Revi of Trl nid Eco ogy and Some Possible Implications

for Archaeological Research

Received 20 1985

BARBARA G. MOIR

INTRODUCTION

THE MANUFACTURE AND USE of Tridacna adzes have long been associated with cul­tures inhabiting atolls and raised coral islands in the Pacific region. A survey of the archaeological and ethnographic literature reveals that Tridacna adze blades have been on numerous islands as establishing broad range island b:;bltats for the of these

the study adzes by archaeologists has almost exclu-sively on typological classification and culture-hIstorical relationships (Kennedy 1931; Thompson 1932; Spoehr 1957; Osborne 1966; Rosendahl 1969; Davidson 1971; Garanger 1972; Craib 1977; Ward 1979; Takayama and Intoh 1980; Kirch and Yen 1982; Sinoto 1984), with primary analytic emphasis given to the form and style of blade. The mateml~ (Tridacna spp. rarely, Hippoplls hippopus) from which have received al attention. to consider their morphological and the relevance of these to tool and efficiency, The Tridacnidae as components of reef ecosystems, together with their microenvi­ronmental requirements for life, suggest a different focus for investigation that may have important implications for archaeological research,

article surveys ecology of two (Tridacna) gigas Tridacna maxima often iden-

tifIed Tridacna adze Jssemblages-and the recorded distribution of Tri-dacna adze blades throughout the Pacific from an ecological perspective. It offers five propositions that might be of use to archaeologists in the development of an ecologi­cally based model of Tridacna adze distribution. Finally, it underscores the signif-icance of;J multidisciplinary approach to the of material and suggests that as well cultural variables affect the form geographical

assemblages.

Moir is alIili,llcd the Anthropology Department of the of Hawaii· Malloa

),1

98 Perspectives, 986-1987

tions are believed to maintain themselves through "recruitment" of juveniles that develop from the gametes of local adults (Yamaguchi 1977)-i.e., the majority of

tcnd to settle metamorphose vIcinity of their and short-term scientific data demonstrate considerable variability in the

of T. gigas. specimens annual increases· shell length of 51 mm (Rosewater 1965), from 50-80 mm (Bonham 1965), from 80-120 mm (Beckvar 1981), and means of 55 mm (J. Munro, pers. comm.) and 142 mm (Ham­ner n.d.) during the respective periods of investigation. There is also evidence for marked variation in tridacnid growth rates relative to latitude (Munro and Gwyther 1981; and Pearson Munro n.d.) habitat-e.g.,

gigas ft)und on and in more turbid sites near areas human activity even arnong rncmbers ofthc same apparent cohort under uniform environmental conditions (McKoy 1980; Beckvar 1981; Munro and Heslinga 1983).

McMichael (1974) measured the growth of different size classes of T. maxima under natural conditions; annual mean increment ranged mm (for the

to 38 rn (srnaUest clams). tridacnids, increment appears be a function size of the first measurement; the rate growth seems to be most rapid during the early years of life, slowmg as the clam grows larger (McMichael 1974; Yamaguchi 1977; Beckvar 1981; Romanek et al. 1987; Pearson and Munro n.d.). Specimens of T. maxima have achieved shell lengths of 35 em, while T. gigas hds been known to re:leb more than 1 m in length (Rose­water but in extant populations of such appear to be rare.

Nourishment

The Tridacnidae, like most bivalves, are filter feeders of phytoplankton, and they absorb dissolved inorgamc nutrients from (Fankboner, Coreau et aL

the gre;ltcst metabolic benefit, is obtained throllgh a highly method of nourishment that is 11l1lque among (Rosewater

1965). In addition to extracting nutrients from phytoplankton and utilizing dis­solved inorganic substances found in seawater, tridacnids also harbor and benefit from vast numbers of zooxanthellae-unicellular algae residing in the haemal

the clam's siphon J 936; Muscatine 1967; Trench et ; 1;lsher et aL and to a lesser extent in the stomach, and

tract (Goreau 1973; Trench 1981). Zooxanthellae are also found m the hermatypic corals. The highly developed association of these algae and the Tridacnidae is reflected in the clams' morphology, habits, and the environmental conditions under which they occur (Rosewater 1965; Goreau et al. 1973; Yonge 1974).

uptake of zooxanthellae from environment occurs ll1

stage of clarn development (Fit! Trench 1981). C::olcHlies of these infest the inner folds of mantle. When by the tide,

these folds extend laterally from the gaping valves and are maximally exposed to sunlight, permitting photosynthesis to take place within the znoxanthellae. Tridac­nids may receive some food energy via digestion of the algae, but the primary meta-bolic believed from released photosynthates et al. 1973;

Trench et specifically and alanine 1967).

