Land Snails from Archaeological Sites in the Marshall Islands,
with Remarks on Prehistoric Translocations in Tropical Oceania
Author(s): Carl C. Christensen and Marshall I. Weisler
Source: Pacific Science, 67(1):81-104. 2013.
Published By: University of Hawai'i Press
DOI: http://dx.doi.org/10.2984/67.1.6
URL: http://www.bioone.org/doi/full/10.2984/67.1.6
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Land Snails from Archaeological Sites in the Marshall Islands, with
Remarks on Prehistoric Translocations in Tropical Oceania1
Carl C. Christensen2,4 and Marshall I. Weisler 3
Abstract: We report the recovery of 11 taxa of nonmarine mollusks from
archaeological sites on Majuro, Maloelap, and Ebon Atolls, Republic of the
Marshall Islands. Pupina complanata (Pupinidae), Omphalotropis fragilis (Assimineidae), Truncatella guerinii (Truncatellidae), Lamellidea pusilla and Pacificella
variabilis (Achatinellidae), Gastrocopta pediculus (Gastrocoptidae), Nesopupa sp.
( Vertiginidae), “Succinea” sp. (Succineidae), Allopeas gracile (Subulinidae), and
Liardetia samoensis (Helicarionidae) arrived in these islands prehistorically; Liardetia sculpta (Helicarionidae) has not yet been recovered from levels of confirmed
prehistoric age. Pupina complanata, O. fragilis, and probably also Nesopupa sp., and
“Succinea” sp. are Micronesian endemics. All other species are widely distributed
in Micronesia and Polynesia and (except for the strand-line species T. guerinii)
were undoubtedly translocated to the Marshall Islands by the prehistoric voyages of Pacific islanders. The precise role of human transport in the dispersal of
the Micronesian endemics remains unclear, but because these atolls have been
emergent for a mere 3,000 yr or so, human transport is likely in view of the
known rarity of natural interarchipelagic dispersal of nonmarine mollusks.
become more widely distributed as a result of
prehistoric voyaging. Matisoo-Smith (2009)
and Storey et al. (2011) summarized how recent advances in phylogenetic analysis have
enabled workers to study the routes by which
rats, pigs, and chickens were transported
throughout Oceania by the Pacific islanders
before the advent of European influence, and
other workers have applied these methods to
determine the histories of reptiles that may
have been similarly translocated (Hamilton
et al. 2010). Lee et al. (2007) and Ó Foighil
et al. (2011) used similar techniques to identify instances of human translocation of tree
snails of the family Partulidae. For the most
part, however, investigations of the prehistoric translocation of Pacific island land snails,
as with insects (Porch 2011), are at the much
more preliminary stage of attempting to identify species that have probably been carried
about by human commerce and using information from archaeological excavations to
determine whether these were introduced to
these islands prehistorically or in modern
times. Malacologists have long been aware of
the substantial role humans played in transporting land snails to various islands in the
Archaeologists have long recognized that
the prehistoric voyagers who colonized the
islands of Micronesia and Polynesia carried
with them a “transported landscape” (Anderson 1952, Kirch 1984) of intentionally and
inadvertently translocated animal and plant
species. Although some of these species have
origins outside this region, others are indigenous to particular islands within it but have
1
Archaeological fieldwork on Maloelap and Ebon
Atolls was financially supported, in part, by the U.S. National Park Service through the Historic Preservation
Office, Republic of the Marshall Islands. Manuscript accepted 3 April 2012.
2
Research Associate, Bishop Museum, 1525 Bernice
Street, Honolulu, Hawai‘i 96817.
3
School of Social Science, University of Queensland,
Saint Lucia, Queensland 4072, Australia (e-mail:
m.weisler@uq.edu.au).
4
Corresponding author (e-mail: carlcc@hawaii.rr
.com).
Pacific Science (2013), vol. 67, no. 1:81 – 104
doi:10.2984/67.1.6
© 2013 by University of Hawai‘i Press
All rights reserved
81
82
Pacific (Reigle 1964, Cowie and Robinson
2003, Cowie 2005), and more than 100 yr ago
Garrett (1879) and Pilsbry (1900) suggested
that this process may have begun with the
pre-Contact voyages of the Pacific islanders.
Only recently, however, has archaeological
evidence become available documenting the
chronology of these introductions. Kirch
(1973) was the first to report humantransported land snails from pre-Contact archaeological sites in Oceania, and works by
Christensen and Kirch (1981), Hunt (1981),
and Sinoto (1983) followed within the next
decade. Although Anderson (2009) has opined
that archaeological data as to the pre-Contact
dispersal of land snails in Oceania were “few,”
in fact a rather substantial body of such reports has accumulated over the last 40 yr.
Articles have presented primary data regarding the prehistoric occurrence of presumably
translocated land snails in the southeastern
Solomon Islands (Christensen and Kirch 1981,
Leach and Davidson 2008), New Caledonia
(Cowie and Grant-Mackie 2004), the Marshall
Islands ( Weisler 1999), Fiji (Hunt 1981), Samoa (Kirch 1993), Tonga (Kirch 1988), the
Cook Islands (Allen 1992, 1997, 1998, Allen
and Christensen 1992, Craig 1995, Kirch et al.
1995, Walter 1998, Brook 2010, Brook et al.
2010), the Society Islands (Sinoto 1983, Orliac 1997), the Gambier Islands (Kirch et al.
1990, 2004, Howard and Kirch 2004, Conte
and Kirch 2008), Henderson Island (Preece
1998), the Marquesas Islands (Kirch 1973,
Rolett 1992, 1998), and the Hawaiian Islands
(Christensen 1984a, Christensen and Kirch
1986, Kirch 1989, 1992, Dixon et al. 1997,
Burney et al. 2001, Burney 2002).
Although some land snails are believed to
have been intentionally transported from island to island (Powell 1938, Hayward and
Brook 1981, Lee et al. 2007, Ó Foighil et al.
2011), most such species are small (<5 mm)
and were undoubtedly transported inadvertently as stowaways on plants carried interisland, just as the modern horticulture industry
disseminates alien snails and slugs throughout
Oceania and elsewhere (Cowie and Robinson
2003, Cowie et al. 2008). The high islands of
Micronesia and Polynesia have diverse native
land snail faunas, most members of which are
PACIFIC SCIENCE · January 2013
endemic to single islands or to the islands of a
single archipelago (Solem 1959, Cowie 1996).
Not all of the terrestrial mollusks of Oceania
are so narrowly endemic, however, and “the
land snails of the Pacific islands may be divided into two groups: those living on low islands and on the shore zone of high islands,
and those inhabiting the forests of high islands. Many of the first group have a wide distribution suggestive of dispersal by human or
other adventitious agency” (Pilsbry 1916:429).
These widely distributed species dominate
the terrestrial mollusk fauna of coral atolls
and other low islands in the Pacific (Cooke
1928, Reigle 1964, Harry 1966) and are sometimes referred to collectively as the “atoll
fauna” (Solem 1959).
The land snail fauna of the Marshall Islands has attracted little attention in the past,
no doubt because of its depauperate nature
and the absence of endemic taxa. The principal references are Pease (1861) and Reigle
(1964), discussing collections from Ebon and
Rongelap Atolls, respectively. The only published report of land snails from archaeological sites in the Marshall Islands is a brief mention by Weisler (1999) of the collections from
Maloelap Atoll that are analyzed here.
The Marshall Islands consist of 29 low
coral atolls and five small coral islands in eastern Micronesia that lack lagoons and are
aligned in two parallel island chains trending
northwest-southeast for about 1,200 km between 4° and 12° north latitude (Figure 1).
The atolls range in size from Jemo ( Jämo),
the smallest at 0.16 km2, to the world’s largest
atoll, Kwajalein (Kuwajleen), with its 16.4
km2 of land surrounding a 2,174 km2 lagoon.
Before human colonization a mixed broadleaf
forest, predominantly of Tournefortia argentea, Guettarda speciosa, and Pisonia grandis,
was presumably common on most atolls,
with salt-tolerant shrubs to small trees such
as Scaevola sericea and Pemphis acidua along
shorelines (Fosberg 1990, Merlin et al. 1997).
