Land Snails from Archaeological Sites in the Marshall Islands, with Remarks on Prehistoric Translocations in Tropical Oceania

Carl C. Christensen; Marshall I. Weisler

This study looks deeply at the atoll, beginning with its young geology, its specific biogeography, its early archaeology, and its ecology. From these different sciences assumptions can be made about its land snail fauna. Natural colonizers had to be highly salt-resistent and able to arrive quickly, but most would be introduced by gardening activities. By careful sampling from the surface of the village islet, from the outer islets, and from column samples throughout its stratigraphy one can distinguish which species are natural, which inhabit the gardens in the poor soil, and which inhabit the gardens in the good soil. One can use these inferences to make statements about the environment of the past. And, in small ways, one can see how the land transformed from a low marginal island into a vital oasis full of splendid people.

s 12 Call for Papers 15 Conferences and Events 16 Member News and Updates 18 Recent Publications 18 In This Issue ARCHAEO + MALACOLOGY GROUP NEWSLETTER Issue 27, February 2017

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 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. 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. Literature Cited Abbott, R. T. 1950. The molluscan fauna of the Cocos-Keeling Islands, Indian Ocean. Bull. Rafﬂes Mus. 22:68–98. Abdou, A., and P. Bouchet. 2000. Nouveaux gastéropodes Endodontidae et Punctidae (Mollusca, Pulmonata) récemment étients de l’archipel des Gambier (Polynésie). Zoosystema 22:689–707. Allen, M. S. 1992. Dynamic landscapes and human subsistence: Archaeological investigations on Aitutaki Island, southern Cook Islands. Ph.D. diss., University of Washington, Seattle. ———. 1997. Coastal morphogenesis, climatic trends, and Cook Island prehistory. Pages 124 –146 in P. V. Kirch and T. L. Hunt, eds. Historical ecology in the Paciﬁc islands: Prehistoric environmental and landscape change. Yale University Press, New Haven, Connecticut. ———. 1998. Holocene sea-level change on Aitutaki, Cook Islands: Landscape change and human response. J. Coastal Res. 14:10–22. Allen, M. S., and C. C. Christensen. 1992. Landsnail analysis. Pages 490–497 in M. S. Allen. Dynamic landscapes and human subsistence: Archaeological investigations on Aitutaki Island, southern Cook Islands. Marshall Islands Land Snails · Christensen and Weisler Ph.D. diss., University of Washington, Seattle. Anderson, A. 2009. The rat and the octopus: Initial human colonization and the prehistoric introduction of domestic animals to Remote Oceania. Biol. Invasions 11:1503– 1519. Anderson, E. 1952. Plants, man and life. University of California Press, Berkeley. Baker, H. B. 1938. Zonitid snails from Paciﬁc islands. Part 1. Southern genera of Microcystinae. Bernice P. Bishop Mus. Bull. 158:1–102, pl. 1–20. Bauman, S. 1996. Diversity and decline of land snails on Rota, Mariana Islands. Am. Malacol. Bull. 12:13–27. Bean, A. R. 2007. A new system for determining which plant species are indigenous in Australia. Aust. Syst. Bot. 20:1–43. Benthem Jutting, W. S. S. van. 1941. Nonmarine Mollusca from the satellite islands surrounding Java. Arch. Neerl. Zool. 5:251–348. ———. 1952. Systematic studies on the nonmarine Mollusca of the Indo-Australian Archipelago. III. Critical revision of the Javanese pulmonate land-snails of the families Ellobiidae to Limacidae, with an appendix on Helicarionidae. Treubia 21:291–435. Bieler, R., and J. Slapcinsky. 2000. A case study for the development of an islands fauna: Recent terrestrial mollusks of Bermuda. Nemouria 44:1–100. Bouchet, P., and A. Abdou. 