ORIGINAL PAPER

Intensified mollusk exploitation on Nevis (West Indies)reveals ~six centuries of sustainable exploitation

Aaron S. Poteate & Scott M. Fitzpatrick & Meagan Clark &

Jessica H. Stone

Received: 26 September 2013 /Accepted: 19 May 2014# Springer-Verlag Berlin Heidelberg 2014

Abstract In this study, we conducted one of the most detailedand comprehensive analyses to date of a pre-Columbian mol-lusk assemblage in the Caribbean. The robust sample, fromthe island of Nevis in the northern Lesser Antilles, comprisedmore than 58,000 individuals recovered from a 25 m2 (40 cmdeep) midden deposit at the Late Ceramic Age (ca. AD 890–1440) site of Coconut Walk. Using this sample, we examinedmollusk exploitation over a ca. 600-year time period, whichrevealed heavy dependence on only a few species. Statisticalanalysis demonstrates that even though mollusk harvestingintensified through time, there was an increase of more than10 % in the average individual weight of the three mainspecies and a ten-fold increase in harvesting generally. Thesedata, in conjunction with a previously observed size increase ofone of the three taxa (Nerita tessellata)—which was increas-ingly preyed on through time—infer a level of sustainabilitycontra to prey-choice models in which over-exploitation is anexpected outcome. Overall, the foraging of mollusks at this siteappears to have been sustainable for ~six centuries based on theabsence of evidence for over-harvesting and increase in sizeduring the time of occupation, regardless of its causation(anthropogenic, climatic, environmental, or otherwise).

Keywords Shellfish . Subsistence . Exploitation .

Caribbean . Pre-Columbian

Introduction

In archaeological midden contexts, prehistoric subsistencestrategies and environmental interaction can be examined

through the analysis of faunal remains, of which mollusksare often the most common constituent (see Stiner andMunro 2002; Giovas et al. 2010; Grayson 1984; Manninoand Thomas 2002; Morrison and Hunt 2007; Botkin 1980; deBoer et al. 2000; Jerardino 2010; Klein et al. 2004; Faulkner2009; Braje 2010; Rick et al. 2008). In recent years, a growingnumber of archaeological studies have demonstrated that theprehistoric exploitation of mollusks through time can result inconsequences (positive, negative, or both) for exploited taxaas a result of increased or expanded predation across time andspace (e.g., Claassen 1998; Stiner and Munro 2002; Manninoand Thomas 2002; Erlandson et al. 2011; Rick 2011; FitzpatrickandKeegan 2007; Fitzpatrick et al. 2008). As such, this providesan excellent opportunity to examine the extent to which variousmollusk taxa were exploited through time and the human andnatural processes that may have influenced predation.

In many of these cases, for example, a decrease in the sizeof a particular taxon through time is used to infer over-harvesting (Allen 2012; Braje et al. 2012; Claassen 1998;Erlandson et al. 2008, 2011; Erlandson and Rick 2010;Faulkner 2009; Mannino and Thomas 2001, 2002; Milneret al. 2007; Morrison and Hunt 2007; Morrison andCochrane 2008; Prummel 2005; Reitz and Wing 2008;Stager and Chen 1996), and may be coupled with trendsobserved in other taxa within the same assemblage to suggestthese processes were in fact occurring. Another trend ob-served archaeologically includes a change in the -age profilethat often follows a decrease in the size of mollusks (Cannonand Burchell 2009; Giovas et al. 2010; Mannino and Thomas2001, 2002; Stager and Chen 1996), a noticeable shift inabundance (Broughton 2002; Cannon 2000, 2003; Graysonand Cannon 1999; Morrison and Hunt 2007; Nagaoka 2005),and a decline in taxonomic richness (Grayson et al. 2001;Nagaoka 2001; Jones 2004).

To provide a framework for examining these and otherissues related to harvesting of faunal resources by humans,

A. S. Poteate (*) : S. M. Fitzpatrick :M. Clark : J. H. StoneDepartment of Anthropology, University of Oregon, Eugene,OR 97403, USAe-mail: apoteate@uoregon.edu

Archaeol Anthropol SciDOI 10.1007/s12520-014-0196-1

Claassen (1986, 1998) introduced a set of criteria to defineover-exploitation in archaeological samples of mollusks,which was later expanded upon by Mannino and Thomas(2002). They noted that the minimum criteria for exploited(preferred) species include (1) a decrease in size, (2) change inage profile, and (3) decline in abundance that form a contin-uum of possibilities instead of an independent set of criteria(Mannino and Thomas 2002). More recent research suggeststhat this theoretical model has exceptions, with several casesthat are counter to these observations (see Giovas et al. 2010,2013; Thakar 2011; McCoy 2008). To understand the com-plex interactions that lead to the consumption and humanover-harvesting of mollusks, we investigate how findings atthe Late Ceramic Age (ca. AD 890–1440) site of CoconutWalk on the island of Nevis in the northern Lesser Antilles(eastern Caribbean) fit into these earlier models (Claassen1986, 1998; Mannino and Thomas 2002). This is especiallyrelevant given that human impacts on mollusks have beenpreviously observed at other pre-Columbian sites in the region(Keegan et al. 2003; Giovas 2013; Giovas et al. 2013).