Economically important plants such as breadfruit and coconut, as well as two kinds of taro
(Cyrtosperma chamissonis and Colocasia esculenta), were prehistoric introductions. Pandanus
tectorius, naturally dispersed, was undoubtedly
selected and developed into several important
Marshall Islands Land Snails · Christensen and Weisler
83
Figure 1. The Marshall Islands, showing the location of Maloelap, Majuro, and Ebon Atolls, where sediment samples
were obtained for the analysis of land snails.
varieties in the Marshalls (Stone 1960, Merlin
et al. 1997:35).
The Marshall Islands emerged about 3,000
yr ago (Dickinson 2003), but the available
land area was probably not sufficiently large
to support human habitation or other terrestrially bound species for several hundred years
after initial emergence ( Weisler et al. 2012).
No credible radiocarbon age determinations
of habitation layers predate ∼2,000 B.P.
( Weisler 1999). This temporal window constrains the time that humans could have transported land snails. The land snail samples reported here are all from the largest islet from
each of three atolls where large expanses of
pits (presumably for Giant Swamp Taro, Cyrtosperma) were formerly under cultivation. It is
these largest islets, with the most substantial
subterranean freshwater lenses, that human
colonists would have targeted for initial settlement of any atoll and brought vital cultigens
for establishing their agricultural systems.
In this paper we (1) present new records
and discuss the biogeography of land snails
systematically collected from archaeological
excavations in the Marshall Islands (some of
these species were translocated into or within
the Pacific islands before the arrival of Europeans, which ranges from the early 1500s
for the Mariana Islands to the late 1700s for
Hawai‘i); (2) discuss the evidence used to elucidate the history and status of cryptogenic
species; and (3) emphasize the importance of
land snails for understanding the process of
human colonization of islands in general, and
Pacific atolls in particular.
materials and methods
Sampling, Data Presentation,
and Taxonomic Analysis
Sediment samples from Maloelap and Ebon
were collected by M.I.W. The sediment
84
weight and volume were recorded before
water-screening with stacked 2, 1, and 0.5
mm sieves. Land snails were then picked from
the resulting residue. Land snails from excavations on Majuro were obtained in 1979 by
Thomas J. Riley; extraction procedures were
not specified. Snails were obtained from seven
separate samples from Maloelap, 11 from
Ebon, and eight from Majuro. All specimens
are held in the malacological collections of
the Bishop Museum and were identified by
reference to those collections and appropriate
literature; individual catalog numbers have
not yet been assigned.
Specimen counts are based on the number
of whole shells or apical fragments found in
samples following the methodology of Evans
(1972); when a taxon is represented in a sample by only nonapical fragments, its presence
is indicated in the lists of material recorded by
a plus sign (+), but the number of fragments is
not quantified.
Archaeological Site and Layer Descriptions
maloelap atoll (8° 45′ N, 171° 3′ E):
Maloelap consists of 71 islets encircling a 970
km2 lagoon; at ∼1.5 km long, Kaven is the
largest islet and is located in the northwestern
corner of the atoll ( Weisler 1999: fig. 2). A
substantial subterranean freshwater GhybenHerzberg lens supported a zone of elongate
pits for giant swamp taro (Cyrtosperma) cultivation just inland of the major ancient village
of the atoll. On Kaven Islet, archaeological
trench excavations through one of the prehistoric spoil heaps at site MLMl-3 that sealed
an ancient A horizon revealed a sequence of
pit construction beginning about 2,000 yr
ago, probably coinciding with the first few
human generations of settlement. Six land
snail samples along the trench, all in prehistoric contexts, are reported here. (See Weisler
1999:634 –640 and fig. 6 for a comprehensive
description of the site and associated dates.)
The brief layer descriptions for site MLMl3, test pits (TP [also referred to as “Units”])
18–27, and context for the land snail samples
are as follows:
Layer IA: A black (10YR2/1, Munsell
Color Charts, taken moist) sandy gravel to
PACIFIC SCIENCE · January 2013
gravelly sand with few prehistoric artifacts,
sparse marine mollusks and bone, combustion
features, and post molds to an average depth
of 27 cm bs (centimeters below surface). Two
radiocarbon age determinations were obtained from two test pits along the 22 m long
trench. An earth oven from TP 19, 20–52 cm
bs, was calibrated to A.D. 1175–1400 (Beta
Analytic, Inc. dating sample number-79575,
720 B.P. ± 70 at 2σ), and another oven from
TP 22, 30–71 cm bs, was calibrated to A.D.
1810–1925 (Beta-77602, 20 ± 50) ( Weisler
1999:table II). This latter charcoal sample,
calibrated to the historic period, was at least 4
m from land snail sample 1 ( Weisler 1999:fig.
6). All radiocarbon age determinations reported here were calibrated using Stuiver et
al. (2005) and rounded to the nearest 5 yr.
Layer IB: A sparse prehistoric cultural
layer consisting primarily of dispersed charcoal and minor amounts of marine mollusks
and bone and no combustion features. The
dark gray (5YR4/1) gravelly sand was only
present in one test pit of the trench.
Layer II: This layer is spoil dirt deposited
from excavating the adjacent cultivation pit in
prehistory. Averaging up to 73 cm thick, this
culturally sterile layer consists of a very pale
brown (10YR7/3) to pinkish white (7.5YR8/2)
sand to gravelly sand with cemented cobbles.
Layer III: This buried A horizon consists
of a very dark grayish brown (10YR3/2) to
very pale brown (10YR7/3) sand to slightly
gravelly sand with scattered charcoal flecks
and an average depth of 119 cm bs. A radiocarbon age determination on unidentified
charcoal flecks was calibrated to 55 B.C.–
A.D. 255 (Beta-79576, 1910 ± 70) ( Weisler
1999:table II).
Layer IV: The culturally sterile subsoil
consists of a very pale brown (10YR8/3)
weakly cemented sand.
ebon atoll (4° 38′ N, 168° 43′ E): Ebon’s lagoon encloses 104 km2 and is surrounded by 22 islets with a total land area of 5.8
km2. Rosendahl (1987:83–88) surveyed Ebon
Islet, by far the largest on the atoll, and briefly
noted six midden sites and collected nearly
400 traditional artifacts. During 1995–1996,
Weisler (2002) surveyed the entire atoll and
recorded 23 additional prehistoric and his-
Marshall Islands Land Snails · Christensen and Weisler
toric sites including middens, burials, cultivation areas, a trail, and house sites.
Land snail samples were taken at two locations at site MLEb-2I on Ebon Islet. Test pits
1–5 excavated across a spoil heap in the cultivation zone near the interior of Ebon Islet
revealed a ∼2,000-yr-old buried A horizon,
making this one of the earliest dated contexts
for the Marshall Islands. Samples for land
snail analysis were taken from this A horizon
(Layer III) as well as from Layer I. A comprehensive description of the site, layers, and associated dates is reported in Weisler (2002).
Here, we briefly describe the sample contexts
from site MLEb-2I, TP 2:
Layer I: A black (10YR2/1) sandy gravel,
surface to about 20 cm thick, with abundant
prehistoric artifacts, features (a human burial,
combustion feature, and post mold), marine
food mollusks, and bones of fish, lizard, and
rat.
Layer III: A buried dark gray (2.5YN4/0)
A horizon consisting of coarse sand with
charcoal flecks, about 55–70 cm bs (averaging about 15 cm thick), A radiocarbon age
determination on dispersed charcoal was
calibrated to 40 B.C.–A.D. 140 (Beta-92123,
1930 ± 40).
At TP 20, a vertical series of nine samples
was taken; the stratigraphy of the test pit is as
follows:
Layer IA: A black (2.5Y2.5/1) compact
dense gravel, averaging about 25 cm thick,
with modern artifacts; bones of the historically introduced chicken, as well as of fish, rat,
and sea turtle; and food marine mollusks. No
sediment samples for land snail analysis were
taken from this disturbed layer.
Layer IB: A very dark gray (2.5Y3/1) dense
gravel, about 50 cm thick, with prehistoric artifacts; bones of fish, seabirds, Pacific rat, sea
turtle, and dog; marine food mollusks; and
several combustion features (samples 6 and 7).
