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A Refined Protocol for Calculating MNI in Archaeological Molluscan Shell Assemblages: a Marshall Islands Case Study

A Refined Protocol for Calculating MNI in Archaeological Molluscan Shell Assemblages: a Marshall Islands Case Study

Accepted Manuscript

A refined protocol for calculating MNI in archaeological molluscan shell assemblages: A Marshall Islands case study

Matthew Harris, Marshall Weisler, Patrick Faulkner

PII: S0305-4403(15)00030-8 DOI: 10.1016/j.jas.2015.01.017 Reference: YJASC 4322

To appear in: Journal of Archaeological Science

Received Date: 2 June 2014 Revised Date: 27 January 2015 Accepted Date: 30 January 2015

Please cite this article as: Harris, M., Weisler, M., Faulkner, P., A refined protocol for calculating MNI in archaeological molluscan shell assemblages: A Marshall Islands case study, Journal of Archaeological Science (2015), doi: 10.1016/j.jas.2015.01.017.

This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT 1 A refined protocol for calculating MNI in archaeological molluscan shell assemblages: 2 A Marshall Islands case study 3 Matthew Harris*1, Marshall Weisler 1 and Patrick Faulkner 2 4 1. School of Social Science, The University of Queensland, St Lucia, Queensland, 4072, 5 Australia 6 2. School of Philosophical and Historical Inquiry, Faculty of Arts and Social Sciences, 7 Department of Archaeology, University of Sydney, Sydney, New South Wales, 2006, 8 Australia 9 * Correspondence to: Matthew Harris, School of Social Science, University of Queensland, 10 St Lucia, Queensland, 4072, Australia. E-mail: [email protected] 11 12 Abstract

13 Comprehensive and transparent protocols for calculating Minimum Number of Individuals 14 (MNI) for archaeological faunal assemblages are critical to data quality, comparability, and 15 replicability. MNI values for archaeological molluscan assemblages are routinely calculated 16 by counting a select range of Non-Repetitive Elements (NREs). Most commonly, only the 17 frequency of the of gastropods and the or hinge of bivalves are recorded. 18 Calculating MNI based only on the frequency of these NREs can underestimate the relative 19 abundance of particular molluscan shell forms. Using archaeological mollusc assemblages 20 from two sites in the Marshall Islands as a case study, we outline a new protocol (tMNI) that 21 incorporates a wider range of NRE and calculates MNI based on the most frequently 22 occurring NRE for each taxon. The principles that underlie the tMNI method can be modified 23 to be regionally or assemblage specific, rather thaMANUSCRIPTn being a universally applicable range of 24 NRE for the calculation of MNI. For the Marshall Islands assemblages, the inclusion of 25 additional NRE in quantification measures led to (1) a 167% increase in relative abundance 26 of gastropods and 3% increase in bivalves (2) changes to rank order abundance, and (3) 27 alterations to measures of taxonomic richness and evenness. Given these results for the 28 Marshall Islands assemblages, tMNI provides more accurate taxonomic abundance measures 29 for these and other archaeological molluscan assemblages with similar taxa. These results 30 have implications for the quality of zooarchaeological data increasingly utilised by 31 conservation biologists, historical ecologists and policy makers. 32 33 Keywords: mollusc quantification, Minimum Numbers of Individuals (MNI), shell middens, 34 Marshall Islands, analytical transparency 35 36 Highlights ACCEPTED 37 • Molluscan MNI values are routinely calculated using a restricted range of Non- 38 Repetitive Elements (NREs) 39 • A new protocol is described that expands the range of NREs for MNI calculation 40 • The new protocol for calculating MNI increased abundance of gastropods by 167% 41 • Richness and evenness measures, as well as rank order were markedly influenced by 42 quantification protocol 43

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ACCEPTED MANUSCRIPT 44 1. Introduction

45 Quantification is fundamental to zooarchaeological analyses, and comprehensive analytical 46 protocols are critical to ensure data quality, comparability, and replicability. As Wolverton 47 (2013:381) noted, it is essential that archaeologists undertake high-quality faunal analyses as 48 zooarchaeological data is increasingly utilised by conservation biologists, historical 49 ecologists, and policy makers (e.g. Augustine and Dearden 2014, Gobalet 2011, Carder and 50 Crock 2012, Erlandson and Fitzpatrick 2006, Groesbeck et al . 2014, Wake et al. 2013). 51 52 The relative merits of various faunal quantification methods and the analytical and 53 interpretive implications for using Minimum Numbers of Individuals (MNI), Number of 54 Identifiable Specimens (NISP) and/or weight have been widely discussed in the 55 zooarchaeological literature, (Claassen1998:106, 2000, Giovas 2009, Glassow 2000, Grayson 56 1979, Gutiérrez-Zugasti 2011, Lyman 2008:21-140, Mason et al. 1998, Reitz and Wing 57 1999:202-213). Post-depositional leaching of from molluscan shell and 58 differential rates of fragmentation within and between taxa can bias NISP values and weight 59 measures of mollusc shell from archaeological deposits. As such, many analysts working on 60 assemblages of invertebrate taxa use MNI to potentially provide a more accurate measure of 61 taxonomic abundance (Ballbè 2005, Mannino and Thomas 2001, Nunn et al. 2007, Poteate 62 and Fitzpatrick 2013, Robb and Nunn 2013, Seeto et al. 2012). 63 64 MNI values for molluscan remains are most commonly calculated by counting the frequency 65 of a restricted number of Non-Repetitive Elements (NREs), such as the spire of gastropods or 66 the umbo and hinge of bivalves (Allen 2012, Ballbè MANUSCRIPT 2005, Claassen 1998:106, Chicoine and 67 Rojas 2013, Mannino and Thomas 2001, Mason et al. 1998, Ono and Clark 2012, Poteate and 68 Fitzpatrick 2013, Seeto et al. 2012). This method, however, has the potential to consistently 69 underrepresent some taxa (Giovas 2009). Differential fragmentation resulting from inter- 70 taxonomic variability in shell architecture, morphology, and human processing of mollusc 71 shell can lead to the under-representation of particular shell forms; especially gastropods that 72 lack robust, readily identifiable spires. 73 74 We propose that current MNI calculation protocols utilising only a restricted range of NRE 75 can influence measures of taxonomic abundance, richness, evenness, and dominance, 76 potentially affecting reconstructions of human behaviour derived from archaeological 77 material. This hypothesis is tested using two assemblages of tropical Indo-Pacific mollusc 78 shells from two prehistoric habitation sites on Ebon Atoll, Marshall Islands. The assemblage 79 was quantified ACCEPTED using current MNI calculation protocols and a new, more comprehensive 80 method for calculating molluscan MNI that utilises a wider range of molluscan NREs and 81 calculates MNI based on the most frequently occurring NRE per taxon. Each quantification 82 protocol is tested by comparing relative abundance and rank order abundance. Alterations to 83 taxonomic richness, evenness, and diversity are measured using the number of taxa 84 (NTAXA) Simpson’s index of Diversity (1-D), Shannon’s Evenness (E), and The Shannon- 85 Weiner index of Diversity ( H’ ). 86

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ACCEPTED MANUSCRIPT 87 Results for the Marshall Islands assemblages analysed indicate that current methods for 88 calculating MNI routinely underrepresent gastropod abundance. Furthermore, the application 89 of the new MNI protocol resulted in increased richness and evenness values for all taxa. 90 Given the importance of molluscs for human subsistence and ecosystem maintenance, it is 91 critical to consider the influence of quantification measures on inferences of human 92 behaviour and long-term impacts to the environment. Analysts should employ quantification 93 methods that result in the most precise and accurate measures of relative abundance. 94 Diversity indices, biomass estimates, and mean trophic level are common measures used by 95 zooarchaeologists and fisheries scientists for examining human impacts to marine 96 environments (Carder and Crock 2012, Cinner 2014, Morrison and Hunt 2007, Perry et al. 97 2011, Wake et al. 2013), and the calculation of these indices using abundance data derived 98 from MNI protocols which utilise only a restricted number of NREs could be misleading or 99 erroneous. The compounding error introduced by such protocols could further bias 100 interpretations and reconstructions of changes in prehistoric subsistence practices and impact 101 modern management of marine resources. 102 103 2. Current methods of MNI calculation in mollusc shell assemblages