In this paper, we discuss a robust spatial (5×5 m) andtemporal sample of invertebrates (MNI=58,000+) recoveredfrom CoconutWalk. This site provides an excellent case studywith which to analyze the criteria for over-exploitation andexplore human-environmental relationships between severalspecies over a ca. 600-year period. To define the level ofexploitation at Coconut Walk, the criteria for over-exploitation were examined by determining the richness ofspecies over time and if the three most common species atCoconut Walk (Nerita tessellata, Cittarium pica, andLithopoma tuber) met the criteria for over-exploitation, thatis, a decrease in size, change in age profile, and decline inabundance.

Research background

The island of Nevis, located in the northern part of theLeeward Islands of the Lesser Antilles, is part of the politicalconfederation of St. Kitts and Nevis (Fig. 1). Nevis was

Fig. 1 Map of the Caribbean

Archaeol Anthropol Sci

known as Oualie (“land of beautiful water”) to its originalAmerindian inhabitants, and is the smaller of the two islandswith an area of 93 km2. At its center, Nevis reaches a height of985 m on the dormant volcano Nevis Peak. Similar to otherislands in the Caribbean, the temperature is tropical with onlyslight seasonal variation.

The first systematic survey of pre-Columbian sites onNevis was conducted by Samuel Wilson (1989) who recordeda number of settlement locations, ranging from one dating tothe Archaic Age (ca. 2500 BC) to numerous others that datedto the Ceramic Age (ca. 500 BC–AD 1400). Along the eastcoast of Nevis, the Coconut Walk site was one of the largestreported by Wilson, in which he noted an abundance ofLate Ceramic Age (post-AD 600), pottery, food remains,and several human burials on the surface (1989: 427–450;2007:62–63). Coconut Walk was initially excavated in1998 as a part of the British-based television show TimeTeam. Their findings revealed cultural material similar towhat was seen on the surface by Wilson (1989, 2006),along with subsurface evidence for household structures(postholes) and a wider array of archaeological remains(Bellamy 2001; Nokkert 2001).

Expanding on the work of the Time Team at Coconut Walk,an archaeological field school was organized in 2010 and co-directed by Scott M. Fitzpatrick, Michiel Kappers, and QuettaKaye. The primary purpose of the fieldwork was to provide amore detailed record of pre-Columbian activity (Kaye et al.2010) and establish a chronological framework for settlementgiven that no 14C dates had yet been reported for the site. Overthe course of the 6-week field project, three 5×5 m trencheswere excavated (3073, 2973, and 2273). The first two wereplaced in the central portion of the site where posthole featureshad been previously found by the Time Team and that likelyrepresented the primary area of household occupation. This

area contained shallow (30 cm) anthropogenic deposits withsterile, sandy subsoil underneath. The third trench (2273) wasfocused on a midden deposit (Fig. 2). Typical of many coastalmiddens in the Caribbean, Trench 2273 did not show visiblelayer distinctions and was excavated in four 10 cm levels(plana) after 5 cm of mixed surface material and topsoil hadbeen removed. Excavation proceeded by dividing the trenchinto smaller 1×1 m units which were excavated by a trowel.Four units designated “environmental squares” were wetsieved through 6 mm mesh with a column sample of nested6 and 1.6 mm sieves taken from the southwestern quarter(50×50 cm) of each of these four units (squares 7, 9, 17,and 19). The midden was later shown to have been depositedbetween ca. AD 890 to 1440 based on four radiocarbon datesfrom marine shell (Table 1), and contained an abundance offaunal remains, artifacts (e.g., pottery, shell and stone beadsand ornaments, chert debitage), and remnants of a partialjuvenile human burial. Of particular interest was the largemollusk assemblage which was analyzed in full. Given thatthe site was excavated with exceptional spatial and verticalcontrol across a wide area, this has afforded a unique oppor-tunity to examine a host of issues related to sampling proce-dures (Poteate and Fitzpatrick 2013), levels of faunal exploi-tation (e.g., Giovas et al. 2013), and general subsistencepractices.

Mollusk assemblage

The complete suite of faunal material recovered in excavationat Coconut Walk was shipped to North Carolina StateUniversity where further sorting, identification, cataloging,and analysis were conducted in the archaeology laboratory.After initial sorting, the shell weighed in at a total of 144 kg(317.5 lbs). Shells were sorted to the lowest level of

Fig. 2 Trench 2273 at CoconutWalk. Raised units (7, 9, 17, and19) are environmental squaresdesignated for wet sieving

Archaeol Anthropol Sci

taxonomic categorization possible (in most cases, down tospecies) following standard identification protocol. Each in-dependent taxon was quantified using the number of identifiedspecimens (NISP), minimum number of individuals (MNI)based on non-repetitive elements (NRE), and weight (g) usingtraditional procedures (see Reitz and Wing 2008; Gassiot2005; Mannino and Thomas 2001; and Morrison andCochrane 2008). Results are shown in Table 2 by taxon. Wealso conducted a quality control check on 12 % of the bags(n=19), representing 14.5 % of the total weight (20.52 kg)after sorting was completed to ensure the accuracy of identi-fication and quantification. The only major issue identifiedwas an error in chiton (Chiton tuberculatus) MNI and NISP,but the total difference was less than 1 % between initialsorting and the subsequent quality control check. There wasalso a marginal difference (1.3 %) observed in the weight ofspecies. This was likely the result of soil in the interior whorls,of gastropods which is often difficult to remove during initialcleaning, that continued to exit the specimens through repeat-ed handling and sorting.