2003. Endemic landsnails from the Paciﬁc islands and the museum record: Documenting and dating the extinction of the terrestrial Assimineidae of the Gambier Islands. J. Molluscan Stud. 69:165–170. Brook, F. J. 1999a. Stratigraphy and landsnail faunas of Late Holocene coastal dunes, Tokerau Beach, northern New Zealand. J. R. Soc. N. Z. 29:337–359. ———. 1999b. Stratigraphy, landsnail faunas, and paleoenvironmental history of coastal duneﬁelds at Te Werahi, northernmost New Zealand. J. R. Soc. N. Z. 29:361–393. ———. 1999c. Stratigraphy, landsnail faunas, and paleoenvironmental history of Late Holocene coastal dunes, Tauroa Penin- 97 sula, northern New Zealand. J. R. Soc. N. Z. 29:395–405. ———. 2000. Holocene distribution, ecology and local extinction of the endemic New Zealand dune snail Succinea archeyi Powell (Stylommatophora: Succineidae). J. R. Soc. N. Z. 30:209–225. ———. 2010. Coastal landsnail fauna of Rarotonga, Cook Islands: Systematics, diversity, biogeography, faunal history, and environmental inﬂuences. Tuhinga 21:161–252. Brook, F. J., R. K. Walter, and J. A. Craig. 2010. Changes in the terrestrial molluscan fauna of Miti‘aro, southern Cook Islands. Tuhinga 21:75–98. Burney, D. A. 2002. Late Quaternary chronology and stratigraphy of twelve sites on Kaua‘i. Radiocarbon 44:13–44. Burney, D. A., H. F. James, L. P. Burney, S. L. Olson, W. Kikuchi, W. L. Wagner, M. Burney, D. McCloskey, D. Kikuchi, F. V. Grady, R. Gage II, and R. Nishek. 2001. Fossil evidence for a diverse biota from Kaua‘i and its transformation since human arrival. Ecol. Monogr. 71:615–641. Butzer, K. W. 1964. Pleistocene palaeoclimates of the Kurkur Oasis, Egypt. Can. Geogr. 8:125–141. Carlton, J. 1996. Biological invasions and cryptogenic species. Ecology 77:1653– 1655. Carr, H. S., and A. Fradkin. 2008. Animal resource use in ecological and economic contact at Formative Period Cuello, Belize. Quat. Int. 191:144 –153. Chapman, J. W., and J. T. Carlton. 1991. A test of criteria for introduced species: The global invasion by the isopod Synidotea laevidorsalis (Miers, 1881). J Crustacean Biol. 11:386–400. Christensen, C. C. 1984a. Analysis of nonmarine mollusks. Pages 355–376 in R. Schilt, Subsistence and conﬂict in Kona, Hawai‘i: An archaeological study of the Kuakini Highway Realignment Corridor. Department of Anthropology, Bernice P. Bishop Museum, Departmental Report Series 84-1. ———. 1984b. Nonmarine mollusks from site TKM-11, Moen. Pages 512–519 in 98 T. F. King and P. L. Parker, Pisekin Nóómw Nóón Tonaachaw: Archaeology in the Tonaachaw Historic District, Moen Island. Micronesian Archaeology Survey Report No. 18. Southern Illinois University at Carbondale, Center for Archaeological Investigations, Occasional Paper 3. Christensen, C. C., and P. V. Kirch. 1981. Nonmarine mollusks from archaeological sites on Tikopia, southeastern Solomon Islands. Pac. Sci. 35:75–88. ———. 1986. Nonmarine mollusks and ecological change at Barbers Point, O‘ahu, Hawai‘i. Bishop Mus. Occas. Pap. 26:52– 80. Clench, W. J. 1949. Cyclophoridae and Pupinidae of Caroline, Fijian, and Samoan Islands. Bernice P. Bishop Mus. Bull. 196. ———. 1968. Additions of the land and freshwater Mollusca of Rennell Island, Solomon Islands. Nat. Hist. Rennell Isl. Br. Solomon Isl. 5:173–179. Clench, W. J., and R. D. Turner. 1948. A catalogue of the family Truncatellidae with notes and descriptions of new species. Occas. Pap. Mollusks Mus. Comp. Zool. Harv. Univ. 1:157–212. Climo, F. C. 1973. The systematics, biology and zoogeography of the land snail fauna of Great Island, Three Kings Group, New Zealand. J. R. Soc. N. Z. 3:565–627. Connolly, M. 1931. The Mollusca from deposits of Gamblian pluvial date, Kenya Colony. Pages 276–277 in L. S. B. Leakey, The Stone Age cultures of Kenya Colony. Cambridge University Press, Cambridge. Conte, E., and P. V. Kirch. 