104 Common protocols for calculating MNI in molluscan assemblages are described in Mason et 105 al. (1998:308-309) and Claassen (1998:106). These protocols lack the sufficient descriptive 106 detail required (e.g., minimum criteria for determining NRE completeness) to ensure 107 accuracy, precision, and replicability of MNI calculations that would allow analysts to 108 reliably compare datasets. The quantification protocol outlined by Mason et al. (1998:309- 109 309), calculates MNI by counting only a limited MANUSCRIPT range of predetermined shell features, 110 emphasising the frequency of gastropod spires and bivalve hinge and umbones. Moreover, 111 Mason et al. (1998:308) assert that analysts should only identify those fragments that 112 contribute toward MNI. Claassen (1998:106) proposes similar methods for MNI calculation 113 including the quantification of gastropod umbilici. Only the pre-selected NREs are identified 114 and quantified for both methods, regardless of the presence of other molluscan NREs that 115 could increase MNI and influence diversity and richness measures. While mollusc shell 116 fractures in ways that are somewhat predictable (Harris 2011, Koppel 2010, Vermeij 1979, 117 Zuschin et al 2003), to determine the element or feature to be counted prior to analysis of the 118 archaeological material assumes that there is no variation in fragmentation or preservation at 119 either the assemblage or taxon level. Differential fragmentation or variation in processing 120 techniques could influence preservation of the pre-selected NRE across taxa, potentially 121 under-representing gastropod taxa that lack thick shells and durable spires, and bivalve shells 122 lacking easily identifiable,ACCEPTED robust umbones and hinges (see also Giovas 2009, Glassow 2000). 123 124 An alternative method proposed by Giovas (2009: Fig. 3) calculates MNI using a part-scoring 125 system based on the presence or absence of portions of mollusc shell (see also Gutiérrez- 126 Zugasti 2011). This method produced ‘no uniform pattern that holds across all taxa and 127 samples’ (Giovas 2009:1560) and remains a valid alternative to NRE MNI. However, NRE 128 based MNI may be less subjective, as NRE are discrete, easily identified features compared 129 to large zones of mollusc shell (Lyman 2008:277). Furthermore, the identification of mollusc

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ACCEPTED MANUSCRIPT 130 shell is often based on diagnostic NRE, and the quantification method proposed here may 131 expedite the quantification process compared to part-scoring methods. 132 133 The method outlined below, referred to herein as tMNI, refines and extends upon the NRE 134 MNI method by: (1) including a wider range of NREs for counting, (2) recording element 135 frequency by taxon, and (3) calculating MNI after the highest frequency NRE is identified for 136 each taxon. Ideally, analysts should calculate MNI by stratigraphic layers, rather than 137 arbitrary aggregates which inflate MNI (Grayson 1984). This method is derived from 138 “Traditional MNI” (Giovas 2009:1558), described originally by White (1953) and elaborated 139 by Grayson (1979, 1984), using a range of well-defined molluscan NREs that often preserve 140 in archaeological deposits rather than only hinges, umbones, and spires. The utilisation of an 141 expanded range of NREs reduces the inherent bias towards particular forms (i.e. overall 142 shape) of mollusc shell that occurs when the NRE for the calculation of MNI is selected prior 143 to analysis. While some researchers have implemented this method, or similar methods, (e.g. 144 Allen 2012, Kataoka 1996, Szabó 2009, Rosendahl 2012), no formal protocol has been 145 outlined in detail for the determination of MNI using a wide range of NREs. 146 147 3. Site description

148 The southernmost atoll in the Marshall Islands, Ebon Atoll (4˚38’N, 168˚42’E) consists of 22 149 islets encircling a 104 km 2 lagoon. Typical of Marshall Islands atolls, prehistoric villages are 150 marked by concentrations of large marine molluscs (Tridacna spp., Conus spp., 151 spp.), shell artefacts, and coral gravel spread to form pavements. Villages are situated parallel 152 to and just inland from the lagoon shore and often MANUSCRIPT run most of the length of the larger islets. 153 TP 18 and 19 (Figure 1: a), forming a 1 x 2 m unit, were excavated ~25 m from the lagoon 154 beach and ~30m west of the council house on a small hill at site MLEb-1 (Rosendahl 155 1987:83; Weisler 1999: Figure 4, 2002:20). Continuous cultural deposits were encountered to 156 a depth of 1.10 m. At the northeast portion of Ebon Atoll, Enekoion islet (1 km long and 300 157 m wide) has one major village site (MLEb-33) where TP7 (Figure 1: b) (1m 2) was excavated 158 100 m from the lagoon shore just inland of a large swampy gardening area. A cultural layer, 159 ~35 cm thick, was encountered. 160 161 Site deposits consist of coralline sands with a neutral pH; consequently, shell is well 162 preserved and minimal evidence of shell degradation via dissolution was noted during routine 163 taxonomic identification. Furthermore, in contrast to many mollusc shell assemblages from 164 middens and mounds (Perry and Hoppa 2011 cf. Weisler 2001:Table 7.7), the assemblages 165 analysed here areACCEPTED both taxonomically rich (many present) and diverse (many taxa 166 represented by many individuals, rather than the assemblage dominated by a single taxon 167 [e.g. Faulkner 2009]). The richness and diversity of the samples will be discussed further in 168 section 5.2. 169

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ACCEPTED MANUSCRIPT 170 4. Methods

171 4.1 A refined MNI protocol for calculating MNI 172 tMNI is calculated by the methods outlined below. All shell should be identified to the lowest 173 taxonomic level and quantified by NISP and/or weight to be broadly comparable with other 174 studies where MNI was not calculated (e.g. Amesbury 1999, Weisler 2001). Here, all 175 specimens were quantified by NISP and weight in grams to two decimal places. The use of a 176 range of quantification measures in concert can highlight the strengths and weaknesses of 177 each method and are useful for addressing additional questions, such as issues relating to 178 taphonomy (see Faulkner 2013, Lyman 2008, Grayson 1984). 179 180 To ensure accurate identification, specimens that could not be confidently assigned to species 181 were only identified to or family, despite having similar morphology to dominant taxa 182 (Szabó 2009:186). All identifications were completed to the lowest possible taxonomic level 183 using modern Indo-Pacific focussed comparative collections held at the University of 184 Queensland archaeology laboratory. Various reference manuals were also consulted, 185 including: Abbott and Dance (1982) Lamprell and Healy (2006), Poppe (2008), and Röckel et 186 al . (1995). For consistency, all taxonomic names follow the World Register of Marine 187 Species (http://www.marinespecies.org). A review of gastropod and features is 188 presented in the following section, followed by a description of the NREs used to quantify 189 archaeological mollusc remains from two prehistoric habitation sites on Ebon Atoll, Marshall 190 Islands. 191 4.2 Gastropod and bivalve shell features MANUSCRIPT 192 There are five major classes of the : the (e.g., whelks and winkles), 193 (e.g., and mussels), Cephalopoda (e.g., , , Nautilidae and 194 ), Polyplacophora (), and Scaphopoda (tusk shells). Only the Gastropoda and 195 Bivalvia are discussed in detail here, as along with the Polyplacophora, they are often the 196 main classes of molluscs recovered from archaeological sites. 197 198 Gastropod shells (Figure 2) most commonly consist of a hollow cone coiled around a central 199 axis, known as the (Stachowitsch 2002, Vermeij 1993, Pechenik 2005). Each full 200 revolution of the shell around the columella is larger than the last as additional shell is 201 secreted along the growth margin (Rupert et al. 2004). This combination of shell coiling and 202 enlargement produces the typical gastropod morphology of a coiled cone with a pointed top 203 (posterior end) and a large opening at the anterior end. Shell architecture (e.g. micro and 204 macrostructure, ACCEPTED teeth, nodes, spines, and ribs) and morphology varies depending on shell 205 growth rate, the overlap of whorls, tightness of coiling, and angle of the in relation to 206 the horizontal plane (Vermeij 1993) (Figure 3). Additionally, environmental and ontogenetic 207 factors can also influence shell architecture and form (Rhoads and Lutz 1980). 208 209 Bivalve shells (Figure 4) consist of two dorsally hinged, articulating valves that enclose the 210 (Gosling 2003:1). Shell shape can be classified broadly depending on the symmetry of 211 pairs and individual valve symmetry. The terms equivalve and inequivalve describe the