Table 1 lists all the land snail samples and
their provenances.
Layer IC: The lowest cultural layer (75
to about 120 cm bs) consisted of very dark
gray (2.5Y3/1) sandy gravel grading to gray
(2.5Y6/1) with abundant prehistoric artifacts;
bones of fish, seabird, dog, Pacific rat, small
lizard, and sea turtle; food marine mollusks;
85
and several combustion features. A charcoal
sample of coconut husk and shell, pandanus,
and wood charcoal yielded a calibrated radiocarbon age of A.D. 1175–1300 (Beta-92127,
750 ± 50) (samples 8, 9, 10, and 11).
Layer II: A culturally sterile white (2.5Y8/2)
lagoon sand subsoil encountered to the maximum depth of excavations at 160 cm bs (samples 12, 13, and 14).
majuro atoll (7° 9′ N, 171° 3′ E): Centrally located in the archipelago, Majuro consists of 64 islets with a land area of 9.2 km2
surrounding a 295 km2 lagoon. In 1979 Riley
(1987) completed three transect excavations
across Majuro Islet at the Laura village site
MLMj-1, and eight sediment samples for land
snail analysis were collected from transect 6,
TP 3, 4, 5, and 7 from ∼10 cm bs to ∼240 cm
bs. Riley described the excavations, stratigraphy, and dating (1987:206–218, 242–243). TP
3, 4, 5, and 6 were excavated in midden areas,
but TP 7 was situated on an aroid pit spoil
heap (Riley 1987:216–218) and revealed a
buried A horizon (Layer VI) 240 cm bs. The
dates for ovens in TP 3 and 5 are roughly
2,000 yr old (see description following). Although the buried A horizon was not dated,
the depth at ∼240 cm bs suggests relatively
early use of this site area. No historic materials were recovered from any of the cultural
layers, and the two radiocarbon age determinations, although not directly associated with
sediment samples analyzed here, suggest a
great antiquity for some of the prehistoric
layers. The layers where sediment samples
were collected for land snail analysis and contexts for two radiocarbon age determinations
are described here:
Layer IA: At TP 6, this layer, averaging
10–12 cm thick, consisted of a black (10YR2/1)
sandy soil with charcoal, coral gravel, shell
midden, and combustion features (Riley
1987:table 2.15). No historic materials were
recovered, and cultural content suggests a
prehistoric date of deposition.
Layer IB: In TP 5, this layer, extending to
60 cm bs overall, consisted of a very dark sand
(10YR3/1) with coral gravel, shell midden, artifacts, and human remains. A human burial
pit extended this layer to 115 cm bs (Riley
1987:213). In TP 5, at the “interface of Layer
86
PACIFIC SCIENCE · January 2013
TABLE 1
Provenance of Land Snail Samples from the Marshall Islands
Sample No.
Atoll
Site
10-2 (96-7)
1
2
3
4
1
3
6
7
8
9
10
11
12
13
14
Maloelap
Maloelap
Maloelap
Maloelap
Maloelap
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Ebon
Majuro
Majuro
Majuro
Majuro
Majuro
Majuro
Majuro
Majuro
MLMl-3
MLMl-3
MLMl-3
MLMl-3
MLMl-3
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLEb-2I
MLMj-1
MLMj-1
MLMj-1
MLMj-1
MLMj-1
MLMj-1
MLMj-1
MLMj-1
a
Transect
Test Pit /Unit
Layer a
Depth below
Surface (cm)
6
6
6
6
6
6
6
6
19
21
19
21
21
2
2
20
20
20
20
20
20
20
20
20
3
3
4
4
5
5
7
7
III (A)
I
III (A)
II
III (A)
I
III (A)
IB
IB
IC
IC
IC
IC
II
II
II
I
II
I
II
IA
IB
VI
VII
144–154
∼25
144–154
∼135
144–154
10
55–65
49–62
61–70
70–80
78–92
90–101
99–110
109–121
120–144
135–150
∼10
∼30
10–15
∼50
∼10
∼50
∼200
∼240
III (A) refers to Layer III, buried A horizon.
IB” (Riley 1987:213), a small oven was radiocarbon dated to 0 B.C.–A.D. 265 ( ISGS [Indiana State Geological Survey]-671, 1970 ± 110
B.P.). This sample probably dates the bottom
of Layer IB.
Layer II: Sediment samples were collected
from this layer in TP 3, 4, and 6, and layer
descriptions varied between each locality.
The layer began 12 to 24 cm bs across the test
pits and varied from 8 to 60 cm in thickness.
The sandy sediments were gray (10YR4/2) to
dark gray (10YR4/1) and contained various
amounts of charcoal flecking, shell midden,
and combustion features (Riley 1987:tables
2.13–2.15).
Layer III: No sediment samples were collected from this layer, but a charcoal sample
from a small earth oven in TP 3 at 72 cm bs
was calibrated to 45 B.C.–A.D. 260 ( ISGS669, 1890 ± 75 B.P.). This date may be
younger than sediment samples collected 70
m distant from TP 7, Layers VI and VII.
Layer VI: Only in TP 7, this layer is a
brownish-gray (10YR4/1) sand, “culturally
sterile; apparent A horizon” (Riley 1987:table
2.16). It is 240–265 cm bs and averaged 25 cm
thick.
Layer VII: In TP 7 this pinkish white
(7.5YR8/3) fine sand was culturally sterile
(Riley 1987:table 2.16).
results
systematic review
Family Pupinidae
Pupina complanata (Pease, 1861)
material: Marshall Islands: Ebon Atoll,
Ebon Islet (“Ebon”), archaeological site
MLEb-2I, sample 1 (3 specimens [spms]);
TP 20, spit 7 (3 spms), spit 8 (1 spm), spit 9
(11 spms), spit 10 (15 spms), spit 11 (14 spms),
spit 12 (14 spms), spit 13 (3 spms), spit 14 (+).
The term “spit” is an arbitrary excavation
Marshall Islands Land Snails · Christensen and Weisler
level, typically 10 cm thick, used to provide
finer stratigraphic control within defined
layers.
remarks: Pupina complanata has been recorded on Ebon and Jaluit ( Jälooj) Atolls in
the Marshall Islands and on Kosrae (formerly
Kusaie), Pohnpei (formerly Ponape), Ulithi,
and Ant Atoll in the Caroline Islands (Pease
1861, Clench 1949, Harry 1966).
Moellendorff (1900:116) contended that
“Ponape is the original habitat of [P. complanata] and that it was introduced on the
different atolls of the Marshall group with
cultivated plants (Pandanus or Musa).” The
species was well represented in pre-Contact
levels of the TP 20 excavations on Ebon, indicating that any translocation as a result of human voyaging, if it occurred at all, must have
taken place prehistorically.
Family Assimineidae
Omphalotropis fragilis (Pease, 1861)
material: Marshall Islands: Maloelap
Atoll, Kaven Islet (“Maloelap”), archaeological site MLMl-3, unit 21, sample 1 (3 spms);
Majuro Atoll, Majuro Islet (“Majuro”), archaeological site MLMj-1, transect 6, TP 3,
north face, Layer I, ∼10 cm bs (6 spms); Majuro, MLMj-1, transect 6, TP 4, north face,
Layer I, 10–15 cm bs (14 spms); Majuro,
MLMj-1, transect 6, TP 5, south face, Layer
IA, ∼10 cm bs (2 spms); Ebon, MLEb-2I,
sample 1 (33 spms); TP 20, spit 6 (71 spms),
spit 7 (11 spms), spit 8 (42 spms), spit 9
(50 spms), spit 10 (21 spms), spit 11 (7 spms),
spit 12 (12 spms), spit 13 (6 spms), spit 14
(3 spms).
remarks: This species was described
from Ebon Atoll (Pease 1861) and has been
reported from Arno, Likiep, Jaluit ( Jälooj),
and Majuro Atolls in the Marshall Islands
(Marshall 1950, Kondo 1961, Reigle 1964,
Wallace and Rosen 1969a, b); it also occurs on
Fais and Ulithi Atoll in the Caroline Islands
(Harry 1966). The species-level taxonomy of
the many Pacific island Omphalotropis is subject to considerable uncertainty, however, and
the relationship of O. fragilis to species occurring outside the Marshall Islands is unclear. In
an unpublished research note dated June 1941
87
held in the Bishop Museum malacology collection, C. Montague Cooke Jr. suggested
that O. fragilis may be conspecific with O. bulimoides (Hombron & Jacquinot, 1854), a species reported by Moellendorff (1900) to occur
on Ruk (= Chuuk), Yap, and Guam.