2008. One thousand years of human environmental transformation in the Gambier Islands (French Polynesia). Terra Australis 29:253–264. Cooke, C. M., Jr. 1928. Notes on Paciﬁc land snails. Proc. 3rd Pan-Pac. Sci. Congr., Tokyo, 30 Oct.–11 Nov. 1926, 2:2276–2284. Cooke, C. M., Jr., and Y. Kondo. 1961. Revision of Tornatellinidae and Achatinellidae (Gastropoda, Pulmonata). Bernice P. Bishop Mus. Bull. 221. Cowie, R. H. 1996. Paciﬁc island land snails: Relationships, origins and determinants of diversity. Pages 347–372 in A. Keast and S. E. Miller, The origin and evolution of PACIFIC SCIENCE · January 2013 Paciﬁc island biotas, New Guinea to eastern Polynesia: Patterns and processes. SPB Academic Publishing, Amsterdam. ———. 1997. Catalog and bibliography of the nonindigenous nonmarine snails and slugs of the Hawaiian Islands. Bishop Mus. Occas. Pap. 50:1–66. ———. 2001. Invertebrate invasions on Paciﬁc islands and the replacement of unique native faunas: A synthesis of the land and freshwater snails. Biol. Invasions 3:119– 136. ———. 2005. Alien non-marine mollusks in the islands of the tropical and subtropical Paciﬁc: A review. Am. Malacol. Bull. 20:95–103. Cowie, R. H., N. L. Evenhuis, and C. C. Christensen. 1995. Catalog of the native land and freshwater mollusks of the Hawaiian Islands. Backhuys Publishers, Leiden. Cowie, R. H., and J. A. Grant-Mackie. 2004. Land snail fauna of Mé Auré Cave ( WMD007), Moindou, New Caledonia: Human introductions and faunal change. Pac. Sci. 58:447–460. Cowie, R. H., K. A. Hayes, C. T. Tran, and W. M. Meyer III. 2008. The horticultural industry as a vector of alien snails and slugs: Widespread invasions in Hawaii. Int. J. Pest Manage. 54:267–276. Cowie, R. H., and D. G. Robinson. 2003. Pathways of introduction of nonindigenous land and freshwater snails and slugs. Pages 93–122 in G. Ruiz and J. T. Carlton, eds. Invasive species: Vectors and management strategies. Island Press, Washington, D.C. Craig, J. 1995. The role of landsnails in Paciﬁc archaeology. M.A. thesis, University of Otago, Dunedin, New Zealand. Davies, P. 2010. Land and freshwater molluscs. Pages 175–180 in T. O’Connor and N. Sykes, eds. Extinctions and invasions: A social history of the British fauna. Windgather Press, Oxford. Dell, R. K. 1955. A tornatellinid land mollusk from the Solomon Islands. Pac. Sci. 9:357– 358. Dickinson, W. R. 2003. Impact of midHolocene hydro-isostatic highstand in re- Marshall Islands Land Snails · Christensen and Weisler gional sea level on habitability of islands in Paciﬁc Oceania. J. Coastal Res. 19:489– 502. Dixon, B., D. Soldo, and C[harles] C. Christensen. 1997. Radiocarbon dating land snails and Polynesian land use on the island of Kaua‘i, Hawai‘i. Hawaii. Archaeol. 5:52–62. Evans, J. G. 1972. Land snails in archaeology. Seminar Press, London. Feldman, L. H. 1994. The mollusks of Copan. Pages 477–479 in G. R. Willey, R. M. Leventhal, A. A. Demarest, and W. L. Fash Jr., eds. Ceramics and artifacts from excavations in the Copan Residential Zone. Pap. Peabody Mus. Archaeol. Ethnol. Harv. Univ. 80. Feulner, G. R., and S. A. Green. 2003. Terrestrial molluscs of the United Arab Emirates. Mitt. Dtsch. Malakozool. Ges. 69/70:23–34. Feulner, G. R., E. Neubert, and S. Green. 2005. Land snails. Pages 222–227 in P. Hellyer and S. Aspinall, eds. The Emirates: A natural history. Trident Press, London. Finsch, O. 1893. Ethnologische Erfahrungen und Belegstuࡇ cke aus der Suࡇ dsee. III. Abteilung: Mikronesien. II. MarshallArchipel. III. Carolinen (1. Kuschai, 2. Ponape). Ann. Naturhist. Hofmus. Wien 8:119–275. Fosberg, F. R. 1990. A review of the natural history of the Marshall Islands. Atoll Res. Bull. 330. Garrett, A. 1879. List of land shells inhabiting Rurutu, one of the Austral Islands, with remarks on their synonymy, geographical range, and descriptions of new species. Proc. Acad. Nat. Sci. Phila. 