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ACCEPTED MANUSCRIPT 212 symmetry of one valve in relation to the other (Pechenik 2005). Lateral symmetry (equilateral 213 or inequilateral) describes the symmetry of the anterior and posterior portion of individual 214 valves (Stachowitsch 2002). Bivalve form and architecture are determined by the way that the 215 animal lays down shell along the ventral margin and surface, influenced during the 216 life of the animal by a range of ecological and ontogenetic factors (Gosling 2003:7, Rhoads 217 and Lutz 1980) (Figure 5). 218 219 A range of morphological elements are defined as NRE. The location of each NRE on the 220 shell and qualitative minimum criteria for counting fragmented NREs are described in the 221 following sections. If multiple fragments can be refitted to form a complete NRE they should 222 be counted as one NRE. The location and descriptions of NREs are drawn from Stachowitsch 223 (2002). 224 4.3 Additional NRE 225 The NREs described below are useful for calculating MNI in archaeological assemblages of 226 tropical marine molluscs from the Marshall Islands. A wider range of NREs may be useful 227 for quantifying other Indo-Pacific molluscan shell assemblages. We recommend that all 228 additional NREs be reported in publication including a clear definition that includes 229 minimum identification and quantification criteria. 230 231 4.4 Calculating Gastropod MNI 232 Gastropod NREs that were used in the quantification of Marshall Islands molluscan remains 233 are the (1) spire (2) anterior notch or canal (3)MANUSCRIPT posterior notch or canal (4) outer (5) 234 aperture (6) , and (7) (Figure 6). A quantification key is provided that 235 illustrates examples of our tMNI method for the quantification of gastropod fragments of 236 different shell forms (Figure 7).

237 4.4.1 The spire 238 The spire consists of the and all shell whorls except the body . The 239 protoconch is the shell laid down at the larval stage and can usually be differentiated from all 240 other whorls by a difference in and smaller size. In molluscan shell from 241 archaeological deposits—if present—the protoconch is generally eroded so that the 242 lines at the intersection of whorls are muted or diminished. For this NRE to be counted, 243 greater than 50% of the —consisting of the smallest whorls of the spire, and if preserved 244 or present, the protoconch —must be present. 245 4.4.2 The anteriorACCEPTED notch or canal 246 The anterior notch is located on the anterior side of the aperture, at the base of the columella. 247 Many taxa feature extended cylindrical forms of this NRE, known as canals (e.g., the 248 subfamily, Muricinae). Anterior canals often fragment in archaeological deposits. 249 To account for post-depositional alteration, the minimum criterion for counting the anterior 250 canal or notch NRE is that the base of the columella is present. Additionally, some taxa (such 251 as the Neritidae) lack anterior notches, and additional NRE should be described and included 252 to account for this variability between taxa.

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ACCEPTED MANUSCRIPT 253 4.4.3 The posterior notch or canal 254 In taxa where posterior notches occur, the NRE is morphologically similar to the anterior 255 notch, but occurs on the posterior margin of the aperture, rather than the anterior margin. Like 256 anterior notches, these NRE can occur as extended cylindrical protrusions, such as the 257 posterior canals of the tropical frog shell Bursa bufonia. More than 50% of the posterior 258 notch or canal must be present for this NRE to be counted.

259 4.4.4 The outer lip 260 The outside edge of the aperture is known as the outer lip. The outer lip can be ornamented 261 with tooth-like protrusions known as denticles (e.g. some Neritidae) or thickened (e.g., 262 , ). The outer lip NRE can be counted if more than 50% is present. The 263 location and morphology of denticles that ornament the outer lip is useful for this 264 determination.

265 4.4.5 The aperture 266 The opening of the shell at the terminal margin (anterior end) of the coiled cone is the 267 aperture. Apertural morphology is variable, with some taxa (e.g., , , 268 Architectonidae) lacking anterior notches or canals. An aperture is counted as whole when 269 the following elements are present: the outer lip, the inner lip (the edge of the aperture that is 270 attached to the columella, often ornamented with folds or plaits), and depending on the 271 specific taxon morphology, the anterior and posterior canals or notches. While this NRE 272 duplicates counts of other NRE noted above, recording aperture frequency contributes to 273 studies of taphonomy (e.g. Szabó 2012) and meat extraction (e.g. Somerville-Ryan 1998). MANUSCRIPT 274 4.4.6 The operculum 275 The aperture of most marine gastropods is capped by a flexible proteinaceous or rigid 276 calcified disk known as the operculum. In most cases proteinaceous opercula (e.g. the 277 opercula of Terebralia spp. and Strombus spp.) will not be preserved in archaeological 278 deposits; however, calcified opercula regularly preserve. The opercula of turban shells ( 279 spp.) are ubiquitous in Pacific island archaeological sites (e.g. Allen 1992, Morrison and 280 Hunt 2007, Walter 1998, Szabó 2009). An operculum may be counted if the nucleus can be 281 confidently identified.

282 4.4.7 The umbilicus 283 If the shell is loosely coiled around the columella an umbilicus (if open, rather than closed) 284 may be present as a small depression or cavity on the base of the columella side of the body 285 whorl. The umbilicus may contribute to MNI only if more than 50% of the element remains. 286 The umbilicus NREACCEPTED limits the underrepresentation of taxa that lack anterior canals or notches, 287 but may be influenced by intraspecific variation in umbilicus form. For example, some 288 members of the genus Turbo exhibit individual variation in umbilicus form (Carpenter & 289 Niem 1998:412-416), which may influence MNI values based on this NRE.

290 4.4.8 Gastropod NRE that are excluded from MNI 291 There are several NRE that are excluded from contributing to MNI calculations as they are 292 highly variable across or within taxa, are strongly influenced by shell age, or are too difficult

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ACCEPTED MANUSCRIPT 293 to confidently identify in fragmented archaeological remains. These NRE are the parietal lip 294 and wall, the columella, and exterior shell sculpture. The columella is excluded as a distinct 295 NRE as this element is incorporated with the spire, anterior canal, and inner lip NREs. Any 296 exterior shell sculpture such as ribs, threads, nodules, and spines are also not considered to be 297 a robust and discrete NRE here. It is important to note that the range NREs outlined here are 298 provided to demonstrate a range of NRE that were useful for the Marshall Islands 299 assemblage, and should be modified to suit the diverse assemblages, taxa, and taphonomic 300 processes which are encountered in the analysis of molluscan remains globally. 301 4.5 Calculating bivalve MNI 302 Bivalve NREs are the: (1) umbo and (2) anterior portion of the hinge (3) posterior 303 portion of the hinge (4) anterior adductor muscle scar, and (5) posterior adductor muscle scar 304 (Figure 8). A quantification key is provided for the Bivalvia (Figure 9). 305 306 All bivalve NREs must be sided in order to be quantified. There are several simple means of 307 siding valves. When teeth are present, they interdigitate on each opposing valve so that where 308 there is a projection on one valve, there will be a socket on the other. The valve side that 309 contains the teeth or sockets is regular and can be used to side valves for quantification. In 310 addition, valve hinges of inequilateral valves are often morphologically distinct on the 311 anterior and posterior sides. 312 313 Adductor muscle scars (See Section 4.5.3) can also be used to side valves. These scars are 314 either equal (isomyarian) or of different sizes (anisomyarian). In some taxa, such as 315 Spondylus spp., only one muscle scar is present MANUSCRIPT (monomyarian). For monomyarian taxa 316 valves are sided according to the presence or absence of a muscle scar. In isomyarian taxa, 317 scar morphology can indicate valve side. Anisomyarian taxa can be sided by comparing the 318 relative size (accounting for shell growth) of muscle scars. Additionally, valves can generally 319 be sided by the prominent direction of the umbo. For prosogyrate taxa where the umbo leans 320 to the anterior, when the shell is placed with the dorsal (exterior) surface facing upward, if the 321 umbo points to the right the valve is from the right side of the shell. A left-pointing umbo 322 indicates a left valve. For opisthogyrate taxa where the umbo leans toward the posterior, the 323 siding procedure is reversed.