100 Perspectives, 1986-1987

For marine fauna in general, temperature ranges favorable to reproduction and juvenile development are far more narrow than those tolerated by adult individuals

a1. 1954). providing a condition spawning, how-temperature important determining ;It which the tridacnid anilnal precIpItate carbonate seawater for the

formation of its shell (Sverdrup et a1. 1954; Romanek et a!. 1987).

SEAWATER CLARITY

The clear water habitat of tridacnids facilitates the transmission of light to their zooxanthcllar symbionts. and Hardy found thJt limiting depth (10~20 for the growth reefs was deterrnincd in large part by water's clarity, the insufficiency light for at depths greater than this (see also Brown and Muskanofola 1985). Their study of Tridacna ecology in Belau revealed the following microhabitat distributions for T. gigas and T. maxima: both were very seldom found in the bays, occasionally in the lagoons, and often in the w;lters of the and barrier T'he primary

difference these microhabltats is that of clarity, with providing condition 1940).

On Fanning Atoll, Kay and Switzer (1974) found the lagoon basIn effectively divided into a clear-water area near the deep reef pass (transparency to 10-15 m), and turbid-water sectors elsewhere (transparency less than 2 m). Distribution of T. max­ima (the only tridacnid species present) was confined to the dear lagoon, where massive cOfJls (Porites permitted settlement above carbonate mud sedirncnts. However, Tridacna found in exposed sections fossil reefs. occurrence in presently devoid livmg Tridacna suggests changes in lagoon character associated with the closure of former reef passes that would have affected water clarity, coral growth, and/or sea level.

Renaud-Mornant et al. (1971) demonstrated the confinement of T. maxima dis-tribution an atoll in Tuarnotus to the shallow, transparent depths of that atoll's Their data underscore the of seawater to tridacnid survivaL Moreover, studies the geomorphological effects of storms on coral reefs suggest that increased turbidity of reef waters following erosion may delay recovery of hermatypic corals (and their faunal associations) for as many as 30 years (Stoddart 1971).

SEA WATER SALINITY

are found with salinity of near 35 mInImUm salinity level tolerated by tridaCiuds is not known, but it may be related to that of the hermatypic corals with which they are associated; tropical corals have been reported to withstand a 20 percent reduction in the salinity of their environment (Sverdrup et a1. 1954). Tridacna gigas has been successfully maintained in raceways at a level of 32

1981; 1984) and sites, together T. maxima, at ppt (Munro Gwyther 1981)

opposite end spectrum, at species of known to tolerate higher saline levels than those normally found in the open ocean. Tridacna maxima occurs in enormous numbers in the closed lagoons ofReao, Maturei Vavao, and Takapoto atolls in the Tuamotus, which have salinity levels of 37.5, 37.5, and 43 respectively 1970; Renaud-Mornant et a1. 1971; J. Newhouse, pers,

102 xxvu(1), 1986-1987

Tridacna populations. normal maintenance of through local recruitment adults, it would be reasonable that intensive might well result in decimation extinction of a species (Wells et a1. 1983). Should this happen, it would be unlikely to recover without reintroduction (Yamaguchi 1977; Hirschberger 1980; Hamner n.d.). Studies by Hardy and Hardy (1969) and by Bryan and McConnel (1976) indi­cate a substantial decline in population densities for larger tridacnid species on the reefs of Belau; this decline is viewed as the direct result of overfishing for the export of shell (see also Hester and Jones 1974). In recent years T. gigas has become extinct in the waters offPohnpei, Truk, and Kosrae, presumably as a consequence of human exploitation (Heslinga Tal\vanese fishing vessels, in known to poach the adductor muscle-abandoning maining flesh of the and vulnerable reefs in the (Pearson 1977; Wankowski and Munro 1981; Hamner n.d.a). In the the success of any Tridacna would have been density oflocal human settlernent extent and rate of rates of tridacnid reproduction, and growth do not keep pace with the demand for this resource, human behavior creates a marginal habitat for the Tridacnidae.