A minimum of 78 individual taxa were identified, whichwas represented by three classes (gastropoda, bivalvia, andpolyplacophora) (Table 2). The assemblage was dominated,however, by three gastropod species: N. tessellata, C. pica,and L. tuber. These would have all lived in the same intertidalarea and likely been harvested and processed in similar ways,perhaps together. Below, we describe the basic attributes ofeach species and their relative importance (based on weight ofthe recovered shell remains) with regard to molluskharvesting.

N. tessellata, commonly known as the checkered nerite forits tessellated (checkered mosaic) pattern (Fig. 3), is common-ly found throughout the Caribbean in pre-Columbian sites(Giovas 2013; Newsom and Wing 2004) and was the mostabundant species by MNI at Coconut Walk. This species iscurrently found in large numbers in intertidal regions alongrocky shorelines in up to 0.5 m of water (Axelsen 1968;Bovbjerg 1984; Chislett 1969; Potts 1980) in the Gulf ofMexico from Florida to Texas as well as through theCaribbean to Brazil (Abbott and Morris 1995; Rehder 1981).This taxon is typically under 20 mm in length (Axelsen 1968;Lewis 1971). Because Nerita tessellata makes up a largemajority of the mollusk collection from Coconut Walk(64.1 % of MNI, 45.6 % of NISP, and 21.5 % of weight), itis justifiably a preferred species (see Table 2).

The taxon was typically found with at least 90 % of theoriginal shell in place and often had a 1:1 MNI to NISP ratio(37,591 to 38,804 for the trench as a whole). N. tessellata isoften found with Nerita peloronta and Nerita versicolor(Axelsen 1968), two species that were also present atCoconut Walk, but in comparatively smaller numbers(MNI=32 and 5,060, respectively). Prehistorically,N. tessellata would have been gathered from rocks and likelyT

able1

Radiocarbon

datesfrom

theCoconut

Walksite

Lab

no.

Sampleno.

Databaseno.

Type

Species

Unit

Square

Planum

Feature

Centim

eter

belowsurface

Weight(g)

13C/12C

ratio

Measure

14C

age

Cal.(2sigm

a)

Beta-290340

Nev-01

10NCW003S

HE

Shell

Eustrom

busgigas(juvenile)

2273

61

TOP

0–10

67.0

1.8

1350

±40

AD970–1170

Beta-324951

Nev-03

10NCW0201SH

ESh

ell

Cittariumpica

2273

253

L001

20–30

55.4

0.3

570±30

AD1320–1440

Beta-290341

Nev-02

10NCW0166SH

ESh

ell

Cittariumpica

2273

84

L003

30–40

59.8

2.6

1410

±40

AD890–1080

Beta-324952

Nev-04

10NCW0212SH

ESh

ell

Cassistuberosa

2273

94

L003

30–40

58.2

2.7

720±30

AD1170–1300

Archaeol Anthropol Sci

Table 2 List of taxa (78 independent species) recovered from trench 2273 at coconut walk

Class Species Weight (g) Percentage of total MNI Percentage of total NISP Percentage of total