324 4.5.1 The umbo and beak 325 Like the spire of gastropods, the umbo and beak are formed during the earliest growth stages 326 of the animal. The beak is present as a small, outwardly protruding feature just above the 327 hinge. The stronglyACCEPTED curved portion of the shell that is laid down after the beak is known as 328 the umbo. The umbo can be distinguished by tightly spaced concentric growth bands that can 329 be muted due to pre and post-depositional erosion. The distinctive morphology of the 330 concavity inside the umbo and beak, however, can be used to confidently identify this NRE 331 which can be counted if more than 50% of the beak is present.

332 4.5.2 The hinge 333 The articulating surfaces which are ventral to the umbo on the interior of the shell valve are 334 known as the valve hinges or hinge. Along with the umbo, the hinge is most commonly

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ACCEPTED MANUSCRIPT 335 preserved in archaeological deposits. The hinge is divided into anterior and posterior NRE at 336 the ventral projection of the beak. By treating the hinge as two NRE in this way, the chances 337 of distinguishing a single individual from fragmented remains are increased and the bias 338 towards taxa with durable umbones is reduced. 339 340 The dentition generally present on the hinge of heterodont (complex hinges with a small 341 number of distinct teeth varying in size and shape) bivalves are classified as either cardinal or 342 lateral teeth. Cardinal teeth are found on the centre of the hinge, ventral to the umbo. Lateral 343 teeth are generally present in the form of ridges that lie anteriorly and posteriorly of the 344 cardinal teeth. Cardinal and lateral teeth can be used to determine if more than 50% of the 345 NRE is present. A hinge NRE (anterior or posterior portion) that is determined to be 50% 346 complete can be counted. Inter-taxonomic variation in the arrangement and number of teeth 347 present on the hinge does occur (such as the taxodont Arcidae, with many alternating teeth 348 and sockets present on the hinge), and quantification methods must be evaluated and refined 349 based on the morphology of taxa using similar methods to those described above.

350 4.5.3 The adductor muscle scars 351 The parts of the live animal responsible for the opening and closing of the shell valves are 352 known as adductor muscles and hinge ligaments. Hinge ligaments rarely preserve 353 archaeologically, but the adductor muscles leave negative impressions or scars on the interior 354 of the shell valve that can be identified in archaeological remains and are considered NREs. 355 Each muscle scar (anterior and posterior) is counted as a single NRE. The minimum criteria 356 for identifying and counting the adductor muscle scar NRE is the presence of more than 50% 357 of the ventral margin of the scar. MANUSCRIPT 358

359 4.5.4 NRE Excluded 360 Cardinal teeth, lateral teeth, and marginal teeth are excluded as NRE due to their inclusion in 361 the hinge NREs. Marginal teeth are also excluded as it is problematic to determine when 362 more than 50% of these features are present in fragmented remains. The and sinus 363 are excluded as these features are often removed by post-depositional alteration of bivalve 364 shell and are difficult to confidently identify. 365 366 4.6 MNI Calculation 367 The frequency of each NRE is recorded as an integer for each taxon in the assemblage. The 368 most frequently occurring NRE for each gastropod taxon is the MNI. For bivalves, only 369 NREs that can beACCEPTED sided are used for quantification. The most frequently occurring NRE for 370 the valve side (left or right) with the highest count is used. Only bivalve NREs that can be 371 confidently assigned a valve side are counted toward MNI. 372 373 The quantification protocol outlined above highlights the range of distinct NRE that appear 374 on mollusc shells. Importantly for this study, a range of NRE other than the commonly used 375 spire are equally suitable for quantification, but not currently widely utilised in gastropod 376 MNI calculation protocols (e.g. Claassen 1998, Mason 1988). For bivalves, the adductor

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ACCEPTED MANUSCRIPT 377 muscle scars of molluscs can be useful NREs, but are also not routinely used. The following 378 case study highlights the benefits of using these methods for the quantification of molluscan 379 shell. 380 381 To compare quantification methods MNI was calculated based on the most frequently 382 occurring NRE for each taxon at each assemblage using the quantification method outlined 383 above (tMNI) and routine quantification protocols using the spire of gastropods and the umbo 384 and hinge of bivalves (NRE MNI), with the anterior and posterior portion counting as distinct 385 NRE. All mollusc NRE counts were aggregated at the pit level prior to the calculation of 386 MNI. All NRE described in section 3.3 are used for tMNI calculations. Additionally, for the 387 , the base and labum adjacent to the aperture were treated as separate NRE (Figure 388 10: a) Similarly, for the nerites ( Nerita spp.), an additional NRE was utilised that counted the 389 frequency of the distinctive whorls on the interior of the shell where the columellar deck joins 390 the outer lip, essentially replacing the posterior and anterior canal NRE for these taxa (Figure 391 10: b: I, I).. While MNI is influenced by aggregation effects (Grayson 1984, 1979), site level 392 totals for all taxa highlight the influence of each calculation method on overall abundance, 393 richness, and diversity. In addition, by examining the values for bivalves and gastropods 394 separately, the influence of shell morphology on MNI value can be clearly established. 395 396 4.7 Testing the influence of quantification protocol 397 To assess the influence of MNI calculation protocols on relative abundance, MNI values 398 derived using NRE MNI was compared with MNI values calculated using tMNI. The 399 resulting rank order abundance of taxa for each MANUSCRIPT quantification protocol is compared, and 400 alterations to the top ten ranked taxa are presented. Any observed differences in relative 401 abundance and rank order depending on quantification method highlight the potential 402 influence of quantification measures on a range of archaeological interpretations. Rank order 403 abundance has important implications for examining foraging practices (e.g. Szabó 2009), 404 tracking and reconstructing environmental change (Amesbury 1999) and applying models of 405 optimal foraging theory (Allen 2012, Bird and Bird 1997, 2000, Stephens and Krebs 1986:17- 406 24, Thomas 1999, 2002, 2007a, 2007b). To test the effect of differential fragmentation, the 407 fragmentation ratio for each taxon (NISP:MNI) was compared for each method. This index 408 allows the approximate calculation of the number of fragments per individual (see Faulkner 409 2010:1946). 410 411 Species richness is the number of species present in the analytical unit of study and evenness 412 is the relative abundanceACCEPTED of species (Magurran 2004:9). Species richness was calculated and 413 compared for each quantification protocol using NTAXA. NTAXA is a count of the number 414 of distinct taxa in each analytical unit calculated by collapsing taxa at the highest common 415 taxonomic level. NTAXA ensures that species richness is not artificially inflated by taxa 416 which are more easily identified to lower taxonomic levels. For example, if an analytical unit 417 consisted of individuals from the Cypraeidae family identified to Cypraea tigris, Lycina lynx 418 and Cypraeidae spp., then the NTAXA value would be one (1) rather than three (3), as