The conditions necessary for tridacnid life are also found to combine in ways that permit a high level of species richness, but where competition for resources may limit the number of individuals the habitat can support. These combinations may be termed competitive habitats. Open atolls (with three or more deep reef passes) in par­ticular, and those fringing and barrier reef areas unaffected by freshwater runoff and siltation, are examples of hahitats, where the basic ments for tridacnid which are characterized by richness but lower for some species. The researches and Richard (Salvat 1971 b, 1971c, 1972, 1973; Richard 1971, 1972) have contrast in molluscan tion and individual atollslreefs and closed systems differ of open seawater and lagoon corresponding differences m clrculation, salinity, and turbidity. Open recf systems apparently offer conditions favorable to a variety of tridacnid species, but competi­tion for available resources may restrict the number of individuals of some species to comparatively low levels.

The findings of Richard and Salvat suggest a third combination of environmental variables that affect the ecological success of the Tridacnidae: the exceptional habitat. Here too are found all the basic requirements for tridacnid survival, but they com­bine with low faunal diversity and minimal competition for resources to comprise a habitat wherein the rnay be exceptionally successfllL example of this habitat IS concerned may be the ing deep reef passes), forms (with one or two appear to offer Radtke et a1. (n.d.) found reef of Rose Atoll support a single tridacnid ima, at a density of per m2 . Studies by Richard the benthic fauna Pukarua, Marutea, Pukapub, Maturei Vavao, Fangataufa, and Vahitahi atolls in the Tuamotus-all of them closed-revealed T. maxima (the only tridacnid present) to be dominant among all

104 xxvu(1),1986-1987

found together on atolls, T. maxima will be reef biotopes while only rarely and only on the These impressions propositions that relate variables (habitat) to ecological variables (success ofTridacnidae):

P1: Open and semi closed atolls (with one or more deep reef passes) will be character­ized by a greater diversity of Tridacnidae species than will closed atolls (lacking such passes), within the distributional overlap of any or all species.

P2: Within the distributional range of T. gigas, open and semiclosed atolls will be found to support viable populations of this species in both reef and lagoon biotopes. Populations of T. gigas will rarely occur on closed atolls within the range of this species. represented on a closed atoll within will be as the outer reef and not in the closed

Tridacna Technology A fundamental tndacnid ecology is essential to

made from this comparative analyses of tween island groups and across habitats. Within the distributional overlap of '1'. gigas and T. maxima, prehistoric human populations on open and semi closed atolls, or on islands with associated reefs, may have had ready access to more species of Tridacna for tool material than did communities living on closed atolls. If this were the case, one might anticipate the tool kits of the former to be representative of more Tridacna species than those of the latter. It might be argued that whereas open reef systems offered competitive habitats for both T. gigas and T. maxima, closed atolls constituted only marginal bhitats for T. gigas but competitive or even habitats for T. differences in the availability of might correspond extent to which they were material.

These speculations relate environmental

the form of the following material culture variables

P3: Within the T. gigas and T. maxima, Tridilwa manufactured and used by the prehistoric inhabitants of open and semi closed atolls, or of volcanic and raised coral islands with associated reefs, will in general include blades made from the valves of both species.

P4: Within the distributional overlap of T. gigas and T. maxima, Tridacna adze kits manufactured and used by the prehistoric inhabitants of closed atolls will in general be characterized only by blades made from the valves of T. maxima. The low frequency with which T. gigas populations occur in this environmental context suggests the possibility of external origins for locally recovered tools made from this materiaL

Artifactual and data (Moir n. d. b, 1989; BPBM lections) indicate gigas valve were utilized than was the case tridacnid valves. Valve features from T. gigas adze hinge line, individual radial line plus anterior umbonal region (Fig. 2). By ima blades were fashioned primarily from the valve body between the dorsal margin (lip) and ventral/hinge lines (Fig. 3)-an area frequently misidentified by

MOIR: TRU)ACNID ECOLOGY AND POSSIBLE IMPLICATIONS

o "0

l\RCHAE010C;ICAL RESEARCH

a: o 0:0 wZ I-W Z «

o a:0 wZ I-W

Q) 0lQ)

- c.:: .c.

o

106

ANTER 10 R E D

Perspectives, 1986-1987

Exterior and intetior of valves of (Chametrachea) maxima ([(c,ding). Valve section most often identified from T. maxima adze blades is outlined.