31:17–30. Gerlach, J. 2006. Terrestrial and freshwater Mollusca of the Seychelles Islands. Backhuys, Leiden. Glover, E. 1995. Molluscan evidence for diet and environment at Saar in the early second millennium BC. Arab. Archaeol. Epigr. 6:157–179. ———. 1997. Food remains: Molluscs. Pages 83–88 in H. Crawford, R. Killick, and J. Moon, eds. The Dilmun Temple at Saar: Bahrain and its archaeological inheritance. Keegan Paul International, London. 99 Grifﬁths, O. L., and V. F. B. Florens. 2006. A ﬁeld guide to the non-marine molluscs of the Mascarene Islands (Mauritius, Rodrigues and Réunion) and the Northern Dependencies of Mauritius. Bioculture Press, Mauritius. Hamilton, A. M., G. R. Zug, and C. C. Austin. 2010. Biogeographic anomaly or human introduction: A cryptogenic population of tree skink (Reptilia: Squamata) from the Cook Islands, Oceania. Biol. J. Linn. Soc. 100:318–328. Hammond, N., and C. H. Miksicek. 1981. Ecology and economy of a formative Maya site at Cuello, Belize. J. Field Archaeol. 8:259–268. Harry, H. W. 1964. The foreign freshwater snails now established in Puerto Rico. Annu. Rep. Am. Malacol. Union 1964:4 –5. ———. 1966. Land snails of Ulithi Atoll, Caroline Islands: A study of snails accidentally distributed by Man. Pac. Sci. 20:212– 223. Hayward, B. W., and F. J. Brook. 1981. Exploitation and redistribution of ﬂax snail (Placostylus) by the prehistoric Maori. N. Z. J. Ecol. 4:33–36. Howard, N. M., and P. V. Kirch. 2004. Zooarchaeological analysis of faunal assemblages. Pages 106–121 in E. Conte and P. V. Kirch, eds. Archaeological investigations in the Mangareva Islands (Gambier Archipelago), French Polynesia. University of California, Berkeley, Archaeological Research Facility, Contribution No. 62. Hunt, T. L. 1981. New evidence for early horticulture in Fiji. J. Polynesian Soc. 90:259–266. Hunt, T. L., and P. V. Kirch. 1997. The historical ecology of Ofu Island, American Samoa, 3000 B.P. to the present. Pages 105–123 in P. V. Kirch and T. L. Hunt, eds. Historical ecology in the Paciﬁc islands: Prehistoric environmental and landscape change. Yale University Press, New Haven, Connecticut. Kay, E. A., and S. Johnson. 1987. Mollusca of Enewetak Atoll. Pages 105–147 in D. M. Devaney, E. S. Reese, B. L. Burch, and P. Helfrich, eds. The natural history of Enewetak Atoll. Vol. 2. Biogeography and 100 systematics. U.S. Department of Energy, Oak Ridge, Tennessee. Kerney, M. P. 1966. Snails and Man in Britain. J. Conchol. 26:3–14. Kirch, P. V. 1973. Prehistoric subsistence patterns in the northern Marquesas Islands, French Polynesia. Archaeol. Phys. Anthropol. Oceania 8:24 –40. ———. 1984. The evolution of the Polynesian chiefdoms. University of California Press, Berkeley. ———. 1988. Niuatoputapu: The prehistory of a Polynesian chiefdom. Thomas Burke Mem. Mus. Monogr. 5. ———. 1989. Non-marine molluscs from the rockshelter sediments. Pages 73–82 in P. V. Kirch, ed. Prehistoric Hawaiian occupation of the Anahulu Valley, O‘ahu Island: Excavations in three inland rockshelters. University of California, Berkeley, Archaeological Research Facility Contribution no. 47. ———. 1992. Anahulu: The anthropology of history in the Kingdom of Hawaii: Vol. 2. The archaeology of history. University of Chicago Press, Chicago. ———. 1993. Non-marine molluscs from the To‘aga Site sediments and their implications for environmental change. Pages 115–121 in P. V. Kirch and T. L. Hunt, eds. The To‘aga Site: Three millennia of Polynesian occupation in the Manu‘a Islands, Samoa. University of California, Berkeley, Archaeological Research Facility Contribution no. 51. Kirch, P. V., C. C. Christensen, and D. W. Steadman. 