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ACCEPTED MANUSCRIPT 419 fragments identified as Cypraeidae spp. might possibly be unidentifiable fragments of 420 Cypraea tigris, Lycina lynx , or other cypraeids . 421 422 Assemblage diversity was calculated using multiple indices, the Simpson’s index of Diversity 423 (1-D) and the Shannon-Weiner index of Diversity ( H’ ) and Evenness ( E), all common to 424 zooarchaeological analyses. 1-D values range from 0 to 1, with higher values indicating even 425 assemblages of many taxa with many individuals from each taxon (Magurran 2004:116). For 426 the Shannon-Weiner index of diversity ( H’ ), theoretical values range between 0 and 5, 427 however, values between 1.5 and 3.5 are most common. Higher H’ values indicate greater 428 diversity and richness. Shannon’s Evenness ( E) values fall between 0 and 1, with values 429 closer to 0 reflecting assemblages dominated by a single taxon, and higher values reflect 430 assemblages with many taxa represented by similar numbers of individuals (Lyman 431 2008:195, Reitz and Wing 2008:111). Random permutation tests of relative abundance data 432 were carried out using the PAST Paleontological Statistics Package, Version 3.04 (Hammer 433 2001), to test for significant difference between Simpson’s Index of Diversity (1-D), 434 Shannon-Weiner Diversity ( H’ ), and Evenness ( E) values reported for both quantification 435 methods. As no alterations to taxonomic measures of dominance and evenness were reported 436 for bivalves, significant difference was tested for in gastropod samples only. 437 438 5. Results 439 5.1 Total MNI 440 Site level totals demonstrate a marked difference in MNI for each quantification method 441 (Table 1). MNI calculated using tMNI for gastropods MANUSCRIPT was three times larger (~108% - 207% 442 increase in MNI) than NRE MNI calculations. However, the tMNI for bivalves was at most 443 6% higher than NRE MNI. The most notable distinction in MNI between quantification 444 protocols was reported for gastropods from TP18 and 19 (Table 1). 445 446 Differences in MNI depending on quantification method were also reported for the top ten 447 gastropod and bivalve taxa ranked by abundance (Table 2) MNI for gastropods calculated 448 using the tMNI method doubled (~78% - 115% increase) compared to NRE MNI. Bivalve 449 MNI was similar for both quantification protocols. 450 451 452 At the site level and for the top ten ranked taxa from each test pit, tMNI resulted in a marked 453 increase in the relative proportion of gastropods to bivalves compared with NRE MNI. Using 454 NRE MNI for TP18ACCEPTED and 19, bivalves accounted for 31% of total MNI, whereas using tMNI, 455 bivalves accounted for 13%. For TP 7, NRE MNI resulted in bivalves accounting for 38% of 456 total MNI, but tMNI reported 23% bivalves and 77% gastropods. 457 5.2 Rank order abundance 458 Each quantification method resulted in different rank order abundance for the top ten 459 gastropod taxa at both sites (Tables 3 and 4). Rank order was distinct for all gastropod taxa 460 for both quantification methods for TP18 and 19. In samples from TP18 and 19 (Table 3),

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ACCEPTED MANUSCRIPT 461 Nerita plicata and Gutturnium muricinum are not reported in the top ten ranked taxa for NRE 462 MNI, but are rank one and two respectively for tMNI. For bivalves, minor alterations to MNI 463 were noted in both contexts and rank order abundance was unaltered by quantification 464 method (Tables 5 and 6). 465 5.3 Species richness and evenness 466 For gastropods, richness at the site level was higher (~17% - 90% increase) in all cases at the 467 family, genus, and species level. NTAXA was slightly lower for TP18 and 19 using the tMNI 468 method as a result of the inclusion of NRE that could not be identified beyond the family 469 level, such as the distinctive of the Muricidae (Table 7). Species richness by 470 NTAXA increased for TP7. For the top ten ranked taxa tMNI calculations decreased richness 471 (~8% - 54%) measured by count of family, genus, and species, and also by NTAXA (Table 472 7). 473 474 Increased evenness and dominance measures for all gastropod taxa are also reported at both 475 sites (Table 8). A significant difference was also reported between Simpson’s Evenness and 476 Shannon’s Evenness values for the top ten ranked gastropod taxa at TP18 and 19 ( p = 477 <0.001) . For the top ten ranked taxa only, evenness was increased for tMNI assemblages at 478 TP18 and 19, but decreased at TP7. A significant difference was also reported between 479 Simpson’s Evenness values for the top ten ranked gastropod taxa at TP18 and 19 ( p = 480 <0.001). The majority of MNI values for the top ten ranked taxa were derived from the apex, 481 leading to similar or lower values when additional elements are included using tMNI. No 482 changes to species richness or evenness due to quantification method were reported for 483 bivalves. MANUSCRIPT 484 485 5.4 Element survivorship 486 Bivalve MNI was not substantially altered by the application of the new protocol at either of 487 the tested sites. In only two cases were the adductor muscle scars the most frequent NRE. 488 However adductor muscles scars were the most frequent NRE only for those taxa with the 489 highest NISP, indicating increased fragmentation relative to other taxa. Furthermore, as many 490 taxa were represented by one or two whole individuals, often umbo, hinge and adductor 491 muscle scar counts were equal. This trend requires further investigation with a larger 492 assemblage of bivalves, assessing the influence of sample size and taphonomic alteration on 493 MNI. 494 495 The contributionACCEPTED of the spire to the total MNI for gastropods was low compared to other 496 elements (Tables 9 and 10). These results reflect the differential survivorship and ease of 497 identification of elements across taxa. The taxa where the spire was the most frequently 498 occurring element were those with easily identifiable, high, (Cerithium nodulosum ) or low 499 dense spires ( Conus spp.). Taxa with low, fragile spires, but identifiable apertures and outer 500 lips (such as the Neritidae) are consistently underrepresented by the NRE MNI method 501 utilising only spires. The MNI for these taxa increased as a direct result of the inclusion of a 502 wider range of NRE, reflected in decreased mean fragmentation ratio values for all gastropod

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ACCEPTED MANUSCRIPT 503 taxa (TP18 & 19: NRE MNI: 10.6/tMNI: 3.4, TP7 NRE MNI: 4.2/tMNI: 2.5) and the top ten 504 ranked gastropod taxa (TP18 & 19: NRE MNI: 9.7/tMNI: 2.3, TP7 NRE MNI: 4.2/tMNI: 505 2.9) when tMNI was implemented. The inclusion of taxa specific NRE, such as the base and 506 labum of the Cypraeidae, and the additional NRE for the Neritidae increased MNI markedly, 507 highlighting the utility of including additional NRE based on the judgment of the analyst. 508 509 6. Discussion

510 The inclusion of a wider range of molluscan NRE has resulted in notable changes in 511 abundance, diversity and evenness. Studies of the quantification of fish from Pacific 512 Island archaeological sites have produced similar alterations to relative abundance and rank 513 order when the range of elements is increased (Lambrides and Weisler 2013). These results 514 highlight the need for analysts to carefully consider the potential loss of information which 515 results from counting a restricted range of NREs. The application of tMNI to an assemblage 516 of mollusc shells from the Marshall Islands has demonstrated that the morphological and 517 architectural complexity of molluscan shell must be considered in MNI quantification 518 protocols. The tMNI and NRE MNI methods are both reliable; if each method was tested 519 multiple times for the same assemblage, results will be consistent (Nance 1987, Giovas 520 2009:1653). However, tMNI increases measurement validity and provides abundance 521 estimates that more closely estimate actual abundance of taxa within an assemblage (Carr 522 1987, Mason 1998:308). 523 524 Total MNI for the NRE MNI method and the tMNI method are markedly different and the 525 relative abundance of gastropods was increased subsMANUSCRIPTtantially. Individuals were more likely to 526 be quantified when a wider range of NRE were utilised, indicated by decreased values for 527 fragmentation ratio for all gastropods and the top ten ranked taxa. The rank order abundance 528 and element survivorship for NRE MNI at MLEb-1 demonstrate a dominance of taxa with 529 high, dense spires relative to overall size, including Cerithium columna , C. nodulosum, and 530 Planaxis sulcatus, and gastropods with dense, low spires, such as Conus spp. and Vasum 531 turbinellus . A more balanced range of shell forms are represented when MNI is calculated 532 using a wider range of NRE. The MNI and rank-order abundance of globoid, weak spired 533 taxa such as Nerita spp. , and small taxa with apertural thickening, such as 534 nicobaricus and Monetaria moneta, were increased. Overall, the spire was rarely the most 535 frequently occurring element when a range of NREs were quantified. Giovas (2009:1563) 536 stated that the NRE method can produce coarse grained estimates of abundance, but the 537 results outlined here indicate that the NRE MNI method provides MNI values that are 538 inherently biasedACCEPTED towards particular forms. 539 540 Quantification results for all bivalves demonstrated that the umbo and hinge are the elements 541 most frequently preserved in this assemblage. It is possible that the NRE MNI method is 542 adequate for quantifying bivalves, but a larger sample of bivalves is needed for conclusive 543 results. The counting of adductor muscle scars is not a time consuming process. Moreover, 544 the application of tMNI allows the analyst to determine the relative contribution of each NRE 545 to total MNI.