Omphalotropis fragilis is well represented in
all pre-Contact levels of the Ebon TP 20 and
is thus either indigenous to the island or a
pre-Contact introduction.
Family Truncatellidae
Truncatella guerinii A. & J. B. Villa, 1841
material:
Marshall Islands: Ebon,
MLEb-2I, sample 1 (4 spms); Ebon, MLEb2I, sample 3, buried A horizon (7 spms); TP
20, spit 12 (6 spms), spit 13 (1 spm), spit 14
(1 spm).
remarks: Truncatella guerinii is widely
distributed in the Indo-Pacific region, occurring from East Africa through the islands of
the Indian Ocean, the East Indies, Philippines, and southern Japan eastward through
Melanesia, Micronesia, and Polynesia to the
Society Islands (Clench and Turner 1948).
Kondo (1961) reported its presence on Jaluit
Atoll in the Marshall Islands, and Reigle
(1964) has reported its occurrence on Rongelap (Ronlap) Atoll. Species of Truncatella
live in debris just above the high-water mark;
they lay eggs on land and thus lack pelagic larvae, but they are amphibious snails and are
therefore well adapted for interisland dispersal by rafting, a circumstance believed to account for the wide distribution of most species (Clench and Turner 1948, Solem 1959,
Rosenberg 1996).
Truncatella guerinii was present in small
numbers in the earliest pre-Contact levels of
the TP 20 site on Ebon and is thus either
indigenous to the island or a prehistoric introduction.
Family Achatinellidae
Lamellidea pusilla (Gould, 1847)
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, 144 –154 cm bs (17 spms);
Maloelap, MLMl-3, unit 19, A horizon (1
spm); Maloelap, MLMl-3, unit 19, A horizon
( land snail 10-2 [96-7]) (2 spms); Maloelap,
88
MLMl-3, unit 21, sample 1 (2 spms); Maloelap, MLMl-3, unit 21, sample 4 (6 spms);
Majuro, MLMj-1, transect 6, TP 3, north
face, Layer I, ∼10 cm bs (2 spms); Majuro,
MLMj-1, transect 6, TP4, Layer I, 10–15 cm
bs (32 spms); Majuro, transect 6, TP 4, Layer
II, ∼50 cm bs (49 spms); Majuro, transect 6,
TP 5, south face, Layer IB, ∼50 cm bs (15
spms); Majuro, transect 6, TP 7, Layer VI,
∼200 cm bs (12 spms); Majuro, transect 6, TP
7, Layer VII, ∼240 cm bs (12 spms); Ebon,
MLEb-2I, sample 1 (28 spms); Ebon, MLEb2I, sample 3, buried A horizon (5 spms); TP
20, spit 6 (2 spms), spit 7 (2 spms), spit 8 (2
spms), spit 9 (18 spms), spit 10 (57 spms), spit
11 (5 spms), spit 12 (9 spms), spit 13 (2 spms).
remarks: In the Marshall Islands, Lamellidea pusilla has previously been recorded from
Ebon Atoll (Pease 1861, Pilsbry and Cooke
1915–1916, Cooke and Kondo 1961), Rongelap Atoll (Reigle 1964), Majuro Atoll ( Wallace and Rosen 1969a, b), Enewetak (Änewätak) Atoll (Kay and Johnson 1987), and
Maloelap Atoll ( Weisler 1999). The species
occurs from the Caroline Islands (Kosrae,
Pohnpei, and Chuuk) and Vanuatu eastward
through the Marshall, Ellice, and Gilbert
Islands, Fiji, Samoa, Tonga, and the Cook,
Society, Marquesas, and Tuamotu Islands to
Mangareva in the Gambier Islands (Solem
1959, Cooke and Kondo 1961). Kondo (1975)
synonymized Lamellidea solomonensis Dell,
1955, with L. pusilla, and thus published records of L. solomonensis from the Solomon Islands (Dell 1955, Solem 1960, Clench 1968,
Turner and Clench 1972) refer to L. pusilla.
The western limits of the range of L. pusilla
cannot yet be determined with precision, because the identities of some apparently related
taxa reported from islands to the west of its
known range remain obscure. Kondo (1975)
suggested that the “Tornatellina microstoma,”
reported by Rensch (1937) from the Bismarck
Archipelago, may be L. pusilla. Lamellidea subcylindrica (Quadras & Moellendorff, 1894),
described from Guam in the Mariana Islands,
has also been reported from two small islands
near Java (Benthem Jutting 1941, 1952) and
from the Krakatau Islands (Smith and Djajasasmita 1988); although Cooke and Kondo
(1961) retained L. subcylindrica as a valid spe-
PACIFIC SCIENCE · January 2013
cies, these Indonesian records may also prove
to be referable to L. pusilla.
“There is little doubt that the wide distribution of L. pusilla is due to human agency.
It was probably transported from island to
island during the Polynesian migrations”
(Cooke and Kondo 1961:188). This species
has been reported from pre-European archaeological sites on Tikopia, southeastern
Solomon Islands (Christensen and Kirch
1981, Kirch and Yen 1982); Maloelap Atoll,
Marshall Islands ( Weisler 1999); Ofu Island,
Samoa (Kirch 1993, Hunt and Kirch 1997);
Niuatoputapu Island, Tonga (Kirch 1988 [as
Lamellidea sp. cf. pusilla]); Huahine and Tahiti
in the Society Islands (Sinoto 1983, Orliac
1997); and in the Cook Islands (Allen 1992,
Allen and Christensen 1992, Walter 1998,
Brook 2010, Brook et al. 2010). Specimens
obtained from pre-European levels on Taumako, southeastern Solomon Islands, and reported as Lamellidea subcylindrica by Leach
and Davidson (2008) are probably also referable to L. pusilla, because those authors stated
that their material was probably identical to
L. solomonensis, a synonym of L. pusilla as
noted earlier. The species is represented in
most pre-Contact levels of the TP 20 site on
Ebon and was thus introduced before European contact.
Cooke and Kondo (1961) opined that L.
pusilla probably originated west of the Marshall Islands and was transported eastward by
the Pacific islanders. Although its origins cannot as yet be determined because no fossils
have been recovered from sites predating
human colonization in Oceania, this scenario
is consistent with its presence on Tikopia at
∼900 B.C. (Christensen and Kirch 1981),
many hundreds of years before the Polynesian
settlement of the eastern Polynesian islands it
now inhabits.
Pacificella variabilis Odhner, 1922
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, 144 –154 cm bs (1 spm);
Maloelap, MLMl-3, unit 21, sample 4 (1
spm); Majuro, MLMj-1, transect 6, TP 4,
north face, Layer II, ∼50 cm bs (1 spm); Majuro, MLMj-1, transect 6, TP 5, south face,
Layer IB, ∼50 cm bs (1 spm); Ebon, MLEb-
Marshall Islands Land Snails · Christensen and Weisler
2I, sample 3, buried A horizon (1 spm); TP
20, spit 13 (1 spm), spit 14 (1 spm).
remarks: This species, placed in the genus Tornatellinops by Cooke and Kondo (1961)
but shown to be generically distinct by Climo
(1973), occurs from the Marshall Islands
(Kondo 1961, National Biodiversity Team of
the Republic of the Marshall Islands 2000),
Ellice and Line Islands, Samoa, and Tonga
eastward to the Marquesas, Tuamotus, Gambier Islands, and Easter Island (Cooke and
Kondo 1961). Specimens tentatively identified as this species have also been reported
from Rota, Mariana Islands (Bauman 1996).