2009. Subfossil land snails from Easter Island, including Hotumatua anakenana, new genus and species (Pulmonata: Achatinellidae). Pac. Sci. 63:105–122. Kirch, P. V., J. Coil, M. I. Weisler, M. E. Conte, and A. J. Anderson. 2004. Radiocarbon dating and site chronology. Pages 94 –105 in E. Conte and P. V. Kirch, eds. Archaeological investigations in the Mangareva Islands (Gambier Archipelago), French Polynesia. University of California, Berkeley, Archaeological Research Facility Contribution no. 62. Kirch, P. V., E. Conte, W. Sharp, and C. Nickelsen. 2010. The Onemea Site (Tara- PACIFIC SCIENCE · January 2013 vai Island, Mangareva) and the human colonization of southeastern Polynesia. Archaeol. Oceania 45:66–79. Kirch, P. V., and R. C. Green. 2001. Hawaiiki, ancestral Polynesia: An essay in historical anthropology. Cambridge University Press, Cambridge. Kirch, P. V., T. L. Hunt, L. Nagaoka, and J. Tyler. 1990. An ancestral Polynesian occupation site at To‘aga, Ofu Island, American Samoa. Archaeol. Oceania 25:1–15. Kirch, P. V., D. W. Steadman, V. L. Butler, J. Hather, and M. I. Weisler. 1995. Prehistory and human ecology in eastern Polynesia: Excavations at Tangatatau Rockshelter, Mangaia, Cook Islands. Archaeol. Oceania 30:47–65. Kirch, P. V., and D. E. Yen. 1982. Tikopia: The prehistory and ecology of a Polynesian outlier. Bernice P. Bishop Mus. Bull. 238. Koken, E. 1903. Das Diluvium im Gebiete der Saltrange (nordwestliches Indien). Centralbl. f. Mineral. Geol. Palaeontol. 1903:433–439. Kondo, Y. 1961. Gastropod mulluscs [sic] collected by J. L. Gressitt. Page 105 in D. I. Blumenstock. A report on typhoon effects upon Jaluit Atoll. Atoll Res. Bull. 75. ———. 1975. Status of Lamellidea solomonensis Dell (Stylommatophora: Achatinellidae: Lamellideinae). Malacol. Rev. 8:114. Leach, F., and J. Davidson. 2008. Archaeology on Taumako, a Polynesian outlier in the eastern Solomon Islands. N. Z. J. Archaeol. Spec. Publ. Leakey, L. S. B. 1931. The Stone Age cultures of Kenya Colony. Cambridge University Press, Cambridge. Lee, T., J. B. Burch, T. Coote, B. Fontaine, O. Gargominy, P. Pearce-Kelly, P., and D. Ó Foighil. 2007. Prehistoric interarchipelago trading of Polynesian tree snails leaves a conservation legacy. Proc. R. Soc. Lond. B Biol. Sci. 274:2907–2914. Leigh, E. G., Jr. 1968. Fossil Mollusca from the Kurkur Oasis. Pages 513–514 in K. W. Butzer and C. L. Hansen, Desert and river in Nubia: Geomorphology and prehistoric environments at the Aswan Reservoir. University of Wisconsin Press, Madison. Marshall Islands Land Snails · Christensen and Weisler Marshall, J. T., Jr. 1950. Vertebrate ecology of Arno Atoll, Marshall Islands. Atoll Res. Bull. 3. Mason, G. M. 1996. The micromolluscs. Pages 239–264 in G. B. Thompson, The excavation of Khok Phanom Di, a prehistoric site in central Thailand. Vol. 4. Subsistence and environment: The botanical evidence (The biological remains, Part II). Reports of the Research Committee of the Society of Antiquaries of London, no. LIII. Matisoo-Smith, E. 2009. The commensal model for human settlement of the Paciﬁc 10 years on—What can we say and where to now? J. Isl. Coastal Archaeol. 4:151– 163. McNiven, I. J., J. Crouch, M. Weisler, N. Kemp, L. C. Martinez, J. Stanisic, L. Brady, S. Hocknull, and W. Boles. 2008. Tigershark Rockshelter (Baidamau Mudh): Seascapes and settlement reconﬁguration on the sacred islet of Pulu, western Zenadh Kes (Torres Strait). Aust. Archaeol. 66:15– 32. Merlin, M., A. Capelle, T. Keene, J. Juvik, and J. Maragos. 1997. Keinikkan Im Meļan Aelön Kein: Plants and environments of the Marshall Islands. 2nd ed. East-West Center, Honolulu. Miksicek, C. H., E. S. Wing, and S. J. Scudder. 1991. The ecology and economy of Cuello: The natural and cultural landscape of Preclassic Cuello. Pages 70–84 in N. Hammond, ed. Cuello: An early Maya community in Belize. Cambridge University Press, Cambridge. Moellendorff, O. F. von. 1900. The land shells of the Caroline Islands. J. Malacol. 