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ACCEPTED MANUSCRIPT 546 547 The application of the tMNI method resulted in variable alterations to richness, dominance 548 and evenness. By including a wider range of elements, richness as measured by count of 549 family, genus, species was increased at both sites for all taxa. NTAXA was increased at TP7 550 for all taxa, as taxa lacking spires which preserve well were represented in MNI calculations. 551 NTAXA decreased at TP18 and 19 for all taxa, as a result of family-level identifications. For 552 example, the Muricidae were represented by individuals from six genera and eight species, 553 but family level identification of the distinctive siphonal notches necessitated a collapsed 554 value of 1 for NTAXA. Quantification of additional NRE is resulted in increased family, 555 genus, and species counts. Family, genus, species and NTAXA values decreased for the top 556 ten ranked taxa at both sites. Using the NRE MNI method, many taxa are represented by 557 single individuals, and many taxa reported the same MNI value. The inclusion of additional 558 NRE added more individuals of each taxon as the bias toward taxa with robust spires was 559 ameliorated. As a result, fewer taxa reported equal MNI values. 560 561 For TP18 and 19, significant differences were noted for Simpson’s Index values for all 562 gastropod taxa, and the top ten ranked taxa only. Simpson’s Index is sensitive to changes in 563 the most abundant species in the sample, while less sensitive to species richness (Maguran 564 2004:115) The significant difference between quantification methods is likely due to the 565 dominance of Cerithidae (MNI = 64, 43.2% of total MNI) in NRE MNI calculations (Table 566 3), However, when MNI is calculated using the tMNI method, no single taxon accounts for 567 more than 20.6% of the total MNI. Shannon’s index is focussed on species richness and 568 relative abundance (Magurran 2004:107), and MANUSCRIPT the minimal difference in NTAXA and 569 differential distribution of individuals across taxa likely accounts for the non-significant 570 difference between quantification methods for all taxa, and the significant difference between 571 top-ten ranked gastropod taxa from TP18 and 19. For TP7, evenness is similar between 572 quantification methods (Table 4), reflected in the non-significant difference between 573 Simpson’s index values derived from both the NRE MNI and tMNI methods. Changes in diet 574 breadth and evenness are critical measures for testing hypotheses relating to declines in 575 foraging efficiency and the long-term impacts of human extraction of molluscan resources 576 (e.g. Broughton 1997, Nagaoka 2002). Measures of taxonomic abundance and heterogeneity 577 are also crucial to understanding forager decision making (e.g. Thomas 2007a), assisting in 578 reconstructing environmental change (Amesbury 1999, Faulkner 2013), and informing 579 modern conservation efforts (e.g. Carder and Crock 2012, Wake et al. 2013). While not all 580 differences were statistically significant, alterations to measures of taxonomic composition 581 highlight the potential influence of quantification protocol on archaeological inference. 582 ACCEPTED 583 Researchers are not able to directly compare the abundances of fish and molluscs using 584 current protocols. The NRE MNI method tend to measure the survival of the pre-selected 585 counting character, rather than the relative abundance of all taxa in the sample. The tMNI 586 method allows invertebrate abundance data to be compared to vertebrate abundance data (e.g. 587 fishbone or terrestrial ) as the most frequently occurring NRE is used for both 588 measures. In many coastal sites, especially in the Pacific Islands, molluscan and finfish 589 resources are important sources of , often captured from the same habitats. Therefore,

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ACCEPTED MANUSCRIPT 590 NRE MNI protocols may be masking changes in subsistence systems that are reflected by 591 shifting abundances of molluscs and finfish. The preservation or lack thereof of particular 592 NREs may also be a useful indicator of a range of human behaviours, including off-site 593 processing or shell-working. 594 595 7. Conclusion

596 The new protocol for the quantification of mollusc shell demonstrates the impact of 597 increasing the range of NRE included in MNI calculations. This new protocol has produced 598 MNI values that more closely match the actual abundance of taxa from an archaeological 599 assemblage in the Marshall Islands. Using this quantification protocol, alterations to richness, 600 diversity, and rank order abundance were reported. Existing quantification protocols for 601 molluscan shell consistently underrepresent the relative abundance of gastropods to bivalves 602 and underrepresent the abundance of particular shell forms. To enhance the study of 603 prehistoric subsistence and provide data that are increasingly used by conservation biologists, 604 historical ecologists, and policy makers, it is critical that the most accurate representation of 605 richness, evenness, and diversity are provided. We recommend that the implications of 606 utilising tMNI quantification protocols be routinely considered. We encourage analysts to 607 locally adapt and refine the protocols outlined here for the analysis of molluscan 608 assemblages. 609 610 8. Acknowledgements

611 We thank the Vice Chancellor of the University MANUSCRIPTof Queensland for strategic funding to the 612 archaeology programme. Marshall Islands fieldwork, conducted under permit from the 613 Historic Preservation Office (Republic of the Marshall Islands) was supported by a grant to 614 Weisler from the Office of the Deputy Vice Chancellor (Research). Harris’ postgraduate 615 studies are supported by an Australian Postgraduate Award. The authors thank the three 616 anonymous reviewers for their useful comments on the manuscript. 617 618 619

620

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ACCEPTED MANUSCRIPT 621 9. References

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ACCEPTED MANUSCRIPT 796 Rhoads, D.C., Lutz, R.A., 1980. Skeletal growth of aquatic organisms: biological records of 797 environmental change, Plenum Press, New York. 798 799 Robb, K.F., Nunn, P.D., 2013. Changing role of nearshore-marine foods in the subsistence 800 economy of inland upland communities during the last millennium in the tropical Pacific 801 Islands: insights from the B ā River Valley, Northern Viti Levu Island, Fiji. Environmental 802 Archaeology 19, 1-11. 803 804 Röckel, D., Korn, W., Kohn, A.J., 1995. Manual of the living Conidae. Verlag Christa 805 Hemmen, Hackenheim. 806 807 Rosendahl, P.H., 1987. Report 1: Archaeology in Eastern Micronesia: Reconnaissance 808 Survey in the Marshall Islands, in: Dye, T. (Ed.), Marshall Islands Archaeology. Pacific 809 Anthropological Records, Vol. 38 Department of Anthropology, Bernice Pauahi Bishop 810 Museum, Honolulu, pp. 17-166. 811 812 Ruppert, E.E., Fox, R.S., Barnes, R.D., 2004. : a functional evolutionary 813 approach, 7th ed. Thomson-Brooks/Cole, Belmont. 814 815 Seeto, J., Nunn, P.D., Sanjana, S., 2012. Human-mediated prehistoric marine extinction in the 816 tropical pacific? Understanding the presence of Hippopus hippopus (linn. 1758) in ancient 817 shell middens on the Rove Peninsula, southwest Viti Levu Island, Fiji. Geoarchaeology 27, 2- 818 17. MANUSCRIPT 819 820 Stachowitsch, M., Proidl, S., 1992. The Invertebrates: an Illustrated Glossary. Wiley-Liss, 821 New York. 822 823 Stephens, D.W., Krebs, J.R., 1986. Foraging theory. Princeton University Press, Princeton. 824 825 Sommerville-Ryan, G., 1998. The Taphonomy of a Marshall Islands' Shell Midden. M.A. 826 Thesis, Department of Anthropology and Archaeology, University of Otago, Dunedin. 827 828 Szabó, K., 2009. Molluscan remains from Fiji, in: Clark, G., Anderson, A.J. (Eds.), The Early 829 Prehistory of Fiji, Terra Australis, Vol. 31, ANU E Press, Canberra, pp. 183-212. 830 831 -, 2012. Terrestrial hermit crabs (Anomura: Coenobitidae) as taphonomic agents in circum- 832 tropical coastal ACCEPTEDsites. Journal of Archaeological Science 39, 931-941. 833 834 Thomas, F.R., 1999. Optimal Foraging and Conservation: The Anthropology of Mollusk 835 Gathering Strategies in the Gilbert Islands Group, Kiribati. Ph.D. Thesis, University of 836 Hawaii at Manoa, Honolulu. 837