As with L. pusilla, there remains some
ambiguity as to the western limits of the range
of P. variabilis; in particular, the identity of
the “Tornatellinops sp.” reported by Mason
(1996:249) from an archaeological site in
Thailand should be investigated, and the affinities of various poorly known species occurring in the Philippines, Indonesia, and
Australia (Cooke and Kondo 1961:170–171)
and in the Mascarene Islands of the Indian
Ocean (Griffiths and Florens 2006:80–81)
should be reexamined.
Pilsbry and Cooke (1933) believed this
species to have been transported interisland
by the Polynesians, and it occurs in preContact archaeological contexts in the Cook
Islands (Brook 2010), the Marquesas Islands
(Rolett 1998 [as “Pacificella sp. cf. variabilis”]),
on Henderson Island (Preece 1998), and on
Easter Island (Kirch et al. 2009). On Henderson Island, however, it occurs also at stratigraphic levels older than initial human colonization of the island, indicating that it is
indigenous there (Preece 1998). If future
studies demonstrate that P. variabilis occurs
naturally only in the islands of easternmost
Polynesia, it would be a rare example of a species being translocated several thousand miles
westward against the flow of Polynesian migration into East Polynesia, but well within
prehistoric postcolonization interaction or
voyaging spheres ( Weisler 1998). In the study
reported here, two specimens were recovered
from the lowest levels of TP 20, site MLEb2I on Ebon, thus establishing its status as a
pre-Contact introduction in the Marshall Islands.
89
Achatinellidae (unidentified)
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, 144 –154 cm bs (7 spms);
Maloelap, MLMl-3, unit 19, A horizon (1
spm); Maloelap, MLMl-3, unit 19, sample 2
(+); Maloelap, MLMl-3, unit 21, sample 1 (+);
Maloelap, MLMl-3, unit 21, sample 4 (5
spms); Majuro, MLMj-1, transect 6, TP 3,
north face, Layer I, ca. 10 cm bs (5 spms); Majuro, MLMj-1, transect 6, TP 4, north face,
Layer I, 10–15 cm bs (15 spms); Majuro,
MLMj-1, transect 6, TP 4, north face, Layer
II, ∼50 cm bs (3 spms); Majuro, MLMj-1,
transect 6, TP 5, Layer IA, ∼10 cm bs (3
spms); Majuro, MLMj-1, transect 6, TP 5,
south face, Layer IB, ∼50 cm bs (1 spm).
remarks: Broken or immature shells of
achatinellid species are commonly encountered in Pacific island archaeological sites and
cannot be more precisely identified than as
“Achatinellidae (unidentified)” (Christensen
and Kirch 1986, Preece 1998), the term used
here.
Family Gastrocoptidae
Gastrocopta pediculus (Shuttleworth, 1852)
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, 144 –154 cm bs (20 spms);
Maloelap, MLMl-3, unit 19, A horizon (1
spm); Maloelap, MLMl-3, unit 19, sample 2
(1 spm); Maloelap, MLMl-3, unit 21, sample
1 (12 spms); Maloelap, MLMl-3, unit 21,
sample 3 (1 spm); Maloelap, MLMl-3, unit
21, sample 4 (8 spms); Majuro, MLMj-1,
transect 6, TP 3, north face, Layer I, ∼10 cm
bs (15 spms); Majuro, MLMj-1, transect 6,
TP 3, north face, Layer II, ∼30 cm bs (1 spm);
Majuro, MLMj-1, transect 6, TP 4, north
face, Layer I, 10–15 cm bs (119 spms);
Majuro, MLMj-1, transect 6, TP 4, north
face, Layer II, ∼50 cm bs (26 spms); Majuro,
MLMj-1, transect 6, TP 5, Layer IA, ∼10 cm
bs (8 spms); Majuro, MLMj-1, transect 6, TP
5, south face, Layer IB, ∼50 cm bs (10 spms);
Majuro, MLMj-1, transect 6, TP 7, Layer VI,
∼200 cm bs (3 spms); Majuro, MLMj-1, transect 6, TP 7, Layer VII, ∼240 cm bs (16 spms);
Ebon, MLEb-2I, sample 1 (22 spms); Ebon,
MLEb-2I, sample 3, buried A horizon (1
spm); TP 20, spit 2 (2 spms).
90
remarks: Gastrocopta pediculus occurs
from the Philippines, Indonesia, and Australia
eastward through Melanesia, Micronesia, and
Polynesia to Hawai‘i and Henderson Island
(Pilsbry 1916–1918, Solem 1959, 1989, 1991,
Preece 1995). In the Marshall Islands, it has
been reported from Ebon Atoll (Pease 1861,
Pilsbry 1916–1918), Jaluit Atoll (Kondo 1961),
Rongelap and Enewetak Atolls (Reigle 1964),
and Majuro Atoll ( Wallace and Rosen 1969a).
Most authors have treated G. pediculus as one
of only two species of Gastrocopta present in
Melanesia, Micronesia, and Polynesia (except
New Zealand) (Pilsbry 1916–1918, Solem
1959, 1989); the other is G. servilis (Gould,
1843), a modern immigrant to the Pacific
(Christensen and Kirch 1986). Pokryszko
(1996), on the other hand, considered Vertigo
pediculus samoensis Mousson, 1865, to be a third
species and separable from G. pediculus. She
has not fully explicated her views, however,
and in the absence of a more definitive analysis we do not distinguish it from G. pediculus.
Pilsbry (1916–1918) believed this characteristic member of the Pacific island “atoll
fauna” to have been transported throughout
Oceania by the Pacific islanders, a view that
has been confirmed by its recovery from preContact archaeological contexts in the Torres
Strait region of Australia (McNiven et al.
2008); eastern Solomon Islands (Christensen
and Kirch 1981, Leach and Davidson 2008);
Reef Islands (Leach and Davidson 2008);
Marshall Islands ( Weisler 1999); Fiji (Hunt
1981); Ofu Island, Samoa (Kirch 1993, Hunt
and Kirch 1997); Niuatoputapu, Tonga
(Kirch 1988); Huahine, Society Islands (Sinoto 1983); Cook Islands (Allen 1992, 1997,
Allen and Christensen 1992, Walter 1998,
Brook 2010, Brook et al. 2010), Marquesas
(Kirch 1973, Rolett 1998); and Henderson Island (Preece 1998).
Gastrocopta pediculus has a long history in
the Pacific islands and was present on Tikopia
by ∼900 B.C. (Christensen and Kirch 1981,
Kirch and Yen 1982), on Taumako by 905–
538 B.C. (Leach and Davidson 2008), and in
Fiji by 2980 ± 90 B.P. (Hunt 1981). Two
specimens were recovered from the uppermost prehistoric level of site MLEb-2I, TP 20
on Ebon Atoll.
PACIFIC SCIENCE · January 2013
Family Vertiginidae
Nesopupa sp.
material: Marshall Islands: Majuro,
MLMj-1, transect 6, TP 4, north face, Layer
I, 10–15 cm bs (1 spm); Ebon, MLEb-2I,
sample 3, buried A horizon (1 spm).
remarks: Members of the genus Nesopupa are widely distributed in Polynesia and
Micronesia, but the non-Hawaiian species
are in need of taxonomic revision. Micronesian taxa recognized by Pilsbry and Cooke
(1915–1916) include N. quadrasi (Moellendorff, 1894) (Guam), N. eapensis (Boettger,
1881) ( Yap), and N. ponapica (Moellendorff,
1900) (Pohnpei).
The identity and affinities of the two specimens of Nesopupa recovered here remain
uncertain; accordingly, Nesopupa sp. must be
regarded as a cryptogenic element in the Marshall Islands fauna.
Family Succineidae
“Succinea” sp.
material:
Marshall Islands: Ebon,
MLEb-2I, TP 20, spit 10 (2 spms); spit 12 (+).
remarks: Excavations on Ebon Atoll
yielded several specimens of the first succineid reported from the Marshall Islands; in the
absence of the anatomical information necessary to allow more precise identification, we
refer to it as “Succinea” sp. Other Micronesian
species include Succinea guamensis Pfeiffer,
1857 (Mariana Islands; also Pohnpei and Koror, Belau [Palau], according to Moellendorff
[1900]), S. quadrasi Moellendorff in Quadras
and Moellendorff (1894) (Mariana Islands),
S. piratarum Quadras & Moellendorff, 1894
(Mariana Islands), and S. philippinica Moellendorff, 1893 ( Yap and Belau [Zilch 1978, Smith
1993, Rundell 2005]). All specimens from the
Ebon, MLEb-2I, TP 20 site were from prehistoric contexts.
