als
o in
Haw
aii
C.
g/ob
ulus
Lin
ne
X
X
Als
o in
Haw
aii
C.
he/v
ola
Lin
ne
X
X
X
X
Als
o in
Haw
aii
C.
isab
ella
Lin
ne
X
X
X
X
X
Sar
igan
A
lso
in H
awai
i C
. ly
nx
Lin
ne
X
X
Ala
mag
an
Ext
rem
ely
rare
in
(US
NM
NH
) H
awai
i C
. m
acu/
ifer
a S
chil
der
X
X
X
X
Als
o in
Haw
aii
<
~
C.
mau
riti
ana
Lin
ne
X
X
X
Ala
mag
an
Als
o in
Haw
aii
(US
NM
NH
) A
grih
an
~
(US
NM
NH
) t:
)
C.
mon
eta
Lin
ne
X
X
X
X
Ala
mag
an
Als
o in
Haw
aii
~ 3 C
. po
rari
a L
inne
X
X
X
X
X
A
lam
agan
A
lso
in H
awai
i g"
A
grih
an
... ~
Sar
igan
0
0
(US
NM
NH
) ....
C.
talp
a L
inne
X
A
lso
in H
awai
i C
. te
stud
inar
ia L
inne
X
C.
tere
s G
mel
in
X
Als
o in
Haw
aii
C.
vite
llus
Lin
ne
X
Als
o in
Haw
aii
Cas
sis
corn
uta
Lin
ne
X
X
Als
o in
Haw
aii
Cas
mar
ia p
onde
rosa
X
(Gm
elin
) C
ymat
ium
hep
atic
um
X
Als
o in
Haw
aii
(Rod
ing)
C
. m
uric
inum
(R
odin
g)
X
Als
o in
Haw
aii
(US
NM
NH
) C
. ni
coba
ricu
m
Agr
ihan
A
lso
in H
awai
i (R
odin
g)
(US
NM
NH
) ... ....
App
endi
x. (
cont
inue
d)
""" """
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
C.
pile
are
(Lin
ne)
X
Agr
ihan
A
lso
in H
awai
i (U
SN
MN
H)
B\s
a b
ufon
ia (
Gm
elin
) X
X
A
lso
in H
awai
i B
. gr
anul
aris
(R
odin
g)
X
X
Als
o in
Haw
aii
NE
OG
AS
TR
OP
OD
A
M u
rico
drup
a fu
nicu
lus
X
Als
o in
Haw
aii
(Woo
d)
Tha
is a
rmig
era
(Lin
k)
X
X
X
X
Inte
rtid
al;
also
in
Haw
aii
T.
tube
rosa
(R
odin
g)
X
X
T. i
nter
med
ia (
Kie
ner)
X
X
X
X
In
tert
idal
; al
so
(US
NM
NH
) in
Haw
aii
::: c;·
Pur
pura
per
sica
(L
inne
) X
X
X
X
X
In
tert
idal
.... 0
Mor
ula
bico
nica
X
X
X
::s ,. "'
(Bla
in vi
lle)
c;· ""
M.
gran
ulat
a (D
uclo
s)
X
X
X
X
Inte
rtid
al;
also
in
Haw
aii
M.
spin
osa
X
Als
o in
Haw
aii
(H.
& A
. Ada
ms)
M
. uv
a (R
odin
g)
X
X
X
X
X
Agr
ihan
In
tert
idal
; al
so
in H
awai
i D
rupe
lla
elat
a X
X
A
lso
in H
awai
i (B
lain
ville
) (U
SN
MN
H)
D.
ochr
osto
ma
X
Agr
ihan
A
lso
in H
awai
i (B
lain
ville
) (U
SN
MN
H)
Dru
pa a
rach
noid
es
X
X
X
X
Sar
igan
A
lso
in H
awai
i; w
e (L
amar
ck)
rega
rd t
his
as a
di
stin
ct s
peci
es
alth
ough
it
is
not
reco
gniz
ed a
s
such
by
Em
erso
n an
d C
erno
hors
ky
(197
3)
D.
clat
hrat
a cl
athr
ata
X
X
X
Mau
g re
cord
tha
t (L
amar
ck)
of
Em
erso
n an
d C
erno
hors
ky
(197
3)
D.