Bivalvia (n=11) 5,566.61 3.9 2,706 4.6 5,732 6.7

Anadara floridana 440.20 0.3 21 0.0 24 0.0

Arca zebra 261.89 0.2 33 0.1 37 0.0

Asaphis deflorata 237.39 0.2 24 0.0 63 0.1

Chama sp. 1,880.81 1.3 50 0.1 76 0.1

Codakia orbicularis 955.25 0.7 193 0.3 475 0.6

Donax denticulatus 1,603.52 1.1 2,367 4.0 5,019 5.9

Lucinoma lamellata 92.91 0.1 6 0.0 13 0.0

Mactrotoma fragilis 30.83 0.0 5 0.0 8 0.0

Raeta plicatella 28.40 0.0 3 0.0 12 0.0

Trachycardium sp. 35.09 0.0 3 0.0 4 0.0

Tucetona pectinata 0.32 0.0 1 0.0 1 0.0

Gastropoda (n=66) 121,499.83 86.1 54,176 92.4 65,948 77.4

Architecnica nobilis 120.82 0.1 27 0.0 40 0.0

Bulla striata 3.05 0.1 1 0.0 2 0.0

Cantharus multangulus 0.21 0.0 1 0.0 1 0.0

Cassis tuberosa 134.81 0.1 1 0.0 1

Cerithium eburneum 1.52 0.0 3 0.0 3 0.0

Cerithium litteratum 24.74 0.0 35 0.1 35 0.0

Cerithium muscarum 0.69 0.0 1 0.0 1 0.0

Charonia variegata 36.89 0.0 3 0.0 3 0.0

Chicoreus brevifrons 71.94 0.1 6 0.0 6 0.0

Cittarium pica 54,003.98 38.2 1,063 1.8 6,450 7.6

Columbella mercatoria 20.11 0.0 34 0.1 34 0.0

Columbella rusticoides 3.04 0.0 7 0.0 7 0.0

Conus flavescens 6.20 0.0 4 0.0 4 0.0

Conus regius 179.86 0.1 31 0.1 47 0.1

Conus sp. 169.82 0.1 62 0.1 75 0.1

Conus spurius 286.66 0.2 53 0.1 79 0.1

Crepidula sp. 2.87 0.0 5 0.0 5 0.0

Cyclostremiscus beauii 0.10 0.0 1 0.0 1 0.0

Cymatium sp. 30.98 0.0 11 0.0 11 0.0

Cyphoma gibbosum 2.33 0.0 2 0.0 2 0.0

Cypraecassis testiculus 328.85 0.2 30 0.1 42 0.0

Cypraeidae sp. 14.64 0.0 3 0.0 8 0.0

Diodora listeri 11.66 0.0 3 0.0 3 0.0

Echinolittorina ziczac 82.57 0.1 408 0.7 685 0.8

Engina turbinella 38.21 0.0 33 0.1 33 0.0

Epitonium lamellosum 0.68 0.0 2 0.0 2 0.0

Eustrombus gigas 5,883.62 4.2 100 0.2 464 0.5

Fissurella nimbosa 61.43 0.0 41 0.1 48 0.1

Fissurella nodosa 85.99 0.1 55 0.1 61 0.1

Hemitoma octoradiata 0.75 0.0 1 0.0 1 0.0

Hipponix antiquatus 19.75 0.0 3 0.0 3 0.0

Impages salleana 1.25 0.0 1 0.0 1 0.0

Leucozonia nassa 36.21 0.0 9 0.0 9 0.0

Leucozonia ocellata 31.44 0.0 23 0.0 23 0.0

Lithopoma caelatum 2,288.11 1.6 196 0.3 347 0.4

Lithopoma tuber 14,588.75 10.3 973 1.7 2,841 3.3

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placed into a pot of boiling water where the meat could moreeasily be extracted. The small size of the species makes itnecessary to collect large quantities to provide sufficient cal-ories, though they are comparatively easy to access, harvest,and consume. Research by Chislett (1969) on Barbadosshows that there was no change in the rate of growth forN. tessellata based on the month (i.e., seasonality), whileAxelsen (1968) found a significant change in seasonal growth

rate for N. tessellata in Barbados with similar results byKolipinski (1964) in Florida. The seasonality of this andother mollusk species appears to be dependent on localizedconditions, given that Axelsen (1968) and Kolipinski (1964)concluded that different environmental factors were the causeof seasonal variation.

C. pica, or the West Indian top snail, is the most abundantspecies byweight at CoconutWalk, contributing 38.2% of the

Table 2 (continued)

Class Species Weight (g) Percentage of total MNI Percentage of total NISP Percentage of total