Family Subulinidae
Allopeas gracile (Hutton, 1834)
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, 144 –154 cm bs (+); Maloelap, MLMl-3, unit 19, sample 2 (+); Ma-
Marshall Islands Land Snails · Christensen and Weisler
loelap, MLMl-3, unit 21, sample 1 (6 spms);
Maloelap, MLMl-3, unit 21, sample 4 (+);
Majuro, MLMj-1, transect 6, TP 3, north
face, Layer I, ∼10 cm bs (25 spms); Majuro,
MLMj-1, transect 6, TP 3, north face, Layer
II, ∼30 cm bs (2 spms); Majuro, MLMj-1,
transect 6, TP 4, north face, Layer I, 10–15
cm bs (69 spms); Majuro, MLMj-1, transect
6, TP 4, north face, Layer II, ∼50 cm bs (9
spms); Majuro, MLMj-1, transect 6, TP 5,
Layer IA, ca. 10 cm bs (5 spms); Majuro,
MLMj-1, transect 6, TP 5, south face, Layer
IB, ∼50 cm bs (11 spms); Majuro, MLMj-1,
transect 6, TP 7, Layer VI, ∼200 cm bs (5
spms); Majuro, MLMj-1, transect 6, TP 7,
Layer VII, ∼240 cm bs (24 spms); Ebon,
MLEb-2I, sample 1 (14 spms); Ebon, MLEb2I, sample 3, buried A horizon (10 spms); TP
20, spit 6 (21 spms), spit 7 (13 spms), spit 8 (28
spms), spit 9 (97 spms), spit 10 (105 spms),
spit 11 (43 spms), spit 12 (87 spms), spit 13 (35
spms), spit 14 (4 spms).
remarks: Allopeas gracile (previously
known as Lamellaxis gracilis, Opeas gracile,
Opeas oparanum, as well as various other
names) is now a circumtropical species widely
distributed through human commerce; indeed, it has been described as “probably the
most widely distributed land snail in the
world” (Pilsbry 1906–1907:124). It occurs
throughout the Pacific islands (Cowie 1997)
and was first reported from the Marshall Islands (without more specific locality) under
the name Stenogyra juncea by Finsch (1893).
Subsequent authors have reported it from
Jaluit Atoll (Kondo 1961), Rongelap and Eniwetok Atolls (Reigle 1964), and Majuro Atoll
( Wallace and Rosen 1969b).
Allopeas gracile is the anthropophilic land
snail most often recovered from pre-Contact
archaeological sites in Oceania and is virtually
ubiquitous in any such sites that contain land
snails. Records include the eastern Solomon
Islands (Christensen and Kirch 1981, Kirch
and Yen 1982, Leach and Davidson 2008), the
Reef Islands (Leach and Davidson 2008),
New Caledonia (Cowie and Grant-Mackie
2004), Fiji (Hunt 1981), Samoa (Kirch et al.
1990, Kirch 1993, Hunt and Kirch 1997),
Tonga (Kirch 1988, Kirch and Green 2001),
the Cook Islands (Allen 1992, 1997, 1998,
91
Allen and Christensen 1992, Walter 1998,
Brook et al. 2010), the Society Islands (Sinoto
1983, Orliac 1997), the Gambier Islands
(Howard and Kirch 2004, Kirch et al. 2004,
Conte and Kirch 2008, Kirch et al. 2010), the
Marquesas Islands (Rolett 1992, 1998), and
the Hawaiian Islands (Christensen 1984a,
Christensen and Kirch 1986, Kirch 1989,
1992, Burney et al. 2001). Like Gastrocopta
pediculus, A. gracile was present on Tikopia by
900 B.C.; on Taumako by 905–538 B.C.
(Leach and Davidson 2008); and at Yanuca,
Fiji, by 2980 ± 90 B.P. (Hunt 1981). It is
by far the most abundant species recovered
from the prehistoric levels of TP 20, site
MLEb-2I on Ebon, establishing its status as
a pre-Contact introduction in the Marshall
Islands.
Unlike the other translocated species discussed here, A. gracile also has an extensive
paleontological and archaeological record in
the Old World tropics outside the Pacific
Basin. It is known from paleontological sites
of uncertain but presumably Quaternary age
in Pakistan (Theobald 1877, Koken 1903,
Thomas 1981, 1986) and India ( Wadhawan
and Kumar 1996). Specimens provisionally
identified by J. P. E. Morrison as Lamellaxis
gracilis have been obtained from a site in
Egypt (Butzer 1964, Leigh 1968). Archaeologically, the oldest known record for the
species is from the “fourth occupation level”
or “Upper Gamblian pluvial period” of Louis
Leakey’s “Gamble’s Cave II” site in Kenya
(Leakey 1931, Connolly 1931, as Opeas tangaense, a synonym of A. gracile according to
Verdcourt [1983]), believed to have an age of
8500–8000 yr B.P. (Protsch 1978). At Allahdino, Pakistan, it has been recovered from a
Bronze Age site dating from 3980–3690 yr
B.P. (Turnbull 1983 [identification by Alan
Solem]), and at Saar, Bahrain, Glover (1995,
1997 [identification by Fred Naggs]) reported
it from a site dating from the early second
millennium B.C. Finally, Feulner and Green
(2003) and Feulner et al. (2005) gave a date of
about A.D. 400–1300 for the species’ occurrence in a site in the United Arab Emirates.
Naggs (1994) remarked on the limits of our
knowledge of the taxonomy of anthropophilic
subulinids, and no attempt has been made
92
PACIFIC SCIENCE · January 2013
here to verify the accuracy of identifications
published in the archaeological and paleontological literature. As a working hypothesis,
however, it appears that the status of A. gracile
as an invasive species and “the most widely
distributed land snail in the world” has its
roots in antiquity. The geographical origin
of this globally invasive species has been a
matter of speculation. Pilsbry’s suggestion
(1946:178) that A. gracile was “probably indigenous in tropical America” has been accepted
by some (e.g., Solem 1964, Bieler and Slapcinsky 2000; see also Neubert 1998) but has
been questioned by others aware of its presence in archaeological contexts in the Pacific
islands (Christensen and Kirch 1986, Smith
et al. 1990) and in the Middle East (Feulner
and Green 2003, Feulner et al. 2005). Although subulinids have been reported from
archaeological sites in Belize (Hammond and
Miksicek 1981: Lamellaxis sp.; Miksicek et al.
1991: Lamellaxis, Opeas; Carr and Fradkin
2008: Lamellaxis), Honduras (Feldman 1994:
Lamellaxis micra; Wells 2007: Lamellaxis
micra), and Mexico (Morrison and CózatlManzano 2003: Lamellaxis martensi), we know
of no reports of any specimens identified as A.
gracile from a pre-Columbian context in the
Americas. Although it may perhaps be premature to regard the issue as conclusively resolved, we believe that the great weight of
the available evidence indicates that A. gracile
originated in the Old World, not the Neotropics.
Family Helicarionidae
Liardetia (Liardetia) samoensis (Mousson, 1865)
material: Marshall Islands: Majuro,
MLMj-1, transect 6, TP 4, north face, Layer
I, 10–15 cm bs (1 spm); Ebon, MLEb-2I,
sample 1 (7 spms); TP 20, spit 6 (1 spm), spit
7 (1 spm), spit 9 (fragment), spit 10 (11 spms),
spit 11 (1 spm), spit 12 (4 spms).
remarks: This species occurs from the
Bismarck Archipelago, Solomon Islands, New
Hebrides, and New Caledonia eastward
through the Marshall and Ellice Islands, Fiji,
and Samoa to the Cook, Society, and Marquesas Islands (Pease 1861, Baker 1938 (as
Liardetia striolata), Solem 1959, 1989, Reigle
1964, Cowie 2001, Brook 2010). This species
has been reported from pre-Contact archaeological sites in the eastern Solomon Islands
on Tikopia and Taumako (Christensen and
Kirch 1981, Kirch and Yen 1982, Leach and
Davidson 2008), in Samoa (Kirch 1993, Hunt
and Kirch 1997), and on Huahine, Society Islands (Sinoto 1983), and its presence in small
numbers in pre-Contact levels of Ebon site
MLEb-2I, TP 20 indicates that it was introduced to the Marshall Islands prehistorically
as well.
