a long-term perspective on biodiversity and marine ......617 a long-term perspective on biodiversity...

617

A Long-term Perspective on Biodiversity and Marine ResourceExploitation in Fiji’s Lau Group1

Sharyn Jones2

Abstract: I present research investigating biodiversity and human interactionwith the local environment through three perspectives on diverse islands in Fiji’sLau Group. First, I generated long-term data on marine diversity and exploi-tation through zooarchaeological analyses of fauna from sites spanning the re-gion’s prehistoric human occupation. The study areas are representative ofregional fauna and local geographic variation in island size and structure. Eachisland also varies in terms of human occupation and degree of impacts onmarine and terrestrial environments. Second, my ethnographic work recordedmodern marine exploitation patterns by Lauan communities. Third, marine bi-ological surveys documented living faunas. Together this information is used toexplore the marine environment over the three millennia of human occupation.Using data derived from my multipronged study I discuss potential causes ofecological change in this tropical marine setting. My findings include the follow-ing: (1) data indicate that relative intensity of human occupation and exploi-tation determines modern composition and biological diversity of marinecommunities because human disturbance occurred more extensively on largerislands than on smaller islands in Lau; (2) Lauans appear to have targeted similarsuites of marine fauna across their 3,000 years of history on these islands; (3)Lauans have had a selective effect on marine biodiversity because particular spe-cies are/were targeted according to local standards of ranking and preference;(4) marine resources existing today have withstood over 3,000 years of humanimpacts and therefore may have life history traits supporting resilience andmaking conservation efforts worthwhile.

Marine ecological studies are increasinglytaking into account the biological complex-ity of the past to identify causes of environ-mental change and demonstrate achievablegoals for resource management and restora-tion ( Jackson et al. 2001, Pandolfi et al.2003). Adding an archaeological dimensionexpands the concept of biodiversity by gen-

erating long-term perspectives on human-environmental interactions. The most pro-ductive scientific programs combine multipleapproaches and methodologies to enhanceunderstanding of environmental changes thatare critical locally and globally ( Jennings andPolunin 1996, Kronen and Bender 2007).Resource exploitation viewed through thelens of historical ecology has been exploredthrough recent research on marine and islandecosystems, including, for example, the workof Thomas (2002, 2007a,b) in Kiribati, Fitz-patrick and Donaldson’s (2007) study focusedon anthropogenic impacts in Palau, Rick andErlandson’s (2008) broad collection of casestudies on human-marine environment inter-actions across the globe, and Steadman’s(2006) massive work reconstructing avianhistorical biogeography in the tropical Pacificislands. These works demonstrate that multi-disciplinary historical approaches hold much

Pacific Science (2009), vol. 63, no. 4:617–648: 2009 by University of Hawai‘i PressAll rights reserved

1 Financial support for this research was providedby grants to S.J. from the National Geographic Societyand UAB-NSF ADVANCE. Manuscript accepted 28December 2008.

2 Department of Anthropology, University of Ala-bama at Birmingham, 1530 3rd Avenue S, Birmingham,AL 35294-3350 (e-mail: [email protected]).

promise for the future of research on islandand marine ecosystems worldwide.

Perhaps one of the biggest debates amongarchaeologists and biogeographers who studythe Pacific islands has centered on naturalversus human causes of environmentalchange. Strong arguments have been madeon both sides of the debate (Nunn 2000,Butler 2001, Allen 2006, Steadman 2006,Morrison and Cochrane 2008). Determiningcausality from archaeological data is difficultdue to equifinality, the principle that a singleoutcome—resource change and depletion—can result from different causes. Some re-searchers have concluded that, ‘‘. . . scientistsshould stop arguing about the relative impor-tance of different causes of coral reef decline:overfishing, pollution, disease, and climatechange. Instead, we must simultaneously re-duce all threats to have any hope of reversingthe decline’’ (Pandolfi et al. 2005:1725). Pan-dolfi and colleagues view the methods ofhistorical ecology as especially valuable inproviding comparisons between the past andthe present, while also providing a baselinefor determining if restoration efforts succeed‘‘in ameliorating or reversing change’’ (Pan-dolfi et al. 2005:1726). (According to thisframework, indicators of success would in-clude the return or presence of key groupssuch as parrotfishes, grazing sea urchins, ma-ture complex coral colonies, sharks, turtles,large jacks, and groupers [Pandolfi et al.2005].)

Historical ecology fundamentally deniesan either/or dichotomy of human versusnatural induced changes in the environment;rather, environmental change should be ap-proached through an understanding of the‘‘landscape,’’ the human-environment inter-action sphere (Crumley 1994, Balee 1998,Balee and Erickson 2006). This perspectiveargues that the landscape is imbued withmeaning and is the product of the collisionbetween nature and culture, which may be ex-amined as a form of architecture, or materialculture (Balee and Erickson 2006). I adoptthis approach, extending the concepts ofhistorical ecology and landscape to the ‘‘sea-scape,’’ associated with islands in Fiji’s centralLau Group.

My multidisciplinary research in the LauIslands contributes a more robust under-standing of biodiversity and long-termhuman interactions with the local marine en-vironment through three perspectives on fourdiverse islands. This multipronged approachincorporates data derived from ethnographic,archaeological, and marine biological re-search with the aims of characterizing thereef environment, documenting species andbiological diversity, and identifying potentialchanges in the marine ecosystem. The Fiji Is-lands are rich in cultural and biological his-tory, with human occupation extending backto approximately 3100 B.P. (Nunn et al.2004, Nunn 2007). Fiji’s Lau Group is an ar-chipelago of 80 islands, located about 200 kmeast and 100 km south of the main Fijian is-lands of Viti Levu and Vanua Levu (Figure1). The Lau Group is composed of volcanicand coralline limestone islands that are lo-cated relatively close together and support ex-tensive reef systems, rich in marine faunalresources. The region has an incredible diver-sity of marine life and a vibrant traditionalculture that is actively engaged in marine-oriented subsistence activities, making it anideal location to focus research examiningmarine biodiversity and human exploitationof marine environments over a broad tempo-ral scale. Food is still largely locally derived inthis contemporary Pacific community, and,compared with other areas in Fiji, the coralreefs of Lau have had relatively little impactby tourism, extensive modern developments,commercial fishing, and high populationlevels. Moreover, central Lau has been thesubject of archaeological and ethnographicresearch by myself and others, which has laidthe groundwork for this project (Hocart1929, Thompson 1940, Best 1984, O’Day etal. 2003, O’Day 2004, Thomas et al. 2004,Jones et al. 2007a,b).

To explore biodiversity and marine exploi-tation over the past three millennia, thisstudy investigated four diverse islands in theLau Group, including Lakeba, Nayau, AiwaLevu, and Aiwa Lailai (Table 1). Each islandvaries in terms of human occupation and thedegree of impacts on marine and terrestrialenvironments. Lakeba is the largest of the

618 PACIFIC SCIENCE . October 2009

study sites (55.84 km2) and is occupied byabout 2,100 people. Nayau (18.44 km2) hasa population of approximately 400 people. Incontrast, the small islands of Aiwa Levu (1.21km2) and Aiwa Lailai (1.0 km2) are currentlyuninhabited, lack a freshwater source, and areused primarily as temporary camps for fisher-people from Lakeba (12 km away). Geologi-cally the Aiwas consist entirely of limestoneand have very little agricultural potential.Lakeba is the most varied topographicallyand geologically, having limestone regionsand large areas of volcanic soils and bedrock(andesitic and dacitic lava) where freshwateris locally available and agriculture is practicedextensively. People living on Lakeba andNayau maintain dryland and wetland agricul-tural crops (especially taro, cassava, sweetpotato, and yam). Conversely, Aiwa supportsdiverse native forests with little if any area

suitable for agriculture ( Jones et al. 2007b).The archaeological sites on these four islandsspan central Lau’s prehistoric occupation, ex-tending from the Lapita period to Europeancontact in 1774 (Best 1984, Jones 2007).Like Lakeba, Nayau is almost entirely modi-fied by human occupation, with continuousman-made features across the landscape (in-cluding gardens, habitation sites, trails, andresource-collection areas). The Aiwas alsoexhibit signs of use islandwide, but this islimited to light surface scatters of artifactsand ecofacts (stone tools, Tridacna shells, andmidden, for example). The reefs surroundingeach of the four islands are similar in termsof structure and potential habitat. However,modern development and occupation onLakeba are much more intensive than thaton Nayau (and the uninhabited Aiwas), there-fore Lakeba’s inshore reefs are likely the most

Figure 1. Map of the Fiji Islands with the study area indicated in the square.