as the lip "-as were H. hippopU5 preforms re-Takuu h) and another Bikini (c. llnpublished

data). same region squamosa also have lIsed for blade material (Sinoto 1984), as well as the thicker UInbonal section (Moir n. d. b). In gener­al, owing to their proportionately greater mass, the umbonal and hinge line areas of a mature T. gigas valve would have yielded a more plastic tool material than the corresponding features of the valves of smaller species. Given this greater morpho-logical and the that more the T. gigas were used for tool than of other t.ridacnid valves, be argued inhabitants of open semiclosed atolls Islands with associated reefs 'T"ridacna adze kits characterized by a greater diversity of blade form than those produced on closed atolls.

This suggests a further environmental-material proposition:

the distributional overlap of T. gl.~ill T. maxima, adze kits manufactured and the prehistOriC .inhahitants of semiclosed

or of volcanic raised coral islands associated character-ized by a greater diversity of blade form than will those produced on closed atolls.

These last three propositions may easily be tested, initially through a careful ex­amination of the available literature and collections, and more rigorously through

investigation. The findings interpreted the perspec-

TABI.J~ J'FIlDACNA ADZES ON VOLCANIC ISLANDS

T.gigas

(Belau)l,z*.*'

Fefan3

Guam4

Kosrae5.6 .7

Lelepa8

Okinawa9

Palawan lO

Pohnpei 11.12.13.14.15

Retoka8

Sanga Sanga 10 Santa Cruz l6

Tikopia 17

To1'8,19

Tongoa8 (Truk)20 Yap7,21

Buka24

(Belau)I.2 Efate8,25

Erromanga22

Fila25

Futuna, West25

Guam4,26.27 Kosrae6,28

Lelepa8

Santa Cruz l6

Tikopia 17.36 To)l8.19

Tongoa8

Vanikoro37 Yap38

• Localities in parentheses indicate island was unspecified .

Buka4')

Efate8

Ishigaki41 ,42 New Britain8

New Ireland43,44

Niuatoputapu45 Pohnpei33,46 Rota34,47 Rotuma35,48,49

• * Data sources: 1, Osborne 1900; 2. Gllmerman et a!. 1981; 3, Sinoto 1984; 4, Hiro & Clayshulte 1981; 5, Bath & Shun 1982; 8, Garanger 1972; 9, O.P.B.E 11, Christian 1899; 12. 1980; 14, Athens 1984; 15, Rosendahl McCoy (unpub!. data); T"kayarn:l & Seki 1973; 19, Takayama & 1 1964; 21, D. Rubinstein Spriggs (unpub1. data); 23, Kirch & Wickler (unpubL data); j 966; 26, Hiro et al. 1982; 27, Hunter-Anderson Athens et aJ. 1983; 29, Clay 1974; 31, Thompson 1932; Ayres et a1. n.d.; 34, Gardiner 1898; 36, Kirch & Yen 38, Gifford & Gifford Specht 1969; 41, Kokubu & Kaneko 1969; 43, White 1972; Rogers 1974; 46, Sarfert 1913; 47. 48, Eason 1951; 49, Parke 1964.

TABLE 2. RECORDS OF TRIDACNA ADZES ON RAISED CORAL ISLANDS

1'. gigas

Banaba I' Fais2

Pakea3

Pur4

Sonsorol4

Tobi5

T. maxima

Banaba l

Bellona6

F~js2

UNIDENTIFIED Tridacna SP.

Fulanga l2

Malden t3

Namllb l2

• Data sources: 1, Lan'lwn (pers. comm.); 3, L Groube, 4, Eilers 1935; 5, Eilers 1936; 6, Poulsen J972; 7, Rosendahl 1987; 8, Krause 1906; 9, Ward 1979; 10, Sarfert & Damm 1935; II, Moir (unpub!. data); 12, Thompson 1938; 13. Dixon 1878; 14, M, Spriggs (unpuhl. data); 15, Loeb 1926; 16, Poulsen 1964.