7:101–126. Morrison, B. A., and R. Cózatl-Manzano. 2003. Initial evidence for use of periphyton as an agricultural fertilizer by the ancient Maya associated with the El Edén wetland, northern Quintana Roo, Mexico. Pages 401–413 in A. Gómez-Pompa, M. F. Allen, S. L. Fedick, and J. J. Jiménez-Osornio, eds. The Lowland Maya area: Three millennia at the human-wildland interface. Food Products Press, New York. Naggs, F. 1994. The reproductive anatomy of Paropeas achatinaceum and a new con- 101 cept of Paropeas (Pulmonata: Achatinoidea: Subulinidae). J. Molluscan Stud. 60:175– 191. National Biodiversity Team of the Republic of the Marshall Islands. 2000. The Marshall Islands—Living atolls amidst the living sea: The National Biodiversity Report of the Republic of the Marshall Islands. RMI Biodiversity Project, Majuro. Neubert, E. 1998. Annotated checklist of the terrestrial and freshwater molluscs of the Arabian Peninsula with descriptions of new species. Fauna Arabia 17:333–461. Ó Foighil, D., T. Lee, and J. Slapcinsky. 2011. Prehistoric anthropogenic introduction of partulid tree snails in Papua New Guinean archipelagos. J. Biogeogr. 38:1625–1632. Orliac, M. 1997. Human occupation and environmental modiﬁcations in the Papeno‘o Valley, Tahiti. Pages 200–229 in P. V. Kirch and T. L. Hunt, eds. Historical ecology in the Paciﬁc islands: Prehistoric environmental and landscape change. Yale University Press, New Haven, Connecticut. Pease, W. H. 1861. Descriptions of six new species of land shells, from the islands of Ebon, Marshalls’s group, in the collection of Mr. Hugh Cuming. Proc. Zool. Soc. Lond. 1860:439–440. Pilsbry, H. A. 1900. The genesis of MidPaciﬁc faunas. Proc. Acad. Nat. Sci. Phila. 52:568–581. ———. 1906–1907. Manual of conchology. 2nd Ser. Vol. 18. Achatinidae: Stenogyrinae and Coeliaxinae. Academy of Natural Sciences, Philadelphia. ———. 1916. Mid-Paciﬁc land snail faunas. Proc. Natl. Acad. Sci. U.S.A. 2:429–433. ———. 1916–1918. Manual of conchology. 2nd Ser. Vol. 24. Pupillidae (Gastrocoptinae). Academy of Natural Sciences, Philadelphia. ———. 1920–1921. Manual of conchology. 2nd Ser. Vol. 26. Pupillidae ( Vertiginidae, Pupillinae). Academy of Natural Sciences, Philadelphia. ———. 1922–1926. Manual of conchology. 2nd ser. Vol. 27. Pupillidae (Orculinae, Pagodulinae, Acanthinulinae, etc.). Academy of Natural Sciences, Philadelphia. 102 ———. 1946. Land Mollusca of North America (north of Mexico). Monogr. Acad. Nat. Sci. Phila. 3. Vol. 2, pt. 1. Pilsbry, H. A., and C. M. Cooke Jr. 1915– 1916. Manual of conchology. 2nd Ser. Vol. 23. Appendix to Amastridae. Tornatellinidae. Index, vols. xxi–xxiii. Academy of Natural Sciences, Philadelphia. ———. 1933. Notes on the land snail family Tornatellinidae. Nautilus 42:59–62. Pokryszko, B. M. 1996. The Gastrocoptinae of Australia (Gastropoda: Pulmonata: Pupilloidea). Invertebr. Taxon. 10:1085–1150. Porch, N. 2011. Prehistoric human impact on the neglected majority: Evidence for human-moderated translocation and catastrophic extinction in the Paciﬁc insect fauna. Page 64 in Proceedings of conference on Evolution of Life on Paciﬁc Islands and Reefs: Past, Present, and Future, 26–30 May 2011, Honolulu, Hawai‘i. Powell, A. W. B. 1938. The Paryphantidae of New Zealand, No. IV and the genus Placostylus in New Zealand. Rec. Auckl. Inst. Mus. 2:141–150. Preece, R. C. 1995. Systematic review of the land snails of the Pitcairn Islands. Biol. J. Linn. Soc. 56:273–307. ———. 1998. Impact of early Polynesian occupation on the land snail fauna of Henderson Island, Pitcairn Group (South Paciﬁc). Philos. Trans. R. Soc. Lond. B Biol. Sci. 353:347–368. Pregill, G. K., and M. I. Weisler. 