gros
sula
ria
(Rod
ing)
X
X
A
grih
an
Als
o in
Haw
aii
but
alm
ost
enti
rely
re
stri
cted
to
the
Nor
thw
est
Haw
aiia
n Is
land
s D
. m
orum
mor
um
X
X
X
X
Ala
mag
an
Inte
rtid
al;
also
(R
odin
g)
Agr
ihan
in
Haw
aii
<
~
D.
rici
nus
(Lin
ne)
X
X
X
X
X
Inte
rtid
al;
also
in
Haw
aii
~
D.
rubu
sida
eus
(Rod
ing)
X
X
X
X
In
tert
idal
; al
so
0 ~ in
Haw
aii
8 N
assa
ser
ta
X
Als
o in
Haw
aii
cr ..
(Bru
guie
re)
.... -M
acu/
otri
ton
brac
teat
a A
lso
in H
awai
i \0
X
X
0
0
w
(Hin
ds)
*Usi
llaf
usco
nigr
a (P
ease
) X
X
In
tert
idal
; al
so
in H
awai
i C
hico
reus
bru
nneu
s X
(Lin
k)
(US
NM
NH
) C
oral
liop
hila
vio
lace
a X
X
A
lso
in H
awai
i (K
iene
r)
Quo
yu/a
mon
odon
ta
X
Agr
ihan
A
lso
in H
awai
i (B
lain
ville
) C
aduc
ifer
dec
apit
ata
X
Als
o in
Haw
aii
fusc
omac
ulat
ti (
Peas
e)
C.
iost
omus
(G
ray
in
X
Als
o in
Haw
aii
Gri
ffit
hs a
nd P
idgi
n)
.;.
v.
App
endi
x.
(con
tinu
ed)
.,. a-
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
Pis
ania
ign
ea (
Gm
elin
) X
A
lso
in H
awai
i C
anth
arus
fum
osus
X
\(D
illw
yn)
(US
NM
NH
) C
. un
dosu
s (L
inne
) X
X
X
X
(US
NM
NH
) N
assa
rius
gau
dios
us
X
Agr
ihan
A
lso
in H
awai
i (H
inds
) N
. pap
illo
sus
(Lin
ne)
X
Als
o in
Haw
aii
N. p
aupe
rus
(Gou
ld)
X
X
X
Als
o in
Haw
aii
Eup
lica
des
haye
si
X
(Cro
sse)
E
. tu
rtur
ina
(Lam
arck
) X
A
lso
in H
awai
i bu
t ra
re i
n th
e ~
r;·
win
dwar
d is
land
s ... 0
E.
vari
ans
(Sow
erby
) X
A
lso
in H
awai
i =
~-P
yren
e ob
tusa
(So
wer
by)
X
X
II>
(US
NM
NH
) O
liva
ann
ulat
a G
mel
in
X
Sem
inel
la v
irgi
nea
X
X
Als
o in
Haw
aii
(Gou
ld)
Sem
inel
la s
p.
X
Vas
um c
eram
icum
(L
inne
) X
X
X
X
X
X
S
arig
an
V.
turb
inel
lus
(Lin
ne)
Ala
mag
an
X
(US
NM
NH
) L
atir
us b
arcl
ayi
(Ree
ve)
X
X
X
X
X
X
Agr
ihan
(U
SN
MN
H)
L.
notk
ltus
(G
mel
in)
X
X
X
Alm
agan
A
lso
in H
awai
i (U
SN
MN
H)
Agr
ihan
(U
SN
MN
H)
L.
noum
eens
is (
Cro
sse)
X
Per
iste
rnia
nas
satu
la
X
X
X
X
Ala
mag
an
(Lam
arck
) A
grih
an
Mit
ra c
hrys
osto
ma
X
X
Bro
deri
p M
. im
peri
o/is
(R
odin
g)
X
Mau
g re
cord
tha
t o
f C
erno
hors
ky
(197
6)
M.
stic
tica
(L
ink)
X
A
lso
in H
awai
i St
riga
tell
a fa
stig
ium
X
X
X
A
lso
in H
awai
i (R
eeve
) S.
/it
tera
ta (
Lin
ne)
X
X
X
Inte
rtid
al;
also
in
Haw
aii
S. p
aupe
rcul
a (L
inne
) X
X
S. p
elli
sser
pent
is (
Ree
ve)
X
X
Als
o in
Haw
aii
<
• Str
igat
el/a
sp.