TRIDACNID ECOLOGY AND POSSIBLE IMPLICATIONS fOR ARCHAEOLOGICAL RESEARCH

TABLE 3. RECORDS OF Tridawi2 ADZES ON OPEN, SE,MICLOSED, AND CLOSED ATOLLS

Open atoll tAilinglaplapl tAnd2

tAur 1

tBikilli3 tEIoaua-Emananus4

(Mussau Is,) tE~rJulapS

tFunafuti6

tJaluit7

tK wajalein8 tLamotrek 1.5.9 tLikiepl tMaloelap' tNgulu 'O

tNukununu"· 12 tNukuria 11 ,12

tOntongJava ll . 13 tSikai.lll:l 1:l,15 tTabltcuca 16

Tongareva 17

tUlithiI8.19.20

tWoleai 21

tWotho l

tWotje l

Semic/osed atoll tArno l •22

Atalu23

tEbon 1

tlfaluk1'()

tKa pi n gam arangi24.25 tLae' tMajuro1.26 tNgatik27

tNonomi l6

tNukufetau28

tNukuorolS.29.30

tOnotoa '6 tPuluwat31

ISatawan32 tTakuu"· 13.33

atoll Cbmtmas34 tEtaj32

Fang:1 Ll" fa ls

Manihiki l5.36

tMokil' tNanumea28

Napuka3S

tNiutao28

tPingelap' Pubpuka37

T gigas

x

X

X X

X X X

X

X

X

X

X X

X

T maxima

x X X X X

X

X

X

X X X X X X

X X

X

X X

X X X

UNlDENTIHED

Tridacna SP,

X

X

X

X

X

X X

X

X

X

X X

X

X

X

X

MOIR: TRJDACNID ECOLOGY AND POSSIBLE IMPLICAHONS FOR ARCHAEOLOCICAL RESEARCH 111

early evidence that such a trade link existed between Belau and Yap (Muller 1917)­T. ~i~as blades or valves were reportedly brought from Belau (where the species was

to Yap was not, except ancient valves in coral)-hypothesize similar situation obtained on Lamotrek. a volcanic with associated situated within distributional

extent of T. gigas, this species may be expected to have inhabited those reefs in the past. Subsequent local extinction of T. gigas may have been related to adverse en­vironmental conditions (e.g., sedimentation of reefs following one of the typhoons

the area) or overcxploitation humans. This may been the case as well, that an extant of T. have been

reduced rather completely ina ted. While and Alkire found no ltve T. gigas specimens on Lamotrek during their research in 1976, a "relict" population was reported there by marine biologists a decade later (Munro n.d.), specimens that probably had been overlooked by the archaeologists. This suggests that Lamotrek may well have been a viable habitat for T. }?~iZaS in the past; that population bivalves cou Id supplied the m~lterial for the adzc recovered and in the sequenee (perhaps more than environment~ll cultural were instrumental reducing the T. gigas population to an extremely low level. It is also possible, of course, that the large Tridacna adze blades recovered by Fujimura and Alkire-or the raw materials themselves-were obtained from external sources.

~amotrek's embedded valves could serve P2 for prehistoric Lamotrek, by to deter-mine Tridacna Vi/ere present on atoll in earlier Evidence for prehistoric local population of T. gigas would simplify the archaeologist's search for the source of this tool material. The possibility of external origins for T. gigas tools recovered from Lamotrek would not be discounted, but might be argued less con­vincingly on environmental-ecological grounds in the face of J local supply of raw material. The artifactual from Fujimura Alkire's (1977, excavations support the adze includes blades from marc one species of Tridama, apparently including T. gigas.

A closed atoll in the eastern Carolines would offer an equally interesting oppor­tunity for tests of P1, P2, and P4. Some 150 km distant from the volcanic island of Pohnpei, Mokil Atoll lacks deep reef passes that can be navigated by ship. Bryan (1971) recorded four "canoe passages" the north and reef sectors, one has been to accom small motorized craft. Mokil is located within the distributional range of but reportedly not toda y support a population of this species (A. Sepedi, pers. comm.). Rosendahl (1979) conducted a reconnaissance survey of Mokil, focusing on a traditional chiefly resi­dence site, and recovered a number of artifacts from the surface and from infor-mants, Among these Tridacna adze (illustrated Rosendahl 1987 and in Table appear to manufactured quite mas-sIve hinges, and from valves , gigas. The survey rep on (1979) does not specify whether these two blades were obtained through surface collection or from informants. In the latter case, information regarding the context and history of the artifacts' possession by informants would provide a more solid basis for establishing their cultural provenience.

survey Mokll's reef and biotopes, both speCImens

) ,

MOIR: TRIDACNID ECOLOCY

rials indigenous to assume a local source with every type