Littoraria irrorata 0.80 0.0 1 0.0 1 0.0

Lottia antillarum 374.36 0.3 608 1.0 754 0.9

Macrocypraea zebra 366.36 0.3 27 0.0 70 0.1

Marginella sp. 0.30 0.0 2 0.0 2 0.0

Mitra barbadensis 0.52 0.0 1 0.0 1 0.0

Monoplex nicobaricus 111.22 0.1 57 0.1 57 0.1

Muricidae sp. 52.99 0.0 60 0.1 60 0.1

Opercula 677.58 0.5 645 1.1 648 0.8

Nassarius vibex 11.85 0.0 18 0.0 18 0.0

Natica sp. 13.15 0.0 10 0.0 10 0.0

Nerita peloronta 29.29 0.0 32 0.1 32 0.0

Nerita sp. 74.37 0.1 99 0.2 439 0.5

Nerita tessellata 30,375.45 21.5 37,591 64.1 38,804 45.6

Nerita versicolor 3,732.33 2.6 5,060 8.6 5,105 6.0

Neritina virginea 34.28 0.0 29 0.0 29 0.0

Oliva sp. 32.11 0.0 12 0.0 15 0.0

Orthalicidae sp. 16.67 0.0 52 0.1 58 0.1

Petaloconhus irregularis 5.05 0.0 2 0.0 2 0.0

Pisanania pusio 1.25 0.0 1 0.0 1 0.0

Polinices lacteus 3.22 0.0 4 0.0 4 0.0

Puperita pupa 3.95 0.0 3 0.0 3 0.0

Purpura patula 300.74 0.2 99 0.2 102 0.1

Rhombinella laevigata 0.88 0.0 3 0.0 3 0.0

Stramonita haemastoma 133.85 0.1 56 0.1 57 0.1

Stramonita rustica 271.99 0.2 126 0.2 127 0.1

Supplanaxis nucleus 170.05 0.1 705 1.2 766 0.9

Tectarius muricatus 1,293.01 0.9 1,804 3.1 2,698 3.2

Tegula excavata 3,350.97 2.4 3,505 6.0 4,364 5.1

Thais deltoidea 1,319.53 0.9 285 0.5 292 0.3

Turbinella angulata 15.10 0.0 1 0.0 1 0.0

Turbo cailletii 0.50 0.0 1 0.0 1 0.0

Turbo castanea 101.70 0.1 40 0.1 41 0.0

Turritella variegata 7.16 0.0 1 0.0 1 0.0

Vasum muricatum 38.08 0.0 2 0.0 2 0.0

Voluta musica 4.64 0.0 2 0.0 2 0.0

Indeterminate shell 9,019.69 6.4 – – – –

Polyplacophora (n=1) 5,109.00 3.6 1735 3.0 13,486 15.8

Chiton tuberculatus 5,109.00 3.6 1735 3.0 13,486 15.8

Total (n=78) 141,195.13 58,617 85,166

The three preferred species are in bold

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weight, 1.7 % of MNI, and 7.6 % of NISP. This taxon is amuch larger species than N. tessellata, with an average lengthof 32–34mm and a maximum recorded dimension of 137 mm(Robertson 2003). The gastropod’s shell is thick and heavywith a pattern of white with reddish/black zigzags (Fig. 3)(Robertson 2003). It is nacreous, however, which makes ithighly susceptible to fragmentation, resulting in many smallerpieces that can still be easily identifiable because of the uniquecolor pattern. At present, C. pica is rarely found along theFlorida coast, with primary habitats situated along the coastsof Mexico south to Venezuela and throughout the circum-Caribbean (Abbott and Morris 1995; Rehder 1981;Robertson 2003). C. pica is a grazer and found in intertidaland shallow subtidal zones with amaximum recorded depth of7 m, though it tends to live in shallower water (Robertson2003). The species is still harvested and eaten in the Caribbeantoday.

The third preferred mollusk, L. tuber, is commonly knownas the green star shell (Fig. 3). L. tuber comprised 10.3 % ofthe total weight, 1.7 % of MNI, and 3.3 % of NISP. Thecurrent distribution of the green star shell is the Gulf ofMexico and Caribbean on rocks in the intertidal and subtidalzones; most range in size between 25 and 75 mm (Abbott andMorris 1995; Rehder 1981). L. tuber is recognizable by itsgreen and white cross-hatched pattern and whorls (Abbott and

Morris 1995). Like the other two key species, the green starshell would have likely been collected, boiled, and then eaten.

Methods

The evaluation of mollusk exploitation by pre-Columbianpopulations living at Coconut Walk included the criteria forover-exploitation (resource depression) as described byClaassen (1986, 1998) and Mannino and Thomas (2002). Asmentioned above, the criteria include the change in richness ofspecies over time and size, age profile, and abundance forpreferred species. Theoretically, in a case of over-exploitation,the criteria should chronologically move from change in sizeto age profile, abundance, and finally richness.

Size and age profile was analyzed through comparing theaverage size (based on weight) of the three preferred species(N. tessellata, C. pica, and L. tuber) over the four plana(layers) of occupation. Research has shown that weight canoften be used as a viable substitute for measured size ingastropods based on research suggesting that size and weightare generally correlated in gastropods (Glassow 2000; Gjerde1986; Jerardino 1997), with some exceptions (e.g., seeGlassow 2000; Mason et al. 1998, 2000; Claassen 2000).We analyzed individual weight (weight/MNI based on non-repetitive elements [NRE] and fragment weight [weight/NISP]) for the three preferred species in the environmentalsquares. These measurements represent the average per unit,preventing us from analyzing variation within a unit, butallowing us to compare averages. We combined results foreach planum to find a trench average, which assumes that theplana in each of the 25 1×1 m squares across the trenchrepresent approximately the same general time period. Theabundance of each species was determined by statisticallycomparing the percentage of molluskMNI and NISP that eachspecies contributed by planum and that should ideally revealthe intensity of mollusk exploitation through time at CoconutWalk. The richness of species over time was measured bycomparing the number of taxa found in each planum.

The use of weight in this study as a unit of analysis is basedupon the taphonomic conditions of the recovered shellfish.

Fig. 3 Image of Nerita tessellata, Lithopoma tuber, and Cittarium pica(left to right, scale bar in centimeters)

Table 3 Descriptive statistics forpreferred species at the CoconutWalk site from environmentalsquares

N. tessellata C. pica L. tuber

Planum 1 Planum 4 Planum 1 Planum 4 Planum 1 Planum 4

Weight (g) 5,046.69 482.15 6,807.94 116.78 818.38 69.5

MNI 6,304 688 120 15 59 7

NISP 6,538 738 986 63 251 27

Individual weight (g) 0.80 0.70 56.73 7.79 13.87 9.93

Fragment weight (g) 0.77 0.65 6.90 1.85 3.26 3.02

Fragmentation (NISP/MNI) 1.04 1.07 8.22 4.20 4.25 3.29

Archaeol Anthropol Sci

Although weight as a measurement of shell size has beendebated within the literature (see Glassow 2000; Masonet al. 1998, 2000; Claassen 2000), it was determined, forexample, that the size (length and width) ofN. tessellata couldin fact be accurately estimated using weight (compared to datafrom Giovas et al. 2013). Patch area, species abundance, andturnover rates are not known for the given locality, but it isassumed that pre-exploitation mollusk population size wasbased on resource availability which would have increasedwith predation. Meat weights and caloric availability for themollusk species were calculated using results from Keegan(1992: Table 6.1), though it should be noted that the averageweight per individual appears to represent a diminished pop-ulation that is smaller than the species found at Coconut Walk.