2007. Lizards from prehistoric sites on Ebon Atoll, Marshall Islands. Micronesica 39:107–116. Price, J. P., and D. A. Clague. 2002. How old is the Hawaiian biota? Geology and phylogeny suggest recent divergence. Proc. R. Soc. Lond. B Biol. Sci. 269:2429–2435. Protsch, R. 1978. The chronological position of Gamble’s Cave II and Bromhead’s Site (Elmenteita) of the Rift Valley, Kenya. J. Hum. Evol. 7:101–109. Reigle, N. J. 1964. Nonmarine mollusks of Rongelap Atoll, Marshall Islands. Pac. Sci. 18:126–129. Rensch, I. 1937. Systematische und tiergeographische Untersuchungen uࡇ ber die Landschneckenfauna des Bismarck-Archipels. II. Arch. f. Naturg. n.f. 6:526–644. PACIFIC SCIENCE · January 2013 Riley, T. J. 1987. Archaeological survey and testing, Majuro Atoll, Marshall Islands. Pages 169–270 in T. Dye, ed. Marshall Islands archaeology. Pac. Anthropol. Rec. no. 38. Bernice P. Bishop Museum, Honolulu. Rolett, B. V. 1992. Faunal extinctions and depletions linked with prehistory and environmental change in the Marquesas Islands (French Polynesia). J. Polynesian Soc. 101:86–94. ———. 1998. Hanamiai: Prehistoric colonization and cultural change in the Marquesas Islands, East Polynesia. Yale Univ. Publ. Anthropol. no. 81. Rosenberg, G. 1996. Independent evolution of terrestriality in Atlantic truncatellid gastropods. Evolution 50:682–693. Rosendahl, P. H. 1987. Report I: Archaeology in eastern Micronesia: Reconnaissance survey in the Marshall Islands. Pages 17– 168 in T. S. Dye, ed. Marshall Islands archaeology. Pac. Anthropol. Rec. 38. Bernice P. Bishop Museum, Honolulu. Rundell, R. J. 2005. The land snails of Belau: Survey of the 16 states. http://www .friendsoftobi.org/misc /documentarchive/ rundelllandsnails2005.pdf. Sinoto, Y. H. 1983. Archaeological excavations of the Vaito‘otia and Fa‘ahia Sites on Huahine Island, French Polynesia. Natl. Geogr. Soc. Res. Rep. 15:583–599. Smith, A. G., W. B. Miller, C. C. Christensen, and B. Roth. 1990. Land Mollusca of Baja California, Mexico. Proc. Calif. Acad. Sci. 47:95–158. Smith, B. D. 1993. Working list of the terrestrial gastropods of Palau, Caroline Islands. Dickinson Memorial Mollusc Collection, Marine Laboratory, University of Guam, Mangilao. Smith, B. J., and M. Djajasasmita. 1988. The land mollusks of the Krakatau Islands, Indonesia. Philos. Trans. R. Soc. Lond. B Biol. Sci. 322:379–400. Solem, A. 1959. Systematics and zoogeography of the land and fresh-water Mollusca of the New Hebrides. Fieldiana Zool. 43. ———. 1960. Non-marine Mollusca from the Florida Islands, Solomon Islands. J. Malacol. Soc. Aust. 4:39–56. Marshall Islands Land Snails · Christensen and Weisler ———. 1964. New records of New Caledonian nonmarine mollusks and an analysis of the introduced mollusks. Pac. Sci. 18:130–137. ———. 1976. Endodontoid land snails from Paciﬁc islands (Mollusca: Pulmonata: Sigmurethra). Part I. Family Endodontidae. Field Museum Press, Chicago. ———. 1983. Endodontoid land snails from Paciﬁc islands (Mollusca: Pulmonata: Sigmurethra). Part II. Families Punctidae and Charopidae, zoogeography. Field Museum Press, Chicago. ———. 1984. Small land snails from northern Australia. III. Species of Helicodiscidae and Charopidae. J. Malacol. Soc. Aust. 6:155–179. ———. 1989. Non-camaenid land snails of the Kimberley and Northern Territory, Australia. I. Systematics, afﬁnities and ranges. Invertebr. Taxon. 2:455–604. ———. 1990. How many Hawaiian land snail species are left? And what we can do for them. Bishop Mus. Occas. Pap. 30:27– 40. ———. 