X
X
X
~ -
Neb
ular
ia a
uror
a jl
orid
ula
Sar
igan
~
(Sow
er by
) 0
N. f
erru
gine
a X
X
A
grih
an
Als
o in
Haw
aii
~ 9 L
amar
ck
(US
NM
NH
) g"
N
. ch
rysa
lis
(Ree
ve)
X
.... -(U
SN
MN
H)
'0
00
Cos
tel/
aria
cro
cata
A
lso
in H
awai
i ..,
X
(Lam
arck
)
C.
coro
natu
m (
Hel
blin
g)
X
C.
unif
asci
alis
A
lam
agan
A
lso
in H
awai
i (L
amar
ck)
C.
paci
ficu
m (
Ree
ve)
X
*Pus
ia s
p.
X
Pus
ia a
mab
i/is
(R
eeve
) X
X
P.
canc
ella
rioi
des
Agr
ihan
A
lso
in H
awai
i (A
nton
) P
. co
nsan
guin
ea (
Ree
ve)
X
Als
o in
Haw
aii
P.
/aul
a (R
eeve
) X
A
lso
in H
awai
i
""" _,
App
endi
x. (
cont
inue
d)
""'"
00
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
P.
mic
rozo
nias
X
X
A
lso
in H
awai
i (L
amar
ck)
P.'f
atri
arch
a/is
(G
mel
in)
X
X
Agr
ihan
A
lso
in H
awai
i (U
SN
MN
H)
P.
spec
iosa
(R
eeve
) X
lmbr
icar
ia o
liva
efor
mis
X
A
lso
in H
awai
i (S
wai
nson
) I.
pun
cta/
a (S
wai
nson
) X
S
arig
an
Als
o in
Haw
aii
Ter
ebra
aff
inis
Gra
y X
A
lso
in H
awai
i T
. cre
nula
ta (
Lin
ne)
X
X
Als
o in
Haw
aii
T.
dim
idia
ta (
Lin
ne)
X
X
Als
o in
Haw
aii
(US
NM
NH
) s::
T. f
elin
a (D
illw
yn)
X
Als
o in
Haw
aii
r;·
T.
laev
igat
a (G
ray)
X
.... 0
Has
tula
so/
ida
Als
o in
Haw
aii
;:;
X
"' "' (D
esha
yes)
r;·
I>
'
Has
tula
str
igil
lata
X
A
lso
in H
awai
i (L
inne
) L
iena
rdia
sp.
X
L.
cf.
L. m
ighe
lsi
X
X
Ired
ale
and
Tom
lin
Car
inap
ex m
inut
issi
ma
X
Als
o in
Haw
aii
(Gar
rett
) M
acte
ola
sege
sta
(Che
nu)
X
Als
o in
Haw
aii
Ired
alea
sp.
X
Tri
tono
turr
is s
p.
X
*Con
us a
ulic
us L
inne
X
C.
balt
eatu
s S
ower
by
X
Ala
mag
an
Agr
ihan
C.
band
anus
Hw
ass
X
X
Ala
mag
an
*C.
cano
nicu
s R
odin
g X
A
grih
an
(US
NM
NH
)
C.
capi
tane
us L
inne
A
lam
agan
A
lso
in H
awai
i
C.
catu
s H
was
s X
X
A
grih
an
Als
o in
Haw
aii
C.
chal
daeu
s (R
odin
g)
X
X
Als
o in
Haw
aii
(US
NM
NH
)
C.
coro
natu
s (H
was
s)
X
X
Als
o in
Haw
aii
(US
NM
NH
)
C.
dist
ans
Hw
ass
X
Agr
ihan
A
lso
in H
awai
i
C.
ebra
eus
Lin
ne
X
X
Als
o in
Haw
aii
C.
jlav
idus
Lam
arck
X
X
X
X
X
A
grih
an
Als
o in
Haw
aii
C.
frig
idus
Ree
ve
X
Agr
ihan
C.
fulg
etru
m S
ower
by
X
C.
gene
rali
s L
inne
X
A
grih
an
<
(US
NM
NH
) 12-
C.
geog
raph
us L
inne
X
~
C.
impe
rial
is L
inne
X
A
grih
an
Als
o in
Haw
aii
0
C. l
itog
lyph
us H
was
s X
X
X
A
lam
agan
A
lso
in H
awai
i ~
C.