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ACCEPTED MANUSCRIPT 838 -, 2002. An evaluation of central-place foraging among mollusk gatherers in western Kiribati, 839 Micronesia: linking behavioral ecology with ethnoarchaeology. World Archaeology 34, 182- 840 208. 841 842 -, 2007a. The behavioral ecology of shellfish gathering in western Kiribati, Micronesia 1: 843 prey choice. Human Ecology 35, 179-194. 844 845 -, 2007b. The behavioral ecology of shellfish gathering in western Kiribati, Micronesia. 2: 846 patch choice, patch sampling, and risk. Human Ecology 35, 515-526. 847 848 Vermeij, G.J., 1979. Shell architecture and causes of death of Micronesian reef . 849 Evolution 33, 686-696. 850 851 Vermeij, G.J., 1993. A natural history of shells. Princeton University Press, Princeton. 852 853 Wake, T.A., Doughty, D.R., Kay, M., 2013. Archaeological investigations provide late 854 Holocene baseline ecological data for Bocas del Toro, Panama. Bulletin of Marine Science 855 89, 1015-1035. 856 857 Walter, R., 1998. Anai'o: the archaeology of a fourteenth century Polynesian community in 858 the Cook Islands. New Zealand Archaeological Association Monograph, Vol. 22, New 859 Zealand Archaeological Association, New Zealand. 860 MANUSCRIPT 861 Weisler, M.I., 1999. The antiquity of aroid pit agriculture and significance of buried A 862 horizons on Pacific atolls. Geoarchaeology: An International Journal 14, 621-654. 863 864 -, 2001. On the Margins of Sustainability: Prehistoric Settlement of Utr ōk Atoll, Northern 865 Marshall Islands. BAR International Series 967, Archaeopress, Oxford. 866 867 -, 2002. Archaeological Survey and Test Excavations on Ebon Atoll, Republic of the 868 Marshall Islands. Historic Preservation Office, Republic of the Marshall Islands, Majuro. 869 870 White, T.E., 1953. A method of calculating the dietary percentage of various food animals 871 utilized by aboriginal peoples. American Antiquity 18, 396-398. 872 873 Wolverton, S., 2013. Data quality in zooarchaeological faunal identification. Journal of 874 Archaeological ACCEPTEDMethod and Theory 20, 381-396. 875 876 Zuschin, M., Stachowitsch, M., Stanton, R.J., Jr., 2003. Patterns and processes of shell 877 fragmentation in modern and ancient marine environments. Earth-Science Reviews 63, 33- 878 82. 879

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ACCEPTED MANUSCRIPT Table 1 Total MNI for all taxa, aggregated at test pit level NRE Site tMNI MNI Gastropoda Eb-1 TP18 &19 188 577 Eb-33 TP7 125 260 Bivalvia Eb-1 TP18 &19 83 88 Eb-33 TP7 78 78 Total 474 1003

Table 2 Total MNI for the top ten ranked taxa only, aggregated at test pit level NRE Site tMNI MNI Gastropoda Eb-1 TP18 &19 170 367 Eb-33 TP7 125 223 Bivalvia Eb-1 TP18 &19 83 88 Eb-33 TP7 78 78 Total 456 756

Table 3 Eb-1 T18 and 19 NRE MNI and tMNI quantification results for gastropod taxa. * = change in rank order; ** = unique to rank order abundance for thatMANUSCRIPT method. NRE MNI tMNI Rank Taxon MNI NISP Rank Taxon MNI NISP

1 Conus spp.* 36 190 1 Nerita plicata** 83 141

2 Cerithium columna * 32 45 2 Gutturnium muricinum** 70 132

3 Cerithidae spp.* 20 37 3 Conus spp.* 36 190

4 Vasum turbinellus ** 11 73 4 Cerithium columna* 32 45

5 Cerithium nodulosum ** 10 176 5 Monoplex intermedius** 31 41

6 Planaxis sulcatus ** 10 16 6 ** 29 37

7 Drupa spp. ** 8 37 7 Nerita polita** 27 75

7 Canarium spp. ** 5 13 8 Monetaria moneta** 22 33

8 Turbo argyrostomus ** 5 51 9 Cerithidae spp.* 20 37

8 Turbo spp. ** 5 85 10 Chicoreous spp.* 17 64

8 Mitra stictica ** 4 18 9 Chicoreous spp.* 3 64 9 Trochus maculatus ** 3 58 9 MonoplexACCEPTED spp. ** 3 46 9 Conus flavidus ** 3 3 9 Nerita signata ** 2 9 9 Melampus flavus ** 2 6 9 Cerithium echinatum ** 2 3 9 Pollia sp. ** 2 3 9 Trochidae spp. ** 2 8 10 Turbo setosus ** 2 49

ACCEPTED MANUSCRIPT Table 4 Eb-33 TP7 NRE MNI and tMNI quantification results for gastropod taxa. * = change in rank order; ** = unique to rank order abundance for that method. NRE MNI tMNI Rank Taxon MNI NISP Rank Taxon MNI NISP

1 Melampus flavus * 35 44 1 Nerita polita * 73 122

2 Conus spp. * 17 71 2 Melampus flavus * 39 44

3 Vasum turbinellus 16 90 3 Vasum turbinellus 28 90

4 Nerita polita* 15 122 4 Conus spp.* 23 71

5 Turbo argyrostomus* 7 60 5 Bursa spp.** 13 36

6 Cerithidae spp.* 6 8 6 Nerita plicata* 9 11

7 Nerita plicata 5 11 7 Turbo argyrostomus* 7 60

7 Cerithium nodulosum 5 48 8 Cerithidae spp.* 6 8

8 Canarium spp. 2 9 8 Monoplex nicobaricus ** 6 6

8 Thais armigera ** 2 8 9 Cerithium nodulosum* 5 48

8 Nerita spp. ** 2 7 9 Canarium spp.* 5 9 9 Drupa ricinus ** 1 4 9 Gutturnium muricinum ** 5 6 9 Mitra stictica ** 1 8 10 Drupa ricinus* 4 4 9 Monetaria moneta ** 1 2 9 Muricidae spp.** 1 6 9 Cerithium columna ** 1 1 9 Sabia conica ** 1 2 9 Pterigya sp. ** 1 1 9 Drupa rubusidaeus ** 1 2 9 Mamilla sp. ** 1 1 9 Nerita albicilla ** 1 1 9 Terebra spp. ** 1 2 9 Trochidae spp. ** 1 2 9 Turbo spp. ** 1 17 MANUSCRIPT 10 - - -

Table 5 Eb-33 TP7 NRE MNI and tMNI quantification results for bivalve taxa. NRE MNI tMNI Rank Taxon MNI NISP Rank Taxon MNI NISP

1 Fragum spp. 48 51 1 Fragum spp. 48 51

2 Asaphis violascens 10 27 2 Asaphis violascens 10 27

3 Chama spp. 10 11 3 Chama spp. 10 11

4 Corculum cardissa 4 4 4 Corculum cardissa 4 4

5 Spondylus sinensis 2 2 5 Spondylus sinensis 2 2

5 Ctena bella 2 2 5 Ctena bella 2 2

6 Arca sp. 1 1 6 Arca sp. 1 1

6 Spondylus sp. 1 1 6 Spondylus sp. 1 1

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ACCEPTED MANUSCRIPT Table 6 Eb-1 TP18 and 19 NRE MNI and tMNI quantification results for bivalve taxa. NRE MNI tMNI Rank Taxon MNI NISP Rank Taxon MNI NISP