Long-term Perspective on Biodiversity in Fiji . Jones 619

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heavily exploited of the study islands. It isinteresting that Lauans travel to exploit reefsnear their islands. This was very likely thecase in the past as well. For example, 17.7km east of Lakeba is an area of shallow waterwhere a reef encloses a lagoon; this reef iscalled Bukatatanoa, a well-known Lakebanfishing ground that is five times the size ofthe reefs surrounding Lakeba. Fisherpeopletravel there in small boats almost daily.Although the most obvious and frequentlyexploited marine areas are those in closeproximity to the villages, Lauans do not lookupon the sea as a barrier; rather they utilize avariety of environments and frequently movebeyond their home shores.

materials and methods

My research contributes to the understandingof biodiversity through three perspectives.First, zooarchaeological analyses of animalbones and shells provide long-term data onmarine diversity and exploitation. Second,ethnographic work recorded modern marineexploitation patterns by Lauan communities.Third, I compare my archaeological andethnographic data to information derivedfrom the examination of living marine faunasthrough detailed biological surveys of thereefs surrounding the study sites. Togetherthis information is used to understand andcharacterize the causes and rates of ecologicalchange in this tropical marine context overthe past 3,000 years. I view the data describedherein as a starting point for the understand-ing of these complicated processes. Althoughresearch on these topics is preliminary, I havebegun to generate a description of the marineenvironment and human exploitation patternsbased on three datasets. At this point my pri-mary goal is to determine if there is evidencefor changes in marine biodiversity and, if so,what the relationship is between these shiftsand human exploitation over time in each ofthe island settings.

Archaeology

Since 2000, 23 sites on the four islands havebeen excavated and analyzed as part of this

research project (O’Day et al. 2003, Jones2007, Jones et al. 2007b). Archaeologicalwork has generated a basic chronology of hu-man occupational history on each island. Allexcavations were conducted in 10 cm levels,using trowels and following natural stratigra-phy whenever possible. All sediments weredry screened through nested sieves of 1/2inch (12.8 mm), 1/4 inch (6.4 mm), 1/8 inch(3.2 mm), and 1/16 inch (1.6 mm) mesh.

Zooarchaeological analysis followed stan-dard procedures described in O’Day (2001)(publications by me from 2001 to 2004 arelisted under my former name of SharynO’Day; fish bone identifications were basedon all diagnostic elements [not limited to so-called ‘‘special elements’’], including atlases,vertebrae, cranial elements, spines, scales,and ‘‘special elements’’) and utilized my com-parative collection housed at the Universityof Alabama at Birmingham’s zooarchaeologylaboratory. (I have assembled a comparativecollection of animal bones, especially fishesand invertebrates [>2,500 specimens], toidentify Pacific archaeological materials tothe most specific level possible. The collec-tion includes multiple individuals of the mostcommon central Pacific families, genera, andspecies and a variety of size and age classes.The majority of these fishes are from Fijiand Lau in particular, but the collection alsoincludes fishes and invertebrates from numer-ous islands in Micronesia, the Philippines,Tahiti, Hawai‘i, Okinawa, and the Carib-bean.) The bones of all taxa were counted,weighed, and modifications recorded. Thenumber of identified specimens per taxon(NISP) is the basic specimen count used; thisincludes bones and shells that were notidentified to a specific taxonomic level. Theminimum number of individuals (MNI) wasdetermined by paired elements and size andage classes whenever possible, following cal-culations described by Reitz and Wing(1999) and O’Day (2001). (Specifically, I esti-mated MNI based on paired elements and evi-dence for symmetry, age and/or body sizes.MNI was estimated for the lowest taxonomiclevel within a systematic hierarchy. Each pro-venience [feature, natural stratigraphic layer,and site]) was considered separately with at-


tention to stratigraphy when appropriate, inan effort to generate conservative MNIs.Within a given provenience [layer, unstrati-fied unit, or feature] excavation levels wereconsidered together to estimate MNI.) Mea-surements of faunal remains, particularlythe fish bones, were taken in an effort to illu-minate long-term changes in the size andmakeup of exploited marine species assem-blages. The anterior width of complete fishvertebral centra from both identified and un-identified remains was measured for compar-ative purposes and to estimate the averageweight and size of fishes in the assemblages.This procedure is based on the assumptionthat both the identified and unidentified fishvertebrae represent a cross section of the spe-cies present in the assemblage (Pandolfi et al.2003, Newsom and Wing 2004).

Shannon’s H, diversity or niche breadthmeasure, was used to quantitatively determinethe variety of animals consumed or exploitedat each of the analyzed sites over time. Theformula used follows Krebs (1999:463).(H 0 ¼ �S½ pi�½loge pi� is the formula for Shan-non’s H, where H 0 is the information contentof the sample and pi is the relative abundanceof individuals or resources for each taxon inthe collection.) Because this diversity measureranges from 0 to y, it can be standardized ona scale from 0 to 1 by using an evenness orequitability measure ðV 0Þ. Equitability ðV 0Þ,or the evenness with which taxa were ex-ploited, and the relative importance of eachtaxon was also calculated. (The equitabilityformula used in the analysis is following Reitzand Wing [1999]. Because this is a compari-son of fauna representative of a wide rangeof organisms that have varied numbers of ele-ments, diversity and equitability calculationsare based on MNI estimates; this places di-verse organisms on a more uniform basis.)V 0 is calculated on a scale of 0 to 1, thusvalues close to 1 indicate an even distributionof taxa, and lower values suggest a dominanceof one taxon.

Ethnography

Modern Lauan fishing expeditions providedopportunities to record species diversity and

the sizes of collected modern fishes. In total,the ethnographic aspect of my research wasconducted over 10 months in Fiji between2001 and 2007. (Over the course of my re-search I devoted about 3 months full-time toarchaeology and the remainder of my time toboth archaeology and ethnography, with themajority of my time spent doing ethnogra-phy.) Fieldwork ranged in duration from 1month to 4 months. The fish catches alsoserve as population cross sections of the exist-ing fauna because Lauans frequently collect awide range of inshore species with gill nets,which are not size and species selective. Forthis study, I recorded the numbers of moderntaxa that are collected and eaten on each is-land. A variety of fisherpeople were inter-viewed to determine what marine resourcesare targeted and why, and to discuss localconservation issues. I recorded how the catchis processed, divided up, consumed, and dis-carded.

Over the course of this research onLakeba, Nayau, and Aiwa, I accompaniedfisherpeople, including individuals andgroups, on over 50 fishing expeditions to re-cord information on modern marine-orientedsubsistence activities. Different types of fish-ing trips were documented (in notes, video re-cordings, and still photos) at a variety of timesand locations on all four of the study islands.Following Thomas (2002:186), fishing expedi-tions were formally documented by collectingthe following information: (1) members com-posing group; (2) collection strategy; (3) occa-sion for which the group is collecting; (4)target prey; (5) actual prey, location, count,weight, and SL (standard length) of collecteditems; (6) collection area (marine zone, typedescription, location); (7) date, time, moonphase; (8) search time; (9) handling time, pro-cessing time; (10) general weather and tidalconditions; (11) how the catch is divided up;(12) items consumed on shore or during fish-ing; (13) patterns of flesh and tissue disposalon the beach; (14) who will consume the fish.

Biological Surveys

Using both formal surveys with standardizedmethods and informal surveys, marine biolo-


gists and I inventoried the marine vertebratesand invertebrates on each island (Maragosand Cook 1995, DeVantier et al. 1998). A va-riety of coastal ecosystems (inshore, forereef,mangrove, sea grass flats, bays, fringing reefs,and platform reefs) were surveyed and de-scribed. The majority of survey transectswere positioned on reef flats, areas that arefrequently fished when in close proximity tovillages. Inshore areas located farther fromthe villages and on the currently uninhabitedislands of Aiwa Levu and Aiwa Lailai werealso sampled. In total, 18 sites were surveyedwith line-transect methodologies on Lakeba,Nayau, Aiwa Levu, and Aiwa Lailai ( Jones2009a). Transects on Lakeba measured 200m long and those on Aiwa Levu, Aiwa Lailai,and Nayau measured 100 m in length fromshore, and all transects extended 10 m deep.Visual scans for fish extended 5 m on eitherside of the transect line and above to the sur-face of the water. Invertebrate scans includedan area extending 1 m on either side of thetransect line. Five forereef scuba dives wereconducted around Lakeba, ranging from 18to 26 m deep. In addition, 15 informal sur-veys were conducted on Lakeba, Nayau,Aiwa Levu, and Aiwa Lailai at various tidelevels and during both day and night.