livi
dus
Hw
ass
X
X
Ala
mag
an
Als
o in
Haw
aii
9 X
~
C.
mag
nifi
cus
Ree
ve
X
X
'1
C.
mar
mor
eus
Lin
ne
X
Als
o in
Haw
aii
:;;
CXI
C.
mil
es H
was
s X
X
X
X
X
A
lam
agan
A
lso
in H
awai
i ....,
Agr
ihan
C.
mil
iari
s H
was
s X
X
X
X
X
A
lam
agan
A
lso
in H
awai
i A
grih
an
C.
mor
elet
i C
ross
e X
X
X
X
A
lso
in H
awai
i
C.
mus
icus
Hw
ass
X
Agr
ihan
C.
nanu
s S
ower
by
X
X
X
X
X
X
Sar
igan
A
grih
an
C.
obsc
urus
Sow
erby
X
A
lso
in H
awai
i
C.
pert
usus
Hw
ass
X
Als
o in
Haw
aii
C.
puli
cari
us H
was
s X
X
X
X
X
A
lso
in H
awai
i
C.
ratt
us H
was
s X
X
X
X
S
arig
an
Als
o in
Haw
aii
Ala
mag
an
Agr
ihan
-!:
> \0
App
endi
x. (
cont
inue
d)
V>
0
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
C.
sang
uino
lent
us
X
X
Quo
y an
d G
aim
ard
~ sp
onsa
lis
Hw
ass
X
X
X
X
Sar
igan
A
lso
in H
awai
i A
grih
an
C.
stri
atus
Lin
ne
X
Agr
ihan
A
lso
in H
awai
i (U
SN
MN
H)
C.
tess
u/at
us B
orn
Agr
ihan
(U
SN
MN
H)
C.
tuli
pa L
inne
X
X
C.
vari
us L
inne
X
C.
virg
o L
inne
X
EU
TH
YN
EU
RA
~
Mic
rom
elo
guam
ensi
s A
lso
in H
awai
i <r
X
... 0 (Q
uoy
and
Gai
mar
d)
1::1 " "'
Smar
agdi
ne/l
a ca
lycu
lata
X
X
X
X
In
tert
idal
; al
so
<r I»
(Bro
deri
p an
d in
Haw
aii
Sow
er by
) H
amin
oea
cym
balu
m
X
X
X
Als
o in
Haw
aii
(Quo
y an
d G
aim
ard)
H
amin
oea
sp.
X
X
X
Act
eoci
na h
awai
ensi
s X
X
A
lso
in H
awai
i Pi
lsbr
y A
cteo
cina
sp.
X
Cyl
ichn
a sp
. X
Ret
usa
sp.
X
X
Aty
s sp
. X
Siph
onar
ia c
f. S.
X
In
tert
idal
guam
ensi
s (Q
uoy
and
Gai
mar
d)
*S.
sir
ius
Pils
bry
X
X
X
Inte
rtid
al
Wil
/iam
ia r
adia
ta
X
X
Als
o in
Haw
aii
(Pea
se)
Mel
ampu
s ca
stan
eus
X
X
X
Sari
gan
Als
o in
Haw
aii;
(Muh
lfel
d)
spra
y z o
ne
M. j
lavu
s (G
mel
in)
X
Spr
ay z
one
Mir
alda
cf.
M.
X
scop
ulor
um W
atso
n P
yram
idel
lida
e sp
. I
X
Pyr
amid
elli
dae
sp.
2 X
X
X
Pyr
amid
elli
dae
sp.
3 X
X
X
Julia
ex
quis
ita
Gou
ld
X
X
Als
o in
Haw
aii
PO
LY
PLA
CO
PH
OR
A
<
Aca
ntho
pleu
ra g
emm
ata
X
X
X.
Inte
rtid
al
~
(Bla
in vi
lle)
(US
NM
NH
) ;o
Pol
ypla
coph
ora
sp.