Results: evaluation of over-exploitation criteria

The first criterion for examining the possibility of over-exploitation is the expected decrease in size of a taxon overtime that would indicate the increased harvesting of lessmature individuals (Claassen 1986, 1998; Mannino andThomas 2002). The length and width measurements of morethan 2,700 N. tessellata specimens statistically analyzed byGiovas et al. (2013) showed an increase in mean and medianshell length from the bottom of the unit in Planum 4 (x =9.96 mm) to the most recent one, Planum 1 (x= 10.44 mm).The trend continued in mean and median shell width fromPlanum 4 (x= 12.70 mm) to Planum 1 (x= 13.27). Because thehigh degree of fragmentation in recovered samples of C. picaand L. tuber prevented a similar analysis from being per-formed, it was necessary to use an alternative for measuringa possible change in size.

Correspondingly, a trend similar to measured size wasfound analyzing the average individual and fragment weightof the preferred species in the environmental squares (7, 9, 17,

and 19) (Table 3).N. tessellata increased in average individualweight (Pl. 4, x = 0.70 g; Pl. 1, x = 0.80 g) and averagefragment weight (Pl. 4, x = 0.65 g; Pl. 1, x = 0.77 g).Likewise, L. tuber had an increase in average individualweight (Pl. 4, x = 9.93 g; Pl. 1, x = 13.87 g) and averagefragment weight (Pl. 4, x= 3.02 g; Pl. 1, x= 3.26 g) over time.C. pica had the largest increase in average individual weight(Pl. 4, x = 7.79 g; Pl. 1, x = 56.73 g) and average fragmentweight (Pl. 4, x= 1.85 g; Pl. 1, x= 6.90 g). The trend continueswhen analyzing the entire trench with the average individualand fragment weight for all three species increasing over time(Figs. 4, 5, and 6). The increase in average fragment weightfor L. tuber and C. pica supports the evidence that individualweight is increasing instead of the number of fragments perindividual. Results demonstrate that all of the three preferredspecies increase in average individual and fragment weightfrom the earliest to most recent plana (see Table 4 for Kruskal-Wallis tests for significance). Though these results suggest theabsence of over-exploitation, we continued to investigate thecriteria for other signs that this may have occurred.

According to the premises required to suggest that over-exploitation was occurring, a decrease in mollusk size isexpected to be accompanied by a change in the age profilevia age being determined by size in most mollusk species.Following an expected decrease in size, an over-exploitedpopulation should show a change in the age profile throughtime. For marine gastropods such as the three preferred spe-cies reported here, this is typically determined through lengthmeasurements as a proxy for age. Because of the preservationof the three most common species, only the tessellated neritecould be measured for age estimation. The length measure-ments reveal an increase in average length (Pl. 4, x= 9.96 mm;Pl. 1, x= 10.44 mm) through time (Giovas et al. 2013), thoughit is still below the average length of 14 to 17 mm in sexuallymature individuals (Chislett 1969; Kolipinski 1964). Based onGlassow (2000) and Mason et al. (1998), we could assume—using weight as a proxy for size—that C. Pica and L. tuber

Fig. 4 Box plots of N. tessellata average individual and fragment weight per unit. Circles represent outliers

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follow trends similar to N. tessellata which show an increasein average size over time.

It might be expected that over-exploitation would producea decrease in preferred species abundance as the average sizeand age profile of the species are declining to create lesssexually reproductive individuals within the mollusk popula-tion. At Coconut Walk, there is a trend of increased abun-dance, as measured through percent NISP and percent MNI ofthe three preferred species, which instead demonstrates a caseof intensified exploitation (see Tables 2 and 5). The increase inabundance (%NISP) for each species in the environmentalsquares was 3.4 % for C. pica, 2.8 % for N. tessellata, and1.7 % for L. tuber. This same trend occurs with percent MNIforN. tessellatawith a 5.4 % increase and 0.8 % in L. tuber. C.pica had approximately the same abundance level with MNIdecreasing 0.1 % (Pl. 4, 1.3 %; Pl. 1, 1.2 %).

The last criterion to determine whether over-exploitationoccurred is a decrease in richness as measured by the numberof taxa. At Coconut Walk, the number of individual taxaincreases from Planum 4 (32) to Planum 1 (61). However,the increase in taxa is correlated with sample size (R=+0.959,

p=0.041), which prevents us from analyzing change in spe-cies richness, though we can speculate that richness remainsrelatively flat because of the correlation with sample size.

While mollusks make up the largest percentage of materialfrom the midden deposit, they are not the only component inthe diet. Coconut Walk is a permanent settlement based onother archaeological evidence such as postholes, with Trench2273 representing only one of several areas of midden depo-sition. Analyzing the caloric contribution to the diet using thetop three species, we found that one person could eat onlymollusks for around 119 days (based on return rates fromKeegan [1992: table 6.1] using NISP as the number of indi-viduals and an estimated consumption of 2,700 cal/day).