results

Archaeological Research

radiocarbon dating and chronol-

ogy. Earlier archaeological research onNayau, Aiwa Levu, and Aiwa Lailai developeda chronology for the islands by obtainingaccelerator-mass spectrometer (AMS) radio-carbon (14C) dates from seven sites. TheNayau chronology was based on six AMSdates from excavations of 12 mid-late prehis-toric archaeological sites (mid-late period ar-chaeological sites in my study area includeprimarily fortified sites and rockshelter occu-pations ranging in time from about 600–280cal B.P. Most of these radiocarbon dates clus-ter around 710–540 cal B.P.) and thereforeextended only to ca. 700–600 cal B.P. (O’Dayet al. 2003). On Aiwa Levu and Aiwa Lailai,11 AMS dates provided evidence of over two

millennia of human activity and occupation,extending from 2,710 to 10 cal B.P., withmost of the dates suggesting site occupationsbefore 500 cal B.P. ( Jones et al. 2007b). How-ever, none of these dates is from Lapita occu-pations; therefore my recent determinationsfrom the Lapita-period site of Na Masimasiare critical for providing a more completechronology of the study area, with datesranging from the first human occupation ofthe islands through the contact and historicperiods. Radiocarbon determinations fromNa Masimasi were provided by Beta Analytic,Inc., Miami, Florida, and are on a single boneor piece of charcoal (Table 2). The conven-tional 14C age is adjusted for 13C/12C ratios.(Calibration for atmospheric variation in 14Cfollows OxCal version 3.3 and INTCAL98Radiocarbon Age Calibration [Bronk Ramsey1995, Stuiver et al. 1998].) The two dates aresimilar, according to a conventional 14C agedetermination, suggesting an early occupa-tion of Nayau by about 2,580G 40 yearsB.P. (A recent article by Beavan Athfield etal. [2008] discussed potential problems withthe radiocarbon dating of human bones andsuggested that these same problems apply tothe dating of chickens, dogs, and pigs, ani-mals that like humans have diets of mixedterrestrial and sea biota. The authors arguedthat it is necessary to predetermine the typeof diet associated with both humans and do-mestic animals that are processed for radio-carbon dates via isotope studies. This shouldbe done in advance of interpreting the radio-carbon results. I have submitted 61 archaeo-logical samples of bone from a broad rangeof Lauan fauna for isotopic analysis. The re-sults are currently pending, but I plan to usethe data to evaluate the Beavan Athfield et al.arguments in relation to data from centralLau.)

zooarchaeological data. To com-pare local marine exploitation patterns andprehistoric environments on each island overthe past 3,000 years, I have summarized thearchaeological sites excavated, radiocarbondates, and the total number of fishes andinvertebrates identified in terms of count(NISP), minimum number of individuals(MNI), measures of diversity and equitability


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610

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au,

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kutu

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280G

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4054

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420

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380

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Pteropustonganus

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aL

evu

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ock

shel

ter

12,

I/1

180G

40�

19.2

280G

4047

0–

280

(.91

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150

(.04

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4258

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aL

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shel

ter

14,

II/3

260G

40�

1936

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osapiens

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410

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290

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270

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150

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aL

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ave

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I/1

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40�

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960G

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760

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aL

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II/2

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sA

iwa

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u,

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180G

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620

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330

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cksh

elte

r1,

II/5

370G

40�

20.3

450G

4055

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430

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330

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aL

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auR

ock

shel

ter

1,II

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0G

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1721

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sA

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lai,

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cksh

elte

r2,

IV/5

2,06

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502,

460

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150

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(H and V ), and the most common fish fami-lies identified (Table 3) as these data relate togeneral periods of prehistory. (The NISPlisted in Table 3 includes fish bones thatwere not identifiable below the level of class;the percentage of specific fish identifications,that is, to the level of family and below, variesfrom 7% to 26%, depending on the site. Forexample 23% of the fish bone from the mid-late period sites from Nayau was identifiableto the level of family, genus, or species.) Thesame classes of data are provided for inverte-brates in Table 4. These tables provide com-parisons of the marine resources that wereexploited prehistorically over the entire pre-historic human occupation of the study sites;the detailed individual site data are publishedin O’Day et al. (2003), Jones et al. (2007a),and in a forthcoming manuscript currentlyunder review ( Jones 2009b). Although theargument could be made that the amalgam-ation of data from multiple sites, especiallythe 12 mid-late period Nayau occupations, istoo general to contribute useful information,I argue that the summary H and V 0 valuesare both relevant and appropriate for thefollowing reasons. Intersite variation ofidentified fish remains is minimal within thetime periods. Moreover, within the mid-lateNayau assemblages, 80% of the identifiedfish bones by count were recovered from twovirtually contemporaneous fortified rockshel-ter occupations (Waituruturu East and Wai-turuturu West produced the majority of themid-late period Nayau fish remains; thesesites also contributed over half of the MNI,87 individuals [see O’Day et al. 2003].) Themid-late period sites include fortified occupa-tions and rockshelters, all of which likely hadsimilar functions and durations of occupation.Finally, the mid-late Nayau occupations alloccur inland, away from the coast, and accord-ing to the zooarchaeological analysis the occu-pants of all these sites exploited the inshoreareas as their primary animal food source.

The comparative abundance of fauna interms of primary and secondary data is evi-dent in the total amalgamated NISPs andMNIs from sites representative of each timeperiod. The Lapita site of Na Masimasi onNayau has a relatively low overall MNI (59)

for bony fishes (Figure 2). In contrast, theNISP from the site (7,570) is very high andabundant enough to interpret the marinefaunal assemblage with some degree of confi-dence (Reitz and Wing 1999). The low num-ber of MNI from Na Masimasi is due to anextremely high frequency of vertebrae in theassemblage and the somewhat preliminarynature of fish bone identifications. I antici-pate that the fish MNI will increase meaning-fully when my analysis is complete.Nevertheless, the other sites yielded ampleMNI, and all of the sites produced sufficientnumbers of NISP to conduct interpretativeanalyses. There are four major trends thatare evident in the faunal data.

First, the identified vertebrate taxa (and es-pecially the families of fishes exploited) fromthe early to late period sites are dominated bytangs, grouper, parrotfishes, triggerfishes, andemperorfishes (Table 3, Figures 2–5). Thesefamilies are common in Pacific island archae-ological assemblages and in material identi-fied from Fiji (Leach and Davidson 2000,Thomas et al. 2004, Jones et al. 2007b). Por-cupinefish occur in high frequencies in theearly period sites on Lakeba and Nayau butwere less frequently identified in sites frommid-late time periods and in deposits fromother islands. Second, for fishes, the V 0 valuessuggest that there is an even distribution oftaxa (note that values close to one indicate aneven distribution). The marine vertebratetaxa exploited remained even over time, anddiversity ðHÞ was moderate to high, rangingfrom 0.88 to 2.03. The number of taxa re-mains high across temporal periods (28–33).

Third, for invertebrates, the diversity andequitability measures are variable (H rangesfrom 0.88 to 2.03 and V 0 ranges from 0.25to 0.6), much more so than in the fish data(Table 4). Sites associated with the earlytime periods on Nayau and Aiwa producedassemblages dominated by particular species(Figure 6); for example, Na Masimasi’s inver-tebrate assemblage is primarily composed ofthe small fighting conch (Strombus gibberulus).On Aiwa Levu and Aiwa Lailai, the inverte-brates are dominated by turban snails (Turbospp.) throughout the sites (Figures 7 andFigure 8). Aiwa’s sites have high diversity


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levels comparable with those of the mid-lateperiod Nayau sites (Figure 9). Exploitationpatterns in terms of diversity and equitabilityfor invertebrates are different than those of

fishes. The number of taxa represented inthe invertebrate assemblages varies from 18to 35; again, this is more variable than thatdocumented in the fish assemblages.

TABLE 4

Summary of Invertebrate Data from Archaeological Sites on Aiwa and Nayau (Sites Listed from Early to Late Period)

Site Island Age Type NISP MNINo. ofTaxa H, V 0

DominantInvertebrate Taxaa

DauRockshelter

Aiwa Early (2,460–1,310 calB.P.)

Rockshelter 1,804 782 18 1.71, 0.6 Turbo sp., Modiolusauriculatus,Cellana spp.

Na Masimasi Nayau Early (2,610–2,760 calB.P.)

Beach site 1,732 1,026 32 0.88, 0.25 Strombidae (esp.Strombusgibberulus),Turbo spp.

12 sites Nayau Mid-late(600–280cal B.P.)

Fortifiedsites androckshelters

2,625 1,703 35 2.03, 0.57 Turbinidae (esp.Turbo spp.),Cypraea spp.,Strombus spp.,Modiolusauriculatus

Aiwa 1 Aiwa Mixed(2,370–280cal B.P.)

Rockshelter 2,726 1,001 21 1.63, 0.54 Turbo sp., Cellanaspp., Chitonidae

Note: Biomass was based on the weight of the shell.a Invertebrates listed in rank order of abundance.

Figure 2. Early Nayau fish remains from the Lapita site of Na Masimasi, listed by family.


Fourth, it is surprising that measurementsof more than 2,500 fish vertebrae, used as aproxy for fish body size and weight, revealthat the fishes collected and consumed at the

earliest occupations sites were actually smallerthan those consumed later in prehistory(Table 5). This trend is particularly evidenton Nayau where the Lapita site of Na Masi-

Figure 3. Mid-late Nayau archaeological fish taxa, listed by family.

Figure 4. Early Aiwa Levu and Aiwa Lailai fish remains, listed by family.


Figure 5. Mid-late Aiwa Levu and Aiwa Lailai fish remains, listed by family.

Figure 6. Early Nayau shellfish taxa from the Lapita site of Na Masimasi, listed by family.

Figure 7. Early Aiwa Lailai shellfish remains, listed by family.