X
0 § B
IVA
LV
IA
9 g-A
rea
ave
/lana
Lam
arck
X
X
"1
-A
. ve
ntr
icos
a L
amar
ck
X
X
X
X
Als
o in
Haw
aii
"' 00
Aca
r di
vari
cata
A
lso
in H
awai
i ""
X
X
(Sow
er by
) B
arba
tia
desc
ussa
ta
X
X
Als
o in
Haw
aii
(Sow
erby
) B
. am
ygdo
lum
tost
um
X
Rod
ing
B.
tene
lla R
eeve
X
A
lso
in H
awai
i M
odi
olus
aur
icul
atus
X
(Kra
uss)
Se
ptif
er e
xcis
us
X
X
(Wig
man
n)
"'
App
endi
x. (
cont
inue
d)
tJ>
N
Ana
taha
n G
ugua
n P
agan
A
sunc
ion
Mau
g U
raca
s O
ther
R
emar
ks
Chl
amys
cor
usca
ns
X
In H
awai
i as
C.
c.
coru
scan
s (H
inds
) ha
wai
ensi
s C
. sq
uam
osa
(Gm
elin
) X
X
Sfto
ndyl
us s
p.
X
X
Lim
a a
nnul
ata
X
(Lam
arck
) L
. li
ma
(Lin
ne)
X
*Gly
cym
eris
sp.
X
Pin
ctad
a m
arga
riti
fera
X
X
A
lso
in H
awai
i (L
inne
) P
inct
ada
sp.
X
lsog
nom
on p
erna
X
A
lso
in H
awai
i (L
inne
) a: <r
I. i
ncis
um (
Con
rad)
X
A
lso
in H
awai
i ..., 0
Pyc
nodo
nte
num
ism
a X
::s .. "'
(Lam
arck
) (U
SN
MN
H)
;:;- "' C
ardi
ta v
arie
gata
X
(Bru
giue
re)
Tra
pezi
um o
blon
gum
X
X
A
lso
in H
awai
i (L
inne
) F
imbr
ia f
imbr
iata
X
M
aug
reco
rd f
rom
(L
inne
) N
icol
(19
50)
Fim
bria
sp.
X
Cha
ma
iost
oma
(Con
rad)
X
X
X
A
lso
in H
awai
i C
ham
a sp
. X
A
grih
an
Tri
dacn
a m
axim
a R
odin
g X
X
S
arig
an
T.
squa
mos
a L
amar
ck
X
X
Lio
conc
ha h
iero
glyp
hica
X
A
lso
in H
awai
i (C
onra
d)
Gaf
rari
um p
ee ti
na tu
m
X
(Lin
ne)
Per
ig/y
pta
reti
cu/a
ta
(Lin
ne)
Nes
obor
nia
bart
schi
C
hava
n Q
uidn
ipag
us p
alat
um
Ired
ale
Pin
guit
elli
na r
obus
ta
Han
ley
Tel
/ina
cru
cige
ra
Lam
arck
T
el/i
na s
p.
Erv
ilia
san
dwic
hens
is
(Sm
ith)
C
tena
bel
la (
Con
rad)
L
imop
sis
sp.
Car
dite
lla
cf.
C.
haw
aien
sis
Dal
l,
Bar
tsch
and
Reh
der
X
X
X
X
X
X
X
X
X
X
X
X
Als
o in
Haw
aii
Als
o in
Haw
aii
Als
o in
Haw
aii
Als
o in
Haw
aii
Als
o in
Haw
aii
Als
o in
Haw
aii
Als
o in
Haw
aii
~ - ~ t:l i -10 0
0
w
v.
w
54 Micronesica
APPENDIX 2 Molluscs common in the southern Marianas but unknown from the northern
Marianas (only species living on hard bottom are included)
Species Intertidal Reef
Cellana radiata orienta/is (Pilsbry) + Patelloida sp. + Trochus ni/oticus (Linne) + Echininus viviparus Rosewater + Planaxis sulcatus (Born) + Alvinia crystal/ina (Garrett) + Cerithium sejunctum Iredale + C. zona tum Wood + C/ypeomorus alveolus (Hombron and Jacquinot) + C. bifasciatus (Sowerby) + C. moniliferus (Kiener) + Hipponix foliaceus (Quoy and Gaimard) + Cypraea annulus Linne + Cymatium gemmatum (Reeve) + Thais acu/eata Deshayes and Milne Edwards + Morula fiscella (Gmelin) + M. nodicostata (Pease) + M . squamosa (Pease) + Morula sp. + Muricodrupa fenestrata (Wood) + Nebularia cucumerina (Lamarck) + Strigatella decurtata (Reeve) +
Eldredge, L. G., R. T. Tsuda, P. Moore, M. Chernin, and S. Neudecker. 1977. A natural history of Maug, northern Mariana Islands. Univ. Guam Mar. Lab. Tech. Rept. 43. 87 p.