Discussion

There are a number of different factors (climate, predators,population density available, food resources, etc.) that couldbe influencing the increase in the individual size of molluskspecies reported here. Many of these factors cannot be easily

Fig. 5 Box plots of L. tuber average individual and fragment weight per unit. Circles represent outliers

Fig. 6 Box plots of C. pica average individual and fragment weight per unit. Circles represent outliers

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analyzed through archaeological evidence, however, and thegeneral paucity of local climatic data also precludes a detailedassessment of how environmental changes may have affectedmollusk populations.

Within the Caribbean, several climatic abnormalities havebeen observed across the region which overlap with prehis-toric occupation at Coconut Walk (Lane et al. 2011). Betweenca. AD 900–950, there appears to be a period of droughtidentified in the paleoenvironmental record (Haug et al.2003), which is broadly consistent with the Terminal ClassicDrought observed in the Yucatan Peninsula (Hodell et al.2005). Following this period, the climate becomes wetter overthe next 300 years (Lane et al. 2009) and follows the trendseen during theMedieval Climate Anomaly (Lane et al. 2011).It is unclear what, if any, effects these changes may havecaused to mollusk populations during the time of occupationat Coconut Walk.

Given the data available from the excavated 5×5 mtrench at Coconut Walk, we attempted to assess whetherwe could discern any impacts on mollusk exploitation overa period of six centuries or so. Because midden depositionlikely reflects predation from numerous sources of shoreposition (and that likely varied through time), the assump-tion is that these remains consist of a reasonable samplewith which to examine trends in subsistence during siteoccupation. Whether this is applicable to contemporaneoussites on the island is presently unknown. While we cannot

completely rule out climatic fluctuations as influencingspecies composition or size of taxa at Coconut Walk, thedearth of locally specific paleoclimatic data—along withquestions relating to how regional trends showing increasedperiods of rainfall would affect these particular species ofmollusks—prevents a clearer understanding of how changesin size of the three species we analyzed may have beeninfluenced by these phenomena.

Regardless, we were interested in better understandingthe exploitation of mollusks at Coconut Walk using thefour criteria outlined above (see Claassen 1986, 1996;Mannino and Thomas 2002) to examine whether inhabi-tants at this Late Ceramic Age site were over-harvestingmollusk resources. These criteria are derived from foragingtheory and resource depression models, whereby increasedpredation is expected to cause a decrease in mean speciessize (Broughton 1994a, b, 1997; Butler 2001; Lupo 2007;Mannino and Thomas 2002). High levels of exploitationamong the tessellated nerite have been shown at otherCeramic Age sites in the Lesser Antilles (Fitzpatricket al. 2009; Newsom and Wing 2004: 87; Serrand andBonnisent 2005; Giovas 2013). The criteria demonstratingwhether over-exploitation of mollusks occurred are notfound at Coconut Walk.

Similarly, two other mollusk studies conclude that there isan increase in preferred species size, but with a decrease inintensity of exploitation. Thakar (2011), for example, found a

Table 4 Kruskal-Wallis test of individual weight

N. tessellata L. tuber C. pica

Comparison Chi-square p value Chi-square p value Chi-square p value

Planum 1–Planum 2 0.083 0.7728 3 0.08326 2.083 0.14890

Planum 1–Planum 3 0.033 0.5637 3 0.08326 2.083 0.14890

Planum 1–Planum 4 5.333 0.0209 0 1 4.5 0.03389

Planum 2–Planum 3 0 1 0.083 0.77280 0.3333 0.56370

Planum 2–Planum 4 5.333 0.0209 2 0.15730 3.125 0.07710

Planum 3–Planum 4 0.750 0.3865 4.5 0.03389 4.5 0.03389

Results that are statistically significant at a 95 % confidence interval are in bold

Table 5 Preferred species NISP,MNI, and assemblage relativeabundance from environmentalsquares

Nerita tessellata Cittarium pica Lithopoma tuber

NISP (%) MNI (%) NISP (%) MNI (%) NISP (%) MNI (%)

Planum 1 6,538 (46.0) 6,304 (65.5) 986 (6.9) 120 (1.2) 449 (3.2) 137 (1.4)

Planum 2 5,169 (38.5) 4,769 (59.9) 499 (3.7) 93 (1.2) 309 (2.3) 94 (1.2)

Planum 3 2,329 (40.1) 2,243 (59.9) 209 (3.6) 42 (1.1) 138 (2.4) 41 (1.1)

Planum 4 775 (43.2) 689 (60.1) 63 (3.5) 15 (1.3) 27 (1.5) 7 (0.6)

14,811 (42.0) 14,005 (62.3) 1,757 (5.0) 270 (1.2) 923 (2.6) 279 (1.2)

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size increase in Pismo clams (Tivela stultorum) in theCalifornia Channel Islands, though this was attributed to ashift to deeper waters for collection.McCoy (2008) showed anincreased size in limpets (Cellana spp.) from Hawaii causedby a massive decline in population and subsequent lesseningof harvests. At Coconut Walk, we found an increase in ex-ploitation (abundance) over time in the three preferred species,concurrent with an increase in size, which suggests some levelof sustainable exploitation (harvests). The results are similar tothose found with the humped conch (Strombus [Gibberulus]gibberulus) in Palau (Giovas et al. 2010), which may berelated to increased horticultural activities on the mainlandthat led to eutrophication and preferential habitat for thespecies. What this demonstrates is that there is a gap in theframework for over-exploitation that excludes the potential forsustainable mollusk exploitation.