Figure 8. Summary of shellfish from the site of Aiwa 1, Aiwa Lailai, representing mixed time periods. Data listed byfamily.

masi contained the majority of small bones(mean anterior vertebral centra width 2.98mm and mean estimated weight 118 g), rep-resenting small-bodied fishes. (I do not attri-bute this variation in the size of fishes todifferences in recovery of fauna because Idirected the excavation and screening of thestudy sites and used the same nested screens

in each context [sieves with 1/2 inch (12.8mm), 1/4 inch (6.4 mm), 1/8 inch (3.2 mm),and 1/16 inch (1.6 mm) mesh].) Na Masimasialso produced the broadest range in the sizeof fishes with vertebral centra measuring0.61–20.2 mm, although the vast majority offish vertebrae are small. Mid-late period siteson Nayau produced a smaller range in the

Figure 9. Mid-late Nayau shellfish taxa, listed by family.

TABLE 5

Fish Vertebral Centrum Widths (mm) from Early and Mid-Late Period Occupations on the Study Islands

Provenience n Mean RangeStandardDeviation

Mean EstimatedWeighta (g)

Early Nayau 1,432 2.98 0.61–20.2 1.76 118Early Aiwa 281 4.65 1.5–13.0 2.05 363Mixed Aiwa 174 4.2 1.7–14.3 2 281Mid-late Nayau 464 4.3 3–18.7 3.9 298Mid-late Aiwa 165 4.85 2.04–13.33 1.9 404

Total 2,516 4.2 0.61–20.2 — 281

a Estimates of total body weight (g) are made using the allometric formula log Y ¼ 2:53ðlog X Þ þ 0:872, where Y is the bodyweight and X is the vertebral width, following Newsom and Wing (2004:71).


size of fishes exploited, a mean estimatedweight of 298 g, and a mean vertebral centrasize of 4.3. The fish remains from Aiwa ex-hibit a less-dramatic variation in size overtime (the mean estimated weight is 363 gwith a 4.65 mm centra width at the earlyperiod sites and 404 g and 4.85 mm in themid-late period sites).

Ethnography

Over 50 fishing expeditions were documentedto record information on modern marine-oriented subsistence activities. Eleven fishingexpeditions were recorded on Lakeba, andseven expeditions were recorded on AiwaLevu and Aiwa Lailai. A total of 37 hr of fish-ing was recorded (about 155 person-hours)on Lakeba. On Nayau, nine fishing trips, cov-ering a total of 20 hr (1,140 person-hours),were formally documented.

Ethnographic investigations on the islandof Lakeba in August 2007 confirmed thatLauans collect and consume virtually everyfish that is caught by nets or hand lines inthe inshore area. Shellfish are sometimes col-lected and consumed on the reef, but theyform a very minor component of the catchand marine portion of the diet overall. Thesefindings support conclusions derived fromprevious research on Nayau (O’Day et al.2003, O’Day 2004, Jones 2007). The totalnumber of fish taxa recorded in modern fish-ing expeditions is 112; 105 of these fishes areeaten, and only seven species are bycatch,which are not consumed (see Appendix). I re-corded the numbers of modern taxa that arecollected and eaten on each island; the resultsare as follows: Aiwa Levu, 24; Aiwa Lailai, 15;Lakeba, 49; Nayau, 94. (Obviously, these datadescribe only a portion of the taxa exploitedand are limited by the fact that I recordedfishing and collection over the duration ofmy fieldwork rather than a lifetime of obser-vation.) These data also suggest that themajority of fish taxa are collected with netsinshore (85.7% of the 112 fishes).

Lauan fishing expeditions that utilize netsinshore produce a broad cross section of theexisting inshore species. On a single expedi-tion the fishing group will exploit numerous

areas that range in depth and substrate. Theyutilize their nets in various ways, for example,to corral the fish or to chase the fish intothe nets. Net fishing is conducted at varioustimes, day and night, and in every season.When and where people fish depends on theoccasion that the fisherpeople are collectingfor, the target taxa (if any), the moon phase,and the tidal phase, among other social issues.The important point is that net catches pro-vide an extensive array of fish taxa that maybe used to understand biodiversity of bonyfishes inhabiting the inshore area.

ethnographic interviews. A numberof important issues were brought up inethnographic interviews with fisherpeople onLakeba and Nayau (Table 6). Findings in-clude information on a wide range of issues,such as the style of fishing particular to cer-tain environments and villages, variations inthe way men and women fish, the types offishes preferred by Lauans, and shifts in thelocal availability of certain resources.

Traditional vono style fishing is the pri-mary mode that people in the northern vil-lages on Lakeba (Nasaqalau and Vakano) useto fish inshore; they state that this has beenthe case for many generations. The methodproduces thousands of small inshore fishesover the course of 2 to 3 days (especiallyrabbitfishes, or Siganus spp.; emperorfishes/Lethrinidae; and convict tangs). Vono is docu-mented in detail in a short ethnographicfilm that I and other individuals involvedin this research project helped to write andproduce. (The fishing expedition docu-mented in the film can be viewed on GoogleVideo at http://video.google.com/videoplay?docid=6257583410432053434&q=vono+fiji&ei=q6eQSOPfAYSGqwPZtt2iCA.) In mod-ern Fiji, this traditional fishing method isonly practiced in the Lau Group on Lakeba.Like all inshore fishing in the region, thiscomplex net method is organized and run en-tirely by women. Many Lauan women spendpart of each day on the inshore reef, collect-ing marine resources. However, in each vil-lage there is a clear division between the ‘‘seapeople’’ (the fisherpeople, yavusa wai or kaiwai) and ‘‘land people’’ (primarily farmers,who rarely if ever fish, the yavusa vanua, or


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kai vanua). Women of the sea people clan arethe keepers of traditional marine ecologicalknowledge (TEK). They hold a wealth of in-formation about marine ecology and biota,and have intimate knowledge of the naturalorder as well as changes and fluctuations inthe system. According to informants, the seapeople have always been the fisherpeople ofLau, from the time of the first human occu-pation. They are the descendents of the firstLauan fishers and the people who collectedthe marine resources that occur in ancient ar-chaeological sites. I hypothesize, although itwould be difficult or impossible to prove,that women were the primary producers ofinshore marine resources in prehistory, justas they are in the present.

Small inshore fish species make up thelargest portion of all animal protein con-sumed by Lauans each day. For example, theconvict tang (A. triostegus triostegus) is a favor-ite fish of Lakebans. It is small, measuringan average of only 18–22 cm in total length(TL). This fish is locally considered to be as-sociated with the high chief of Lakeba andthe Lau Islands, or king, the late Tui Lau, be-cause he was very fond of this fish and com-missioned vono expeditions whenever he wason Lakeba. People on Nayau also claim tolove eating the convict tang ( Jones 2009a).Interviews indicate that the majority of sur-vey respondents on Nayau favor inshore reefspecies that are typically small-bodied in themodern environment (a35 cm in TL). Thesingle most sought after fish among women Iinterviewed is a group referred to collectivelyas kawakawa (small groupers, including Epi-nephelus merra and Cephalopholis sp., thehoneycomb and hind groupers). Epinephelusmerra in Fiji measure 20 cm (TL) on averagewhen caught, and Cephalopholis averages 24–40 cm in TL (Froese and Pauly 2006).

Most of those interviewed from Lakebaand Nayau stated that they have noticed a de-cline in the local availability of some fishesand invertebrates (Table 6). For instance, thevillagers of Vakano and Nasaqalau on Lakebacomplained that people from other villagesfrequently come to their inshore reefs to col-lect fish. This practice is becoming increas-ingly common as the marine resources in

larger villages on Lakeba, such as Tubou, arein more obvious decline. Lauans, and espe-cially the fisherpeople, are keenly aware ofworldwide environmental shifts such as globalwarming, the heating of the oceans, and sea-level rise. The long-term sustainability ofLauan fishing practices is something that allthe fisherpeople I interviewed on Lakebamentioned, and in each village on Lakebathere are active representatives from the FijiMinistry of Fisheries. This is not the case onNayau, where there is comparatively littlediscussion of conservation and the effects ofglobal warming; however, people are awareof changes in the marine environment. Thelong-term impacts of intensive marine har-vesting, fishing, and collecting are a frequentsubject of conversation among the villagechiefs and fisherpeople.

Biological Surveys

Over 200 species of marine vertebrates(sharks, stingrays, sea snakes, bony fishes,and turtles) from over 50 families, and almost200 species of macroinvertebrates, were ob-served around the study sites. The reefs ofAiwa Levu and Aiwa Lailai have the widestrange and abundance of observed vertebratesand invertebrates (Table 7). The detailedresults of this biological survey will be pub-lished in the future. Most of the identifiedfishes are relatively common in the waters ofthe tropical Pacific (Myers 1991) and belongto the most abundant coral reef families in-cluding the following: Acanthuridae, Chaeto-dontidae ( butterflyfishes), Labridae (wrasses),Lethrinidae, Mullidae (goatfishes), damsel-fishes (Pomacentridae), Scaridae, Lutjanidae(snappers), and Serranidae. On average, theinshore scuba and snorkel surveys identified41 fish species on Lakeba (per 1,000 m2

area), 55 species on Nayau (per 500 m2), 67species on Aiwa Levu (per 500 m2), and 66species on Aiwa Lailai (per 500 m2).