Emerson, W. K. 1978. Molluscs with Indo-Pacific faunal affinities in the Eastern Pacific Ocean. Nautilus 92: 91- 96.
Emerson, W. K., and W.O. Cernohorsky. 1973. The genus Drupa in the Indo-Pacific. Indo-Pacific Mollusca 3(13): 801-863.
Grigg, R. W. 1981. Acropora in Hawaii. Part 2. Zoogeography. Pac. Sci. 35: 15-24. ltono, H . 1980. The genus Ga/axaura in Micronesia. Micronesica 16: 1-19. Kay, E. A. 1967. The composition and relationships of marine molluscan fauna of the Hawaiian
Islands. Venus, Japan. J. Malac. 25: 94-104. 1971. The littoral marine molluscs of Fanning Islands. Pac. Sci. 25(2): 260--281. 1979. Hawaiian marine shells. Bishop Museum Press, Honolulu. 653 p. 1980a. Little worlds of the Pacific: an essay on Pacific basin biogeography. Harold L. Lyon
Arboretum Lecture Number 9. University of Hawaii, Honolulu, Hawaii. 1980b. Micromolluscs: techniques and patterns in benthic marine communities. In
Environmental survey techniques for coastal water assessment conference proceedings. Water Resources Research Center and Sea Grant College Program, University of Hawaii.
1984. Patterns of speciation in the Indo-West Pacific. In Raven, P., F. Radovsky and S. H . Sohmer, Biogeography of the Tropical Pacific: Proceedings of a Symposium. Association of Systematics Collections and B. P. Bishop Museum.
Ladd, H . S., J. l.;Fracey, Jr., and G. Gross. 1970. Deep drilling on Midway Atoll. Geol. Surv. Prof. Pap. 680--A.
Vol. 19. December 1983 55
Nicol, D. 1950. Recent species of the lucinoid pelecypod Fimbria. J. Washington Acad. Sci. 40: 82-87. Randall, J. E. 1982. Examples of antitropical and antiequatorial distribution of Indo-West Pacific
fishes. Pac. Sci. 35: 197- 209. Rehder, H. A. 1980. The marine molluscs of Easter Island (Isla de Pascua) and Sala y Gomez.
Smithsonian Contrib. Zoo!. 289: 1- 167. ' Rosewater, J. 1970. The family Littorinidae in the Indo-Pacific. I. The subfamily Littorininae. Indo
Pacific Mollusca 2(11): 417-506. Rosewater, J. , and D. Kadolsky. 1981. Rectifications in the nomenclature of some Indo-Pacific
Littorinidae II. Proc. Bioi. Soc. Washington 94: 1233-1236. Safriel, U., and Y. Lipkin. 1975. Patterns of colonization of the eastern Mediterranean intertidal zone
by Red Sea immigrants. J. Ecol. 63: 61-63. Springer, V. G. 1981. Comments on Solem's land-snail biogeography with an hypothesized expla
nation of the distribution of the Endondontidae. pp. 225-230./n G. Nelson and D. E. Rosen (eds.). Vicariance biogeography. Columbia University Press, N . Y.
1982. Pacific plate biogeography, with special reference to shorefishes. Smithsonian Contrib. Zoo!. 367: 1-182.
Stanley, S. M. 1979. Macroevolution. Pattern and process. W. H. Freeman, San Francisco. Talmadge, R. R. 1963. Insular haliotids in the Western Pacific (Mollusca: Gastropoda). Veliger 5:
129- 139. Tsuda, R. T., and W. J. Tobias. 1977a. The marine benthic algae from the northern Mariana Islands,
Chlorophyta and Phaeophyta. Bull. Japan. Soc. Phycol. 25: 19-24. 1977b. The marine benthic algae from the northern Mariana Islands, Cyanophyta and
Rhodophyta. Bull. Japan. Soc. Phycol. 25: 155-158. Vermeij, G. J. 1972. Endemism and environment: some shore molluscs of the tropical Atlantic. Amer.
Nat. 106: 89-101. 1978. Biogeography and adaptation: patterns of marine life. Harvard University Press,
Cambridge. 332 p. Zinsmeister, W. J., and W. K. Emerson. 1979. The role of passive dispersal in the distribution of
hemipelagic invertebrates, with examples from the tropical Pacific Ocean. Veliger 22: 32-40.