1991. Distribution and diversity patterns of Australian pupilloid land snails (Mollusca: Pulmonata: Pupillidae, s. l.). Veliger 34:233–252. Spenneman, D. H. R. 2006. Extinctions and extirpations in Marshall Islands avifauna since European contact: A review of historic evidence. Micronesica 38:253–266. Steadman, D. W. 2006. Extinction and biogeography of tropical Paciﬁc birds. University of Chicago Press, Chicago. Stone, B. C. 1960. The wild and cultivated Pandanus of the Marshall Islands. M.A. thesis, University of Hawai‘i at Mänoa, Honolulu. Storey, A. A., A. C. Clarke, and E. A. Matisoo-Smith. 2011. Identifying contact with the Americas: A commensal-based approach. Pages 111–138 in T. L. Jones, A. A. Storey, E. A. Matisoo-Smith, and J. M. Ramírez-Aliaga, eds. Polynesians in America: Pre-Columbian contacts with the New World. Altamira Press, Lanham. Stuiver, M., P. J. Reimer, and R. W. Reimer. 2005. CALIB 5.0. http://radiocarbon.pa .qub.ac.uk /calib/. 103 Theobald, W. 1877. On the occurrence of erratics in the Potwar, and the deductions that must be drawn therefrom. India Geol. Surv. Rec. 10:140–143. Thomas, K. D. 1981. Palaeoecological studies in the Bannu Basin: The sources of evidence. Pages 227–232 in H. Hlrtel, ed. South Asian archaeology 1979. Dietrich Reimer Verlag, Berlin. ———. 1986. Environment and subsistence in the Bannu Basin. BAR Int. Ser. 310:13– 33. Turnbull, P. F. 1983. The study of clams and snails from archaeological sites. BAR Int. Ser. 183:53–65. Turner, R. D., and W. J. Clench. 1972. Land and freshwater Mollusca of Savo Island, Solomons, with anatomical descriptions (Mollusca, Gastropoda). Steenstrupia 2:207–232. Verdcourt, B. 1983. A list of the non-marine Mollusca of East Africa (Kenya, Uganda, Tanzania, excluding Lake Malawi). Achatina 11:200–239. ———. 1992. The identity of two euconuloid snails recently found in the Seychelles. J. Conchol. 34:169–174. Wadhawan, S. K., and V. Kumar. 1996. Subsurface Quaternary aeolian stratigraphy in the Ghaggar Basin of Thar Desert, India. J. Arid Environ. 32:37–51. Wallace, G. D., and L. Rosen. 1969a. Experimental infection of Paciﬁc island mollusks with Angiostrongylus cantonensis. Am. J. Trop. Med. Hyg. 18:13–19. ———. 1969b. Studies on eosinophilic meningitis. V. Molluscan hosts of Angiostrongylus cantonensis on Paciﬁc islands. Am. J. Trop. Med. Hyg. 18:206–216. Walter, R. 1998. Anai‘o: The archaeology of a fourteenth century Polynesian community in the Cook Islands. N. Z. Archaeol. Assoc. Monogr. 22. Webb, D. A. 1985. What are the criteria for presuming native status? Watsonia 15:231– 236. Weisler, M. I. 1998. Hard evidence for prehistoric interaction in Polynesia. Curr. Anthropol. 39:521–532. ———. 1999. The antiquity of aroid pit agriculture and signiﬁcance of buried A 104 horizons on Paciﬁc atolls. Geoarchaeology 14:621–654. ———. 2002. Archaeological survey and test excavations on Ebon Atoll, Republic of the Marshall Islands. Report prepared for the Historic Preservation Ofﬁce, Republic of the Marshall Islands, and on ﬁle there. Weisler, M. I., H. Yamano, and Q. Hua. 2012. A multi-disciplinary approach for dating human colonization of Paciﬁc atolls. J. Isl. Coastal Archaeol. 7 (1): 102–125. Wells, E. C. 2007. Faenas, ferias, and ﬁestas: Ritual ﬁnance in ancient and modern Hon- PACIFIC SCIENCE · January 2013 duras. Pages 29–65 in C. E. Wells and K. L. Davis-Salazar, eds. Mesoamerican ritual economy: Archaeological and ethnological perspectives. University Press of Colorado, Boulder. Zilch, A. 1978. Die Typen und Typoide des Natur-Museums Senckenberg, 60: Mollusca: Succineacea. Arch. Molluskenkd. 109:109–136. Zimmerman, E. C. 1948. Insects of Hawaii. Vol. 1. Introduction. University of Hawai‘i Press, Honolulu.

RELATED PAPERS

RELATED TOPICS

Log In

Land Snails from Archaeological Sites in the Marshall Islands, with Remarks on Prehistoric Translocations in Tropical Oceania 1

Land Snails from Archaeological Sites in the Marshall Islands, with Remarks on Prehistoric Translocations in Tropical Oceania 1

Related Papers

RELATED PAPERS

RELATED TOPICS