The presence of juvenile fishes in the in-shore areas varies according to differences inthe marine environments, including the tide,the moon phase, and the substrate. On thenorth side of Lakeba, villagers have plantedextensive areas of mangrove trees. This ap-


pears to have increased the local abundanceof fishes on the reef flats. Juveniles of severalspecies were frequently recorded in closeproximity to the planted mangrove trees, andthey are especially abundant around AiwaLailai and in the intertidal area of AiwaLevu. Aiwa Levu’s mangroves also act asnurseries for juvenile fishes. Around Nayau,the presence of young fishes appears to behighly dependent on the tide, but the inshoreflats around all three of Nayau’s villages boasthigh frequencies of juveniles in the familieslisted here.

The effect of harvesting invertebrates isclearly visible on most of Lakeba and Nayau’sreef flats. Species of giant clam (Tridacnaspp.) and large snails (e.g., Trochus spp.) arerarely encountered (note that when large-bodied invertebrates such as Tridacna andTrochus are encountered by Lauans they areimmediately collected and often consumedright on the reef ). In contrast, on Aiwa Levuand Aiwa Lailai, where regular harvesting isless common, several species that are rare onLakeba flourish (Tridacna spp.). The majorityof forereef dive sites had less than 5% coralcover, which is likely due to a recent bleach-ing event. Since records began, the first time

the reefs of central Lau were affected bybleaching was in 2003 (Leon Zann, Univer-sity of the South Pacific, pers. comm., 2006).(Apparently Lauan reefs were minimally af-fected or unaffected by the 1997/1998 ElNino–Southern Oscillation mass-bleachingevent.)

On both Lakeba and Nayau the reefs gen-erally exhibit a notable lack of gorgoniansand neptheids (soft tree corals, finger softcorals, etc.). The environments evidence se-vere leather soft coral diebacks (Sinularia,Sarcophyton, and Lobophytum spp.). The scaleand condition of the coral bleaching anddeath on Lakeba and Nayau is so severe thatalthough local fishing pressures over the last3,000 years have undoubtedly affected it, re-cent climatic shifts are likely to blame. Never-theless, small coral recruits were documentedand thus natural partial recovery of oncelarger colonies is beginning at all the studyreefs. Despite the poor state of much of thecoral on these islands, substantial populationsof fishes were observed in pockets of healthyreef and in association with the recoveringreef colonies, including large-bodied speciesof groupers, juvenile groupers and parrot-fishes, wrasses, large-bodied parrotfishes, and

TABLE 7

Summary of Marine Biological Survey Data from Lakeba, Nayau, Aiwa Levu, and Aiwa Lailai

Island Survey SiteSurvey Type, Transect

MeasurementTotalFish

No. ofFish Species

Lakeba Oru Scuba, 200 m/1,000 m2 988 54Lakeba Tubou Snorkel, 200 m/1,000 m2 711 37Lakeba Nasaqalau Mangrove Walked, 200 m/1,000 m2 37 2Lakeba Waciwaci Snorkel, 200 m/1,000 m2 807 33Lakeba Nukunuku Mangrove Walked, 200 m/1,000 m2 27 3Lakeba Vakano Mangrove Walked, 200 m/1,000 m2 210 16Lakeba Selesele Point Forereef scuba, 18 m maximum depth 366 54Lakeba Vakano Forereef scuba, 20 m maximum depth 339 54Lakeba Ucuiboagi Point Forereef scuba, 26 m maximum depth 430 64Lakeba Nasaqalau Forereef scuba, 20 m maximum depth 480 64Lakeba Oru Forereef scuba, 20 m maximum depth 357 62Aiwa Levu Aiwa 1 Scuba, 100 m/500 m2 938 76Aiwa Levu Aiwa 2 Scuba, 100 m/500 m2 651 58Aiwa Lailai Aiwa LL1 Snorkel, 100 m/500 m2 703 60Aiwa Lailai Aiwa LL2 Snorkel, 100 m/500 m2 784 72Nayau Salia Snorkel, 100 m/500 m2 618 49Nayau Narocivo Snorkel, 100 m/500 m2 831 66Nayau Liku Snorkel, 100 m/500 m2 587 50


a small number of sea turtles. Large-bodiedfishes were relatively uncommon in the in-shore areas and common but not abundanton the forereef of all the study sites. Reefsharks and jacks are most frequently observedaround the inshore reefs of Aiwa. These ani-mals were also documented off Nayau’s northand south forereefs.

discussion

This research has produced important results,including descriptions of modern exploitationpatterns, extant marine biodiversity, and theoutcomes of inshore fishing expeditions. Inaddition, ethnoarchaeological studies haveilluminated several long-term trends in ma-rine resource exploitation and biodiversity.There appears to be relatively little variationin the types of marine vertebrate species ex-ploited over time. The vertebrates are rela-tively diverse and do not show a markeddecline in size according to the zooarchaeo-logical data. However, both the invertebratearchaeological assemblages and modern in-vertebrate communities provide evidence forshifts in exploitation and availability of ma-rine resources over time.

Archaeology

Laboratory analysis on archaeological boneand shell samples from Nayau and Aiwahas contributed to the understanding of themakeup and sizes of exploited marine taxa.Detailed analysis of over 24,000 fish bonesand shellfish remains indicates that theindigenous inhabitants of the study areas in-tensively exploited relatively small-bodiedinshore marine resources over the course oftheir 3,000-year history. However, to date,there is no solid zooarchaeological evidencesuggesting a major decrease in the overallsizes of exploited fishes and invertebratesover time. Rather, the data from fish vertebralmeasurements suggest an increase in the sizesand weights of fishes exploited on Nayaufrom the early to mid-late time periods, andrelatively little change on Aiwa Levu andAiwa Lailai. On Nayau this bimodal distribu-tion pattern in the Lapita-period site of Na

Masimasi may be indicative of two distinctfishing technologies utilized contemporane-ously, such as trolling (resulting in the cap-ture of large-bodied fishes) and inshore netfishing (resulting in the small-bodied fishes).

The most obvious differences are in thetaxa exploited and changes in the compositionof the assemblages through time. There islittle temporal variation in the overall num-bers of fish taxa exploited, regardless of theNISP and calculated MNI. By comparingfish assemblages with measures of diversityand equitability, it becomes apparent that thetaxa exploited remained relatively even overtime, and diversity ðHÞ was moderate tohigh. More variation is evident in the inverte-brate assemblages, but the shifts are minorwith the exception of the early exploitationof fighting conch (Strombus gibberulus) onNayau at Na Masimasi. On Aiwa Levu andAiwa Lailai, the invertebrate assemblagesare consistently dominated by turban snails(Turbo spp.). The invertebrate pattern maybe attributed to preference; that is, the firstinhabitants of the islands selected choice in-vertebrates based on taste and/or ease of ac-cess to these items. For example, the inshorearea by Na Masimasi is composed of coralreef on sand and limestone flats, exactly thehabitat preferred by fighting conch. It shouldbe noted that today both fighting conch andturban snails are relatively rare on Lakebaand slightly less rare on Nayau and the Aiwas(this is based on biological surveys and ethno-graphic data documented during fishing expe-ditions in the inshore area). Lauans claim thatthese taxa were more abundant 20–50 yearsago. The change in prehistoric and modernavailability could be due to a long history ofintensive exploitation, resulting in overexploi-tation, or a combination of environmentaland human-induced changes. However, inthe absence of data supporting a species-sizereduction in middens over time it is difficultto distinguish human impacts from the effectsof natural disturbance on invertebrate popu-lations or decreased shell bed density. Ar-chaeologists have identified sites exhibitingreductions in shellfish abundance that arelikely the result of terrestrial/agricultural de-velopment, including increased siltation into


the marine environment (Kirch and Yen1982, Spennemann 1987). Unfortunately, inmost situations there is no simple explanationfor the presence or absence of a marine spe-cies in a given habitat because this dependsheavily on natural stochastic recruiting events(Sale 1980).

habitat. Between 80% and 95% of theidentified fishes from all the study sites in-habit inshore coral reef environments ( bymeasures of NISP and MNI). The vast ma-jority of the identified invertebrates inhabitthe intertidal reef, the splash zone abovethe high-tide line, tide pools, sand flats, andfringing reefs. Turban snails (Turbinidae),the most common invertebrate at all the sites,with the exception of Na Masimasi wherefighting conchs are more frequent, inhabitcoral reef habitats and rocky coral areas.Turbo setosus, the most frequently identifiedTurbo on Aiwa, is known to prefer exposedareas of coral reef and the sublittoral zone inshallow waters (it accounts for 40–60% of theinvertebrate assemblages by measure of MNIand NISP). The identified bivalves can befound in shallow-water habitats, includingsilty or sandy inshore areas on fringing reefs(Kay 1979, Colin and Arneson 1995). Bivalvesmake up a small portion of all the zooarch-aeological assemblages (<20% at any site).