The notion of over-exploitation as a continuum suggestedby Mannino and Thomas (2002) appears to be a suitablemodel for explaining this process. Examining exploitationalong a spectrum of possibilities allows for a better under-standing of long-term harvesting, which, as is demonstratedby this study and others (e.g., Cannon and Burchell 2009;Whitaker 2008; Jerardino 2012), is more complex in naturethan might be expected. Yet, when attempting to analyzesustainability within this framework, it becomes muddledwithin the constraints of the model. What is required is anew model of exploitation that accounts for the vast range ofpossible interactions between humans and mollusk taxa as arange from no human interaction, to being sustainably har-vested, and in extreme cases, to extirpation or extinction.Ideally, archaeological data would fit perfectly into a model,yet because of the breadth of locations and cultures, each siteis inherently unique. We emphasize that sustainability is not auniversal term, and must be placed into context both locallyand specifically. A sustainable practice at one locale within agiven period of time may not be sustainable for a longerduration, nor at other sites given myriad cultural and naturalprocesses.

Our investigation at Coconut Walk, along with other recentstudies, suggests that contrary to what would be expected inhuman subsistence strategies if increased predation of selectspecies occurred—in which over-exploitation is the result—mollusks in this large assemblage from the Caribbean actuallyappear to have been harvested sustainably. This suggests thatcurrent models for assessing intensive harvesting of specifictaxon or a suite of taxa in an assemblage should bereevaluated. Further research at Coconut Walk and othercoastal sites on Nevis, and in the Caribbean in general, shouldprovide more information on the temporal framework andspatial distribution of faunal exploitation to aid in evaluatingwhether there are any correlations between cases of sustain-ability (perhaps due to conscious conservation efforts) orover-exploitation regionally.

Conclusions

The exploitation of mollusks was a common feature amongcoastal societies worldwide, and as such, they provide one ofthe best resources for analyzing human-environmental inter-action and subsistence strategies across time and space. Theirnear ubiquity, relative ease of quantification, and frequentharvesting all provide excellent opportunities for archaeolo-gists to investigate these and a host of other issues related tohuman predation patterns. While some researchers have sug-gested that mollusks are uneconomic to collect, relegatingthem to a different level of predation (Keegan 1992:130)—with certain species also known to have been seasonallyrestricted (Kirch 1982; Erlandson 1988)—data shows this tonot necessarily be the case (McCoy 2008; Chislett 1969). Inthe case study of Coconut Walk on the island of Nevisduring the Late Ceramic Age, we found an increase inboth exploitation and size of the three preferred species,suggesting that exploitation at Coconut Walk was generallysustainable between ca. AD 890–1440. This suggests thatwhile mollusks may occasionally be expensive, they werestill increasingly attractive resources to local inhabitantsgiven their relative ease of collection and inability toavoid predation.

Because the criteria for examining over-exploitation vis-à-vis prey-choice modeling (Broughton 1994a, b, 1997;Grayson and Cannon 1999; Butler 2001) do not fall in linewith what is seen at Coconut Walk, we suggest the need for arevision of existingmodels of faunal exploitation that considera much broader range of possibilities and that includes sus-tainability as a potential outcome.While there is an abundanceof evidence that clearly shows prehistoric people in coastalregions around the world were impacting local marine re-sources to the point of depression or even extirpation orextinction (Braje et al. 2012; Butler 2001; Erlandson andRick 2010, ; Faulkner 2009; Jerardino 1997; Mannino andThomas 2001; Morrison and Hunt 2007; Morrison andCochrane 2008)—with the Caribbean being no exception(Fitzpatrick and Keegan 2007; Keegan et al. 2003)—thisoutcome should not be considered inevitable (see Carderet al. 2007; Giovas 2013; Giovas et al. 2013; Thomas 2014).The reasons behind these phenomena are in some cases quitevisible, whereas in others it can be much more difficult totease out the variables that may be causing changes to faunalassemblages. Future investigation of midden deposits in theCaribbean should help ameliorate some of the complexitiesinvolved with assigning causality. Overall, the results of thisstudy suggest the need to revise models of exploitation toemphasize that these processes can take place along a contin-uum that may wax and wane through time. Research dedicat-ed to expanding the commonly used exploitation models, withthe inclusion of sustainability as a potential outcome in mol-lusk and other types of faunal harvesting, will prove useful in

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discerning the degree to which humans interacted with theirlocal environments in the Caribbean and elsewhere.

Acknowledgments We thank the Nevis Historical and ConservationSociety for their support and logistical help in the 2010 fieldwork atCoconut Walk. Thanks also go to the many students who assisted in fieldand lab processing activities on Nevis and at North Carolina StateUniversity and Michiel Kappers and Quetta Kaye who co-directed theproject. The project described in this publication was supported in part byGrant/Cooperative Agreement Number G10AC00624 from the UnitedStates Geological Survey. Its contents are solely the responsibility of theauthors and do not necessarily represent the official views of the USGS.Comments from William Keegan and an anonymous reviewer helped toimprove the clarity and content of the arguments made.

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