Liardetia (Liardetia) sculpta (Moellendorff,
1883)
material: Marshall Islands: Majuro,
MLMj-1, transect 6, TP 4, north face, Layer
I, 10–15 cm bs (3 spms); Majuro, MLMj-1,
transect 6, TP 5, Layer IA, ∼10 cm bs (+).
remarks: In Micronesia, this species has
been reported to occur on Majuro in the
Marshall Islands ( Wallace and Rosen 1969a,
b), on Guam in the Mariana Islands (Baker
1938), and on Pohnpei and Chuuk in the
Caroline Islands (Baker 1938, Christensen
1984b); in Polynesia, it has been reported
from Rarotonga, Cook Islands (Brook 2010).
It has been reported from Kwangtung and
Macao in China (Baker 1938, Abbott 1950)
and in the Indian Ocean on Cocos Island
(Abbott 1950), the Seychelles ( Verdcourt
1992, Gerlach 2006), and on Mauritius and
Réunion in the Mascarene Islands (Griffiths
and Florens 2006). In the study reported here,
this species was found only in the uppermost
10–15 cm of sediment in two test pits on
Majuro, and thus its presence in the Marshall
Islands before the modern era cannot be demonstrated.
Unidentified
material: Marshall Islands: Maloelap,
MLMl-3, unit 19, A horizon (1 spm); Maloelap, MLMl-3, unit 21, sample 1 (3 spms);
Majuro, MLMj-1, transect 6, TP 3, north
face, Layer I, ∼10 cm bs (5 spms); Majuro,
MLMj-1, transect 6, TP 4, north face, Layer
I, 10–15 cm bs (12 spms).
remarks: A small number of specimens
too immature or fragmentary even for familylevel identification are reported here simply
as “unidentified.”
Marshall Islands Land Snails · Christensen and Weisler
discussion
The Role of Human Translocation
Any biogeographic analysis of the truly terrestrial mollusks of the Marshall Islands (i.e.,
excluding Truncatella and other widely distributed strand-line dwellers such as Assiminea
and members of the family Ellobiidae) must
attempt to distinguish indigenous species
from those that owe their presence to humanmediated translocation, not an easy task. Examination of material from dated archaeological or paleontological sites can demonstrate
that a particular species inhabited an island at
a particular time, but information that a species was absent from an island before initial
human settlement is rarely available. Cooke
and Kondo (1961:202) could state with confidence that Lamellidea oblonga was absent from
the Hawaiian Islands before initial human
colonization because of their unique familiarity with the Pleistocene fossil deposits of
those islands. Elsewhere in the tropical Pacific, however, there are very few published
analyses of fossil assemblages in their entirety
(as distinct from taxonomic treatments of
particular elements of such assemblages) before human arrival of the kind that Brook
(1999a, b, c, 2000) presented for sites in New
Zealand.
Lacking paleontological or archaeological
evidence of the absence of various species
from particular Pacific islands before human
arrival, inferences as to their status must be
drawn from indirect evidence. Various criteria have been proposed for distinguishing
plant and animal species as being either indigenous or introduced in a particular location
(e.g., Webb 1985, Chapman and Carlton
1991, Bean 2007, Hamilton et al. 2010), and
Harry (1964, 1966) addressed the issue in the
specific context of insular nonmarine mollusks. Biogeographical and ecological characteristics indicative of introduced status include an absence of related endemic species in
the area in question, presence in disturbed
habitats, and a known propensity for human
transport in other locations. Genetic analysis
may show dispersal to have occurred recently
(Lee et al. 2007, Ó Foighil et al. 2011), and, of
particular relevance here, archaeological evi-
93
dence may document the chronology of dispersal (Kerney 1966, Christensen and Kirch
1981, Davies 2010). Nevertheless, drawing
the distinction between native and introduced
taxa can be difficult. Species of doubtful status
are “cryptogenic” in the sense of Carlton
(1996), a category that includes several of the
species considered here. The status of others
can be inferred from available information
with a greater or lesser degree of confidence.
None of the sites under study provide the
sequence of multiple well-dated samples that
would be required for a detailed analysis of
faunal change over time; accordingly, we seek
only to identify those species that can be
shown to have inhabited the Marshalls before
the modern era and are thus indigenous to
these islands or were transported to them
by the pre-Contact voyages of the Pacific islanders.
Of the species represented in the material
presented here, Liardetia sculpta may be a
modern introduction, because it is in the
Cook Islands (Brook 2010); it has not yet been
recorded from a prehistoric context anywhere
in Oceania. Lamellidea pusilla, Pacificella variabilis, Gastrocopta pediculus, Allopeas gracile, and
Liardetia samoensis, on the other hand, were
all present in the Marshall Islands prehistorically. Gastrocopta pediculus and A. gracile lack
native congeners elsewhere in Micronesia,
Melanesia, or Polynesia, and A. gracile is noteworthy for its well-established relationship
with human activities (Pilsbry 1946). Their
status as prehistoric introductions seems
clearly established. Lamellidea, Pacificella, and
Liardetia, on the other hand, do have endemic
species in the Pacific islands. The absence of
L. pusilla, P. variabilis, and L. samoensis from
the Marshall Islands before human settlement
has not been demonstrated, and Brook (2010)
regarded them as cryptogenic in the Cook
Islands. Because these species are restricted to
Oceania, they must be native to at least some
of the islands within that area. Preece (1998)
demonstrated that P. variabilis is indigenous
to Henderson Island, but the origins of these
species within that region are otherwise unknown. Given the very limited period of time
(3,000 yr) since the Marshall Islands were
last submerged (Dickenson 2003), however,
94
we believe it is probable that these three
species owe their presence in the Marshall
Islands and their widespread distribution
elsewhere in Oceania to pre-Contact human
transport.
More problematic are Pupina complanata,
Omphalotropis fragilis, Nesopupa sp., and “Succinea” sp. None of these are generally distributed members of the Pacific “atoll fauna” and
(except for “Succinea” sp.) are not closely related to taxa known for their invasive propensities. Nesopupa sp. and “Succinea” sp. have
been reported in the Marshall Islands only
from archaeological contexts; it is unclear
whether this indicates recent extinctions of
the kind reported for birds by Spenneman
(2006) or merely the inadequacy of our knowledge of the islands’ modern land snail fauna.
Pupina complanata and O. fragilis were also
present in the Marshall Islands prehistorically, however, so all four taxa may be either
indigenous or the result of prehistoric translocations.
In general, natural interarchipelagic dispersal of Pacific island land snails is an extremely rare event. Although from 750 to
1,000 species of land snails are native to the
Hawaiian Islands (Solem 1990, Cowie et al.
1995), that highly diverse fauna is believed to
be the result of explosive speciation following
a mere 22–24 original colonization events
in which species from outside the Hawaiian
Islands became established there following
natural over-water dispersal (Zimmerman
1948:104). Although geological evidence indicates the existence of emergent islands in
the vicinity of the present Hawaiian Islands
for at least 32 million years, genetic evidence
suggests that most existing lineages that have
been studied originated within the last 5 million years (Price and Clague 2002). Using 5
million years as a conservative estimate, this
means that a new lineage of land snails has
successfully colonized the Hawaiian Islands
on average about every 208,000 to 227,000 yr.
The Marshall Islands are less isolated from
potential sources of immigrant snails (e.g.,
Pohnpei and Kosrae) than are the Hawaiian
Islands, but in light of the very limited time
available for natural immigration to the Marshalls (ca. 3,000 yr since last submergence) a
PACIFIC SCIENCE · January 2013
major role for human-mediated translocations seems probable.