In his analysis of archaeological marine re-sources from Lakeba, Best (1984:498) arguedthat there is little evidence of any noteworthychanges in the fish taxa exploited over a3,000-year sequence of data. Best’s conclu-sions are supported by the archaeologicaldata in this study. One possible explanationfor this phenomenon is that given fairly smalland stable human population sizes over time,the Lauan data represent a relatively stablesystem of exploitation, where some of thetraditional fishing methods utilized by Lauansand the animals exploited (e.g., convict tangsand rabbitfishes that are targeted in vonoexpeditions have short population-doublingtimes) are suited to long-term sustainability(also see Jennings and Polunin 1996). Al-though this may be true for some bony fishes,the exploitation and availability of particularshellfish species appears to have decreasedthrough time.

A number of complex ecological processesmay be related to the documented changes infish and shellfish exploitation. These include,but are not limited to, variations in fishingtechnologies, shifting climatic conditions, re-cruitment, and changing predator-prey rela-tions. Undoubtedly the marine biota of Lauexperienced both natural and human-inducedshifts across space and time; these phenomenaare well documented throughout the Pacificislands (Allen 2006). Allen (2006) discussednew data from long-lived Pacific corals andgeneral climate modeling for the central Pa-cific. She described climatic shifts in the re-gion and areas of particular instability. Thedata suggest increased El Nino–Southern Os-cillation (ENSO) variability from A.D. 1100to 1400, with a shift from cool conditionsafter A.D. 1200 and a short warm period atabout A.D. 1300 (Allen 2006:530). In Fiji thetransition from the Little Climatic Optimum(LCO) (1250–700 B.P.) to the Little Ice Age(LIA) (700–200 B.P.) at around 700 B.P. wasa period of frequent and intense ENSOactivity, resulting in increased storminess,low-pressure systems, sea-level decline, andheightened sea temperatures (Nunn 2000,Hughes et al. 2003, Field 2004). Increasedtemperatures and ocean salinity are known toheavily impact coral reefs through bleachingand, ultimately, tremendous coral die-offs asseen in modern ENSO events (Hughes et al.2003, Allen 2006). The high level of declinein marine productivity due to these naturalclimatic events is extreme and may outweighthe potential negative impacts that relativelysmall populations such as those occupyingAiwa and Nayau at any given time couldhave exerted upon their rich marine environ-ments (Thomas makes a similar observationin his paper in this volume, regarding the po-tential marine impacts of small populationsinhabiting atolls in Kiribati. However, thisview and that of Allen stand in contrast towork by Pandolfi and colleagues [2003:957],who argue that their data trajectories of de-cline in fisheries worldwide through time aresimilar and that ‘‘All reefs were substantiallydegraded long before outbreaks of coral dis-ease and bleaching,’’ and therefore, humanoverfishing, along with associated land-based


pollution and runoff, is the only reasonableexplanation for the pre-1900 decline ofworldwide coral reefs); additional faunal datafrom sites dating to the LCO-LIA transi-tion and fine-grained radiocarbon dates areneeded to further illuminate this issue.

In D’Arcy’s (2006) recent book he dis-cussed potential impacts of highly unpredict-able annual and seasonal climatic variationson the lives of Pacific islanders inhabiting Re-mote Oceania. In terms of fishing patterns,biomass of fish taxa also changes annuallyand over longer-term periods, with variabilityvisible in terms of individuals. ‘‘The fact thatinstability in individual species’ numbers canoccur alongside stability in the overall size ofthe ecosystem biomass suggests that there canbe continuity in marine harvests for fisher-folk not rigidly tied to harvesting a fewspecies only’’ (D’Arcy 2006:21). In a highlyvariable environment flexibility in terms oftechnologies, habitats exploited, and targetedcatch appears to be a great advantage. In par-ticular, net technologies are well suited tocapture a broad spectrum of available inshorespecies. I argue that Lauan fisherpeople’sflexibility and natural shifts may both accountfor the zooarchaeologically observed patternsin the Lau marine fauna. In addition, Lauanshave long-standing elaborate systems of ma-rine tenure and rules regarding the use ofspecific areas of the reef. They consider evenminor shifts in marine variability and shift fo-cus accordingly in ways that manage use andtemporarily conserve particular resources.This may be achieved by declaring areas ofthe reef taboo or ‘‘off limits’’ for some timewhile local populations of marine resourcesare allowed to recover. Or a village elder maydecide that a particular type of fish should notbe captured over a given time period.

Ethnography

The inclusion of indigenous behaviors andknowledge is a unique and critical componentof this project. Interactions between scien-tists and local communities are undoubtedlycrucial to understanding and the long-termmaintenance and stewardship of marine bio-logical communities. Inshore fishing expedi-

tions are coordinated and directed by elderwomen, who pass knowledge of how andwhere to fish on to the younger generations.I worked closely with these fisherwomen inan attempt to learn what marine resourcesare harvested and how the diversity of avail-able fishes and invertebrates has changed. Ar-chaeological fish bones indicate that Lau’sfirst inhabitants intensively harvested rela-tively small inshore fishes, just as their de-scendants do today. There appears to be adirect correlation between the fish taxa thatLauans claim to prefer and those that areactually collected and consumed. The evolu-tionary perspective afforded by combiningmultiple perspectives has illuminated long-term trends in Lauan exploitation and prefer-ences. It is possible that some of the Lauantraditional systems for marine exploitationand management allow for long-term sustain-ability (especially in the case of small-bodiedinshore bony fishes), but others, such as theharvesting of shellfish, do not. Pandolfi et al.(2005) suggested that indicators of a healthyreef and successful management might in-clude the presence of taxa such as parrot-fishes, grazing sea urchins, sharks, turtles,large jacks, and groupers. These taxa arefound on all the study reefs but infrequentlyon Lakeba and more frequently on Nayauand the Aiwas.

Biological Surveys

The results of the marine biological surveysin association with this project build on previ-ous marine biological surveys from the Lauregion by Wells (1977), Jennings and Polunin(1995, 1996), Vuki et al. (2000), Dulvy et al.(2004), Kuster et al. (2005, 2006), and Turneret al. (2007). These workers and others havenoted that traditionally managed fisheries inFiji have expanded to accommodate escalat-ing demands for fish in emerging marketeconomies ( Jennings and Polunin 1995,1996), and that indigenous or informalknowledge is extremely useful for marine bi-ologists and ecologists in Fiji and the Pacificislands in general (Dulvy and Polunin 2004,Kuster et al. 2006, Middlebrook and Wil-liamson 2006).


On each of the study islands the biologicalsurveys revealed that there are patches ofhealthy, recovering reef associated with rela-tively large populations of diverse fish com-munities. However, the overall diversity andthe numbers of fishes recorded varied de-pending on the island (Table 7). The dataindicate that Aiwa and Nayau have morediverse and possibly more abundant marinecommunities. For example, the Lakeba sur-veys found relatively fewer total fishes andfewer fish species in larger survey areas(1,000 m2) than the surveys on Nayau andAiwa (500 m2). The Aiwa reefs have somehealthy colonies of soft tree corals and fingersoft corals, which are lacking on Lakeba andin most areas on Nayau. Reef sharks andlarge-bodied fishes are also more abundantaround Aiwa than the larger islands.

conclusions

Using the past as a baseline for comparisonwith the present to explore change, the inter-view and fishing expedition data provide im-portant information about the status of thecoral reefs; methods and technologies of ex-ploitation; changes in this environment; andpresence, absence, or shifts in the local avail-ability of particular resources. A wide rangeof species is regularly exploited, that is, a totalof 112 taxa for all four islands. The biologicalsurveys recorded over 200 species of fishes,and the archaeological bone identificationsinclude over 50 fish taxa. Although there aremany factors that influence the outcome ofthe summary data for each line of evidence,these measures do indicate an overlap in theavailable resources past and present, and con-tinuity in the way that the marine fauna havebeen used throughout Lauan history. In thefollowing section I discuss four key findingsbased on the data derived from the fieldworkand laboratory analyses. More research isneeded to adequately address the complexissues raised, but I begin with the followingobservations and interpretations.