1 Asaphis violascens 25 145 1 Asaphis violascens 26 145

2 Fragum spp. 10 11 2 Fragum spp. 10 11

2 Gafraium spp. 10 23 2 Gafrarium spp. 10 23

3 Ctena bella 6 6 3 Ctena bella 6 6

4 Tridacna spp. 3 4 4 Tridacna spp. 3 4

5 Arca spp. 2 3 5 Arca spp. 2 3

5 Barbatia spp. 2 5 5 Barbatia spp. 2 5

5 Tridacna maxima 2 5 5 Tridacna maxima 2 5

5 Chama spp. 2 7 5 Chama spp. 2 7

5 Codakia divergens 2 2 5 Codakia divergens 2 2 5 Lucinidae spp. 2 2 5 Lucinidae spp. 2 2 5 Spondylus spp. 2 4 5 Spondylus spp. 2 4 6 Tridacna crocea 1 2 6 Tridacna crocea 1 2 6 Tridacna cf. gigas 1 2 6 Tridacna cf. gigas 1 2 6 Vasticardium elongatum 1 1 6 Vasticardium elongatum 1 1 6 Codakia tigerina 1 1 6 Codakia tigerina 1 1 6 Pectinidae spp. 1 2 6 Pectinidae spp. 1 2 6 Pinctada spp. 1 20 6 Pinctada spp. 1 20

Table 7 Unique family, genus, and species counts for gastropods aggregated by test pit. For each test pit, the counts for all taxa and the top ten ranked taxa are presented. All taxa Top ten taxa MANUSCRIPT Eb-1 TP18 & 19 NRE MNI tMNI NRE MNI tMNI Families 18 21 13 6 Genera 24 32 14 8 Species 21 40 12 7 NTAXA 26 25 14 9

Eb33 TP7 NRE MNI tMNI NRE MNI tMNI Families 14 17 14 11 Genera 15 23 15 9 Species 14 21 14 11 NTAXA 15 19 15 12

ACCEPTED ACCEPTED MANUSCRIPT Table 8 Gastropod Evenness and dominance measures.1-D = Simpson’s Evenness, H’ = Shannon’s index, E = Shannon’s evenness All Taxa TP18 & 19 TP7

NRE MNI tMNI p NRE MNI tMNI p

1-D 0.787 0.884 <0.001 0.822 0.830 0.995 H’ 2.273 2.412 0.301 1.988 2.182 0.297 E 0.698 0.749 0.852 0.734 0.741 0.827

Top ten ranked TP18 & 19 TP7

NRE MNI tMNI p NRE MNI tMNI p

1-D 0.788 0.858 <0.001 0.841 0.825 0.476 H’ 1.975 2.069 0.208 2.084 2.057 0.903 E 0.748 0.942 <0.001 0.770 0.828 0.084

Table 9 Gastropod NRE frequency distribution, Eb-1, TP18 and 19, top ten ranked taxa by tMNI; AC = anterior canal; PC = posterior canal; OL = outer lip; Ap. = aperture; Op. = operculum; I = Cypraeidae: base / Neritidae: posterior columellar deck/outer lip intersection II = Cypraeidae: labum / Neritidae: anterior columellar deck/outer lip intersection. MNI NREMNI tMNI NISP Spire AC PC OL Ap. Op. Um. I II Nerita plicata 1 83 141 0 - - 54 31 0 - 76 83 Gutturnium muricinum 1 70 132 MANUSCRIPT 1 50 - 70 31 0 - - - Conus spp. 36 36 190 36 30 - 0 0 0 - - - Cerithium columna 32 32 45 32 6 - 1 0 0 - - - Monoplex intermedius 0 31 41 0 15 - 31 2 0 - - - Monoplex nicobaricus 0 29 37 0 10 - 29 2 0 - - - Nerita polita 0 27 75 0 0 - 7 1 0 - 27 19 Monetaria moneta 0 22 33 0 0 - 2 0 0 - 4 22 Cerithidae spp. 20 20 37 20 0 - 0 0 0 - 0 0 Chicoreous spp. 3 17 64 3 17 - 2 1 0 - - -

ACCEPTED ACCEPTED MANUSCRIPT Table 10 Gastropod NRE frequency distribution, Eb-33, TP7, top ten ranked taxa by tMNI; AC = anterior canal; PC = posterior canal; OL = outer lip; Ap. = aperture; Op. = operculum; I = Cypraeidae: base / Neritidae: posterior columellar deck/outer lip intersection II = Cypraeidae: labum / Neritidae: anterior columellar deck/outer lip intersection. MNI NREMNI tMNI NISP Spire AC PC OLR Ap. Op. Um. 1 2 Nerita polita 15 73 122 15 0 - 35 24 0 - 73 24 Melampus flavus 35 39 44 35 38 - 39 38 0 - - - Vasum turbinellus 16 28 90 16 28 - 0 1 0 - - - Conus spp. 17 23 71 17 23 - 1 1 0 - - - Bursa spp. 0 13 36 0 11 9 13 6 0 - - - Nerita plicata 5 9 11 5 0 - 8 7 0 - 9 8 Turbo argyrostomus 7 7 60 7 1 - 0 0 5 - - - Cerithidae spp. 6 6 8 6 5 - 5 5 0 - - - Monoplex nicobaricus 0 6 6 0 6 - 6 5 0 - - - Cerithium nodulosum 5 5 48 5 3 - 3 0 0 - - -

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ACCEPTED ACCEPTED MANUSCRIPT Figure 1 a. Stratigraphic section of MLEb-1, TP19 (left, 1 m wide) & TP18 (right, 1 m wide) south profile with a maximum depth of ~115cmbs. Scale is 1 m long.; b. North profile of TP7, with a maximum depth of 40cmbs at site MLEb-33 on Enekoion islet. The dense cultural deposit is ~35 cm thick. Scale is 1m long. (Both photos, M. Weisler)

Figure 2 Gastropod terminology.

Figure 3 Examples of gastropod shapes. a. globoid b. involute c. tubular d. trochoid e. turbinate f. patelliform h.disjunct i. turriform. Note the substantial variation in spire height between shell forms.

Figure 4 Bivalve terminology

Figure 5 Examples of bivalve shapes a. orbicular b. alate c. auriculate d. subquadrate e. trigonal f.fan-shaped g.ensiform h. elongate-elliptical.

Figure 6 Gastropod NRE (1 = spire; 2 = anterior canal; 3 = posterior canal; 4 = outer lip; 5 = aperture; 6 = operculum; 7 = umbilicus). Hatched areas represent areas of shell included in quantification of MNI. Note the presence of NRE on some shell forms, but not others.

Figure 7 The tMNI method of gastropod MNI calculation. Hatched areas represent a fragment of shell. (Sp = spire; OL = outer lip; Ap = aperture; AC = anterior canal/notch; PC = posterior canal /notch; Um = umbilicus; Op = Operculum).

Figure 8 Bivalve NRE (1 = umbo; 2 = posterior hinge; 3 = anterior hinge; 4 = posterior adductor muscle scar; 5 = anterior adductor muscle scar). NoteMANUSCRIPT the presence of only a single adductor muscle scar on the monomyarian shell valve of b.

Figure 9 The tMNI method of bivalve MNI calculation. Hatched areas represent a shell fragment. (Um = umbo; AH = anterior hinge; PH = posterior hinge; AAMS = anterior adductor muscle scar; PAMS = posterior adductor muscle scar)

Figure 10 Additional NRE included in tMNI quantification. Hatched areas represent areas of shell included in quantification of MNI. a. view of Cypraeidae spp. aperture and base showing additional NRE; b. view of Neritidae spp. aperture and columellar deck showing location of additional NRE, I = anterior columellar deck/outer lip intersection Neritidae NRE, II = posterior columellar deck/outer lip intersection.

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