Patterns of Prehistoric Dispersal
The land snail species that were translocated
to the Marshalls show several distinct patterns
of dispersal, patterns that also characterize
several species occurring elsewhere in Oceania but not present in the Marshalls:
species originating in africa and /or
south asia: Allopeas gracile is the only taxon
clearly demonstrating this pattern, with an
extensive paleontological and archaeological
record in East Africa, the Persian Gulf,
Pakistan, and India in addition to numerous
records from prehistoric archaeological contexts in eastern Melanesia, Micronesia, and
Polynesia. Although we know of no paleontological or archaeological records of the occurrence of Pupisoma orcula (Benson, 1850) (Pupilloidea) in the Old World tropics, its broad
modern range in that region (Pilsbry 1920–
1921, 1922–1926) and its prehistoric presence in the Cook Islands (Brook 2010), Henderson Island (Preece 1998), and possibly also
Hawai‘i (Christensen 1984a, Brook 2010)
suggest that it may eventually prove to have a
similar distribution.
species originating in island southeast asia: Gastrocopta pediculus appears to
have been transported into Oceania prehistorically from Island Southeast Asia. Several
other species not represented in our Marshall
Islands samples have similar distributions.
Discocharopa aperta (Moellendorff, 1888)
(Charopidae) occurs from the Philippines, Indonesia, and northern Australia eastward to
Vanuatu, Fiji, the Kermadec Islands, Samoa,
the Society Islands, and the Austral Islands
(Solem 1983, 1984, 1989). It has been recorded from pre-Contact archaeological contexts in the Torres Strait region of Australia
(McNiven et al. 2008); on Taumako and the
Reef Islands in the eastern Solomons (Leach
and Davidson 2008 [as Discocharopa cf. planulata]); on Niuatoputapu, Tonga (Kirch 1988);
and in the Cook Islands (Brook 2010, Brook
et al. 2010). Wilhelminaia mathildae Preston,
1913 (Helicarionidae), has a similar though
less-extensive distribution (Solem 1959) and
Marshall Islands Land Snails · Christensen and Weisler
has been reported prehistorically from Tikopia, eastern Solomon Islands (Christensen
and Kirch 1981). The prehistoric occurrence
of Costigo saparuana (Boettger, 1891) ( Vertiginidae) in the Cook Islands (Brook 2010)
indicates that it probably belongs in this category as well, because the species is otherwise
known only from Indonesia.
species originating in oceania and
now widely distributed within the
region: Lamellidea pusilla, Pacificella variabilis, and Liardetia samoensis are believed to be
endemic to the Pacific Basin but have become
widely dispersed throughout this region, apparently as a result of prehistoric human
transport. Although the location of their origin within the Pacific islands cannot yet be
established, their presence in the Marshall
Islands is apparently the result of prehistoric
human introduction. Lamellidea oblonga, not
present in the Marshall Islands, also belongs
to this category. The modern range of L. oblonga extends from the Hawaiian Islands west
and south to Fiji and the Ellice Islands and
eastward to the Marquesas, Gambier, and
Austral Islands as well as Pitcairn Island
(Cooke and Kondo 1961). It has been reported from pre-Contact archaeological sites
in the Cook Islands (Allen 1992, 1997, 1998,
Allen and Christensen 1992, Walter 1998,
Brook 2010, Brook et al. 2010), Society Islands (Sinoto 1983, Orliac 1997), Gambier
Islands (Howard and Kirch 2004, Conte and
Kirch 2008), Marquesas Islands (Kirch 1973,
Rolett 1992, 1998), Henderson Island (Preece
1998), and on Kaua‘i in the Hawaiian Islands
(Dixon et al. 1997, Burney 2002) but was
apparently absent from the Hawaiian Islands
before Polynesian settlement (Cooke and
Kondo 1961:202).
species originating in oceania and
translocated to at least one additional archipelago but not widely
distributed: Lee et al. (2007) and Ó
Foighil et al. (2011) demonstrated that two
species of Partula (Partulidae) were translocated between the islands of their origin and a
number of nearby islands, apparently prehistorically. No partulid species has become
generally distributed in Oceania in the manner of the members of the “atoll fauna,” how-
95
ever. Occasional events of interisland translocation of this sort may also account for the
presence of Pupina and Omphalotropis in the
Marshall Islands and should be considered as
well when considering the anomalous occurrence of the endodontid species Australdonta
degagei (Garrett, 1879) on Mauke in the Cook
Islands and on Rurutu and Rimatara in the
Austral Islands (Solem 1976).
The Potential of Land Snail Analysis
in Pacific Archaeology
Although we have cited a substantial number
of reports of the occurrence of land snails
in archaeological sites in the Pacific islands,
most of these appeared as incidental remarks
in studies having a principal focus on other
topics. Nevertheless, the observed widespread
presence of prehistorically introduced land
snail species in such sites suggests that researchers studying introduced vertebrates
such as Rattus exulans or lizards as indicators
of initial human colonization and of the process of settlement (Pregill and Weisler 2007,
Matisoo-Smith 2009) should recognize that
snails can play a similar role. Genetic analysis
of island populations of translocated land
snail species can supplement studies of vertebrates that are intended to elucidate the routes
of prehistoric migration and commerce. As
with birds (Steadman 2006), many Pacific
land snail species have become extinct since
the advent of human settlement in the region
(Christensen and Kirch 1986, Solem 1990,
Abdou and Bouchet 2000, Bouchet and Abdou 2003), and investigations of snails from
archaeological sites can demonstrate the extent and chronology of this process of extinction; it is also critical to document the original
diversity of island faunas, essential to any biogeographic studies.
conclusions
Harry (1966) subtitled his review of the land
mollusks of Ulithi Atoll in the Caroline
Islands “A study of snails accidentally distributed by Man.” The study reported here
demonstrates that human activities, and specifically prehistoric human activities, have had
96
an important influence on the land snail faunas of the Marshall Islands. A review of the
surprisingly rich body of literature on the
topic demonstrates that this is equally true
elsewhere in Oceania, although the effect will
be most evident on atolls and other low islands lacking substantial endemic land snail
faunas. Furthermore, in the case of Allopeas
gracile, the record in Oceania and elsewhere
demonstrates that a species once said to be
“probably the most widely distributed land
snail in the world” (Pilsbry 1906–1907:124)
has a history of association with humans at
least 8,000 yr long.
Unfortunately, land snails in tropical environments have only rarely attracted the serious attention of archaeologists and Quaternary paleontologists, and although several of
the species prehistorically translocated into
Oceania appear to have originated in Island
Southeast Asia, there are few if any records of
their pre-modern occurrence in that critical
source area. We hope that future researchers
in the Philippines, Indonesia, and elsewhere
in that region will expand their studies of
translocated vertebrates to include the analysis of those land snail species that may share a
similarly long-standing history of association
with human activities.
Unlike continental and high volcanic islands that have great age and corresponding
environmental and biotic diversity, the hundreds of low-lying Pacific atolls found across
Oceania exhibit the most precarious and
relatively recent landscapes that support an
impoverished biota. This canvas provides a
unique opportunity to investigate the human
translocation of plants and vertebrates, and, as
we have demonstrated here, land snails are an
important part of these efforts.
Although the hundreds of low-lying atolls
found across Oceania are comparable in age
to the continental and high volcanic islands of
the region, until several thousand years ago
the atolls were submerged and thus uninhabitable by terrestrial organisms (Dickenson
2003). Accordingly, they lack the environmental and biotic diversity of their higher
neighbors. Their simpler ecology and the
shorter time span within which their terrestrial biota has developed may, however, make
PACIFIC SCIENCE · January 2013
them particularly suitable for studies attempting to distinguish the effects of natural and
human-mediated dispersal.
acknowledgments
We thank Carmen Bigler, Hemley Benjamin,
and Clary Makroro for their efforts to make
the archaeological fieldwork projects run
smoothly. M.I.W. also acknowledges support
from the Deputy Vice Chancellor (Research),
University of Queensland. Tom Riley collected sediment samples during his 1979 excavations on Majuro Atoll, and Tom Dye
assisted in making them available to us. We
greatly appreciate the Bishop Museum for curating this material since then. This is contribution 2012-010 from the Pacific Biological
Survey at the Bishop Museum.
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