First, the data from central Lau suggestthat the relative intensity of human exploita-tion (intensity is related to population size,land area and use, and the degree of agricul-

tural development) will determine the currentcomposition and biological diversity of ma-rine communities. It appears that biodiversitydoes not vary as a function of physical factorsalone. Although Lakeba should be the mostdiverse of the islands surveyed, due to thelarge size and the natural physical variationin the types of reefs, substrates, and landforms, Aiwa and Nayau appear to have morediversity in modern marine communities (ac-cording to catch diversity in Appendix andthat observed in marine surveys [Table 7]).Lakeba has the highest overall human popu-lation for the study sites, and it likely hassince the island was settled. However, byland area/m2 and reef area Lakeba is cur-rently less densely settled than Nayau. Adetailed comparison of the faunal materialfrom Lakeba, presented in Best (1984) (thecomparison is hampered by the fact thatBest’s collection and identification methodsdiffer from my own in fundamental ways.For example, Best did not use fine screens tocollect faunal samples, shellfish was not col-lected using the methods he used for verte-brates, and he did not identify fishes usingthe same elements that I use, rather he reliedonly on so-called ‘‘special elements’’ for iden-tification), and that from Nayau and Aiwa willdetermine if this pattern of diversity is appar-ent throughout the archaeological data. It isinteresting that the diversity of the zooarch-aeological samples of both bony fishes andshellfish is greater on Nayau than on Aiwa.Assuming that inshore catches represent across section of the available marine re-sources, this pattern may reflect greater di-versity on Nayau in prehistory and a shift incontemporary times to a less-diverse marineenvironment. Alternatively, the data could beindicative of a broader exploitation pattern bythe prehistoric occupants of Nayau versusthat on Aiwa.

Second, human disturbance may have oc-curred more extensively and continuously onlarge islands, and therefore small islands mayharbor species-rich communities. This find-ing is evidenced on the islands of Aiwa Levuand Aiwa Lailai and relates to the pointalready made. Lakeba has a larger humanpopulation, more advanced technologies and


infrastructures (roads, cars, educational andmedical compounds, an airport, a large pinetree farm, generators), and more develop-ment and runoff from agriculture than theother islands studied. Nayau, Aiwa Levu, andAiwa Lailai are much less disturbed in mod-ern times, but even in prehistory theseenvironments appear to have had less im-pacts resulting from human occupation. It islikely that these smaller islands had contin-ually smaller population densities and less-intensive exploitation over time (note thatthe size of the reef on Nayau is approxi-mately the same size as the reef associatedwith Aiwa Levu and Aiwa Lailai). The diver-sity of fishes and invertebrates species ex-ploited on Nayau and Aiwa is greater acrossprehistory; the reefs of these islands currentlyhave more diverse biological communitiesthan those of Lakeba.

Third, humans appear to have a selectiveeffect on marine biodiversity as particularspecies are/were targeted according to localstandards of ranking and preference. For ex-ample, people sometimes target large taxa,but they also target fishes with specific physi-cal characteristics such as big eyes and redlips; this includes, for example, emperorfishes(O’Day 2004). It should be noted that optimi-zation criteria derived from foraging theorydo not entirely dictate cultural preference inthis setting. In addition to exploiting large-bodied fishes with spears and while trolling(trolling and fishing outside the reef is amale activity; women fish in the inshorearea), Lauan women target fishes in the in-shore area (which are primarily small-bodied)and are motivated by characteristics thatextend beyond the body size of particularanimals (Tables 3–5 and Figures 2–9). Thisfinding was apparent in the archaeologicaldata (small-bodied fishes occur in the highestfrequencies in all sites across time periods),and the small-bodied fighting conch was in-tensively exploited at Nayau’s earliest knownLapita occupation site, Na Masimasi. Themajority of the animal protein consumedeach day by Lauans comes from the inshorearea; therefore small fishes contribute themost animal protein to the diet and havesince the earliest occupation of the islands

examined in this study, according to thezooarchaeological data. (This statementneeds to be evaluated by isotope analysis,which is forthcoming.) By determining localstandards of preference and rank and examin-ing social and environmental motivationsfor exploiting particular food items, we canbetter understand how humans make deci-sions about using their environment and itsresources.

Fourth, vulnerable species are currentlyoverexploited or locally extinct; even rareresources have withstood 3,000 years of hu-man impacts and thus may have life historytraits supporting resilience, making conserva-tion efforts worthwhile ( Jackson et al. 2001,Steadman 2006). In all of the archaeologicalsites, fish bones constitute the majority ofthe recovered fauna, suggesting a heavy reli-ance on the sea and relatively small inshorereef fish in particular. The bony fish familiesthat contribute the most to zooarchaeologicalassemblages throughout the archaeologicalsequence by count (NISP) and the numbersof individuals (MNI) are Acanthuridae(tangs), Serranidae (groupers), Scaridae (par-rotfishes), and Lethrinidae (emperorfishes).Modern subsistence systems also rely heavilyon these families of fishes, and they form amajority of the modern fauna, according tothe biological surveys. Invertebrates make upa less-important portion of the diet. Themost commonly exploited archaeological in-vertebrates include species in the family Tur-binidae (turban snails), which are now rare onLakeba and Aiwa and becoming increasinglyrare on Nayau; Strombidae (strombs); Coni-dae (cones); and Patellidae (limpets). Mystudy did not note any locally extinct re-sources (species that occurred in the past butwere absent in the modern community over-all), but according to ethnographic and bio-logical survey data, many large-bodied fishesand invertebrates appear to be in decline afterthree millennia of exploitation (groupers,parrotfishes, sweetlips, snappers, mullet, co-conut crab, giant clam, strombids, trumpettriton). These species and those that areknown to be locally productive on Lakeba,Nayau, and Aiwa would be excellent targetsfor conservation (see Table 6). For example,


in the study area the local diversity of lethri-nids, despite 3,000 years of intensive exploita-tion, is remarkable. There are 14 species oflethrinids documented in all of the FijiIslands (World Wildlife Foundation 2003),and my research documented 10 speciesfrom ethnographic observations, six from thearchaeological sites, and four in the modernmarine surveys.

Convict tangs (Acanthurus triostegus) andrabbitfishes (Siganus spinous) are well suitedto the types of exploitation practiced in thenorthern villages of Lakeba and on Nayaubecause they are less vulnerable than otherspecies—their population doubling times are1.4–4.4 years and less than 15 months, re-spectively (Froese and Pauly 2006). Appar-ently, a component of the Lauan traditionalsystem for marine management, movingfrom various fishing grounds exploited bygroup fishing practices, allows for long-termsustainability of particular types of resources.These ecological practices and traditionalecological knowledge (TEK) may be used toguide the development of programs for sus-tainable use of marine resources in Fiji andelsewhere in the tropics. In fact, social atti-tudes toward marine resource managementare critical to the success of managementand conservation programs (Middlebrookand Williamson 2006, Turner et al. 2007).

Thomas Lovejoy, who coined the termbiodiversity, claims that this is the singlemost important measure for evaluating theimpacts of humans on the environment(Lovejoy 1997:10). To understand the extentof the human impact on marine ecosystemsin Lau and the complicated relationships sur-rounding this issue, long-term research isneeded. It is clear that modern surveys andinterviews, as well as ethnographic documen-tation of fishing practices are all useful modesof determining the local reliance on and im-portance of marine resources. Each line ofevidence contributes varied but overlappinginformation, which should be compared withthe archaeological data.

Allen (2006) predicted that marine pro-ductivity declines due to heightened ENSOactivity during the LCO-LIA shift around700 B.P. had an obvious signature on marine

zooarchaeological assemblages; that is, theseclimatic changes resulted in environmentallyinduced cultural change. On Viti Levu Field(2004) documented shifts to fortified settle-ments around A.D. 1300–1500 that wereaccompanied by a broadening diet breadth.The mid-late period archaeological datafrom Nayau and Aiwa exhibit minor shifts inthe diversity of exploited taxa. A slightlygreater diversity in bony fish exploitation oc-curred during the mid-late period occupa-tions on Nayau, where a more diverse groupof invertebrates was exploited over time.However, there is no evidence for intensivedeclines in marine productivity from theLauan archaeological data. More obvious de-clines in invertebrates and large-bodied fishesappear in the modern data, rather than in thearchaeological past. Perhaps the data illus-trate both Lauan stability and flexibility inmarine exploitation patterns, just as D’Arcy(2006:21) suggested for all the inhabitants ofRemote Oceania, who have been subject tohighly variable climatic shifts throughouttheir history.

The responsible management and conser-vation of marine ecosystems is critical for Fi-jians. In the remote Lauan archipelago, theentire population lives on the coast and reliesheavily on marine resources for food andtheir livelihoods. Like coastal ecosystemsworldwide, Fiji’s marine biodiversity facesthe growing threat of overfishing and impactsresulting from pollution, land development,climatic change, and coral bleaching. Com-pared with other areas of Fiji, the Lau Groupis less well studied but exhibits great marinediversity as the result of smaller human pop-ulations, less development, partial isolation,little or no commercial fisheries, and lack ofa tourist infrastructure. My multidisciplinaryapproach contributes to the understandingof Lauan biodiversity through three perspec-tives, archaeological, ethnographic, and bio-logical. Together this information was usedto better understand the marine environmentand human interactions with it over the threemillennia of human occupation on the studyislands. Ultimately, I expect that an evolu-tionary perspective will facilitate the devel-opment of programs for sustainable use of


marine resources in the study area and be-yond.

Methodologically, this study indicates thateach level of data (ethnographic, archaeologi-cal, and marine biological) produces differentbut overlapping results. A better understand-ing of environmental problems and solutionsfor dealing with them will come from multi-disciplinary collaborations and the examina-tion of biological complexity over the longterm (Lovejoy 1997, Jackson et al. 2001,Briggs et al. 2006). Combining multiple ap-proaches and methodologies will enhanceunderstanding of the issues related to marinechanges on local and global scales (Kronenand Bender 2007, Turner et al. 2007, Rickand Earlandson 2008). A multidisciplinaryhistorical ecological approach holds muchpromise for the future of research in Fiji andin marine ecosystems worldwide.

acknowledgments

I gratefully acknowledge the late Na GoneTuraga Na Tui Lau, Tui Nayau Ka Sau niVanua ko Lau, The Right Honourable RatuSir Kamisese Mara and his family for allow-ing me to work on Lakeba, Nayau, andAiwa. I thank the Fiji Museum and especiallySepeti Matararaba for their important assis-tance. I am grateful to the people of Nayauand Lakeba for welcoming and facilitatingmy research, in addition to providing mewith a wealth of local knowledge. Vinakavakalevu Tui Liku, Tui Naro, Jack, Sera,Caka, Meli, Mote, Sina, Semeti Guvaki,Colati, and Rusila. The majority of the radio-carbon determinations (excluding those fromNa Masimasi) were funded by NationalScience Foundation awards to David Stead-man. I also thank Scott Fitzpatrick, LorettaCormier, and three anonymous reviewersfor constructive comments on this paper;Heather Walsh-Haney of Florida Gulf CoastUniversity for identifying the human remainssubmitted for radiocarbon dating; JosephOrtega for assistance with the illustrations;and Lisa Kirkendale and Peter Middlefartfor conducting marine biological surveys inAugust 2008.

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Appendix

Fish Species Collected in Modern Fishing Expeditions on the Islands of Aiwa Levu, Aiwa Lailai, Nayau, and Lakeba

Taxa: Scientific Name Common NameAiwaLevu

AiwaLailai Lakeba Nayau

Carcharhinidae Requiem sharksCarcharhinus spp. Reef sharks La L LGaleocerdo cuvier Tiger shark L L

Dasyatidae StingraysDasyatis kuhlii Bluespot stingray G

Muraenidae Moray eelsGymnomuraena zebra Zebra moray G*Gymnothorax favagineus Blackblotched moray S, GG. meleagris Guineafowl moray G*G. zonipectus Bartail moray S, GEnchelynassa canina Longfang moray S, GEnchelycore schismatorhynchus Whitemargined moray G*Echidna nebulosa Snowflake moray G S, G

Ophichthidae Snake eelsMyrichthys colubrinus Harlequin snake eel G* G*

Elopidae LadyfishesElops sp. Ladyfish G

Plotosidae Eel catfishesEuristhmus lepturus Longtailed catfish G


Appendix (continued)



Clupeidae HerringsHerklotsichthys quadrimaculatus Bluestripe herring G

Batrachoididae ToadfishesChelonodon patoca Mikspotted toadfish S, G

Exocoetidae FlyingfishesCypselurus sp. Flyingfish N, T

Belonidae NeedlefishesPlatybelone argalus platyura Keeled needlefish G GStrongylura incisa Reef needlefish GTylosurus crocodilus crocodilus Hound needlefish G

Hemiramphidae HalfbeaksHyporhamphus acutus acutus Halfbeak N

Holocentridae Squirrelfishes, soldierfishesMyripristis sp. Soldierfish S S, G S, G S, GSargocentron spiniferum Longjawed squirrelfish S G S S, GS. violaceum Violet squirrelfish S, G

Platycephalidae FlatheadsOnigocia spinosa Spiny flathead S, G

Scorpaenidae Scorpionfishes G*Serranidae GroupersCephalopholis argus Peacock grouper (rockcod) S S G, TC. sonnerati Tomato hind G, TEpinephelus merra Honeycomb grouper S, G, H S, G S, G GE. lanceolatus Giant grouper S, TE. howlandi Blacksaddle grouper GE. polyphekadion Marbled grouper S, G G, TE. tauvina Reef cod S

Therapontidae GruntersTerapon jarbua Crescentbanded grunter G G

Carangidae Jacks, trevallysAlectis ciliaris Pennantfish S, G, TCaranx ignobilis Giant trevally S, G, TC. melampygus Bluefin trevally H, T S S, G S, G, T

Gerreidae MojarrasGerres oyena Common silver-biddy G

Lutjanidae SnappersLutjanus spp. Snappers S, H, TLutjanus gibbus Paddletail S, H, T

Haemulidae Sweetlips, gruntsPlectorhinchus spp. Sweetlips S, H, T S, H, T

Nemipteridae Threadfin breamScolopsis bilineata Twolined monocle bream S G

Coryphaenidae DolphinfishesCoryphaena hippurus Common dolphinfish T T

Lethrinidae EmperorfishesGnathodentex aureolineatus Yellowspot emperor G GGymnocranius grandoculis Bluelined largeeye bream S S, GLethrinus conchyliatus Redaxil emperor GL. erythropterus Orangefin emperor S, G GL. harak Blackspot emperor G GL. miniatus Trumpet emperor GL. obsoletus Yellowstripe emperor GL. xanthochilus Yellowlip emperor G GL. atkinsoni Yellowbrown emperor GMonotaxis grandoculis Bigeye bream S, G G G





Mullidae GoatfishesMulloidichthys flavolineatus Yellowstripe goatfish G GMulloides vanicolensis Yellowfin goatfish G GParupeneus barberinoides Half and half goatfish G G GP. cyclostomus Yellowsaddle or Yellow goatfish GP. indicus Indian goatfish GP. macronemus Longbarbel goatfish GP. multifasciatus Multibarred goatfish S GP. trifasciatus Goatfish SP. pleurostigma Sidespot goatfish G G

Kyphosidae Sea chubsKyphosus bigibbus Gray sea chub S, G

Ephippidae Batfishes, spadefishesPlatax teira Longfin spadefish G

Chaetodontidae Butterflyfishes S G G GChaetodon vagabundus Vagabond butterflyfish G G

Pomacanthidae AngelfishesPygoplites diacanthus Royal angelfish S G

Pomacentridae DamselfishesAbudefduf septemfasciatus Banded sergeant S S

Labridae WrassesAnampses spp. Wrasses GBodianus vulpinus Blackspot pigfish S, GCheilinus undulatus Doubleheaded Maori wrasse S, GHologymnosus doliatus Pastel ringwrasse GLabrichthys unilineatus Tubelip wrasse GNovaculichthys taeniurus Carpet wrasse S, GThalassoma spp. Wrasses G GIniistius pavo Peacock wrasse S, G

Scaridae ParrotfishesCalotomus carolinus Carolines parrotfish GChlorurus microrhinos Steephead parrotfish S, GC. sordidus Bullethead parrotfish S S, GScarus globiceps Globehead parrotfish SS. rubroviolaceus Redlip parrotfish S GS. prasiognathos Dusky parrotfish S, GS. schlegeli Yellowband parrotfish GS. oviceps Blue parrotfish S, G G S, G

Mugilidae MulletsMugil cephalus Flathead mullet G GValamugil buchanani Bluetail mullet G GLiza vaigiensis Squaretail mullet G

Sphyraenidae BarracudasSphyraena barracuda Great barracuda G S, H, TSphyraena obtusata Striped seapike S, H, T

Blenniidae BlenniesSalaris sinuosus Fringelip blenny G

Acanthuridae SurgeonfishesAcanthurus lineatus Blueband surgeonfish S GA. guttatus Whitespotted surgeonfish G GA. nigricauda Epaulette surgeonfish SA. triostegus triostegus Convict tang G, S G S, G GCtenochaetus striatus Striped bristletooth S G S S, GNaso lituratus Orangespine unicornfish G, S S, GN. unicornis Bluespine unicornfish S G, S S, G





Zanclidae Moorish idolZanclus cornutus Moorish idol G G

Siganidae RabbitfishesSiganus argenteus Forktail rabbitfish G GS. spinus Little spinefoot rabbitfish S G G

Scombridae Tunas, mackerels, bonitoScomberomorus spp. Mackerel T

Bothidae and Soleidae Flounder and sole S, GBalistidae Triggerfishes S, G, HBalistoides viridescens Bluefinned triggerfish SPseudobalistes fuscus Yellowspot triggerfish GMelichthys niger Ebony triggerfish S, G

Ostraciidae TrunkfishesOstracion cubicus Yellow boxfish G* G*Lactoria diaphana Roundbelly cowfish G* G*

Tetraodontidae PuffersArothron reticularis Reticulated pufferfish GArothron stellatus Starry pufferfish G S, GTriodon macropterus Threetoothed puffer S, G

Diodontidae PorcupinefishesChilomycterus reticulatus Porcupinefish S, GDiodon hystrix Porcupinefish G S, G

Istiophoridae Billfish, marlin T

Note: Only seven species are bycatch, which are not consumed; these are marked with an asterisk (*).a Modes of collection: L, longline; G, gill net; S, spear; N, hand net; T, trolling; H, hand line.


a long-term perspective on biodiversity and marine ......617 a long-term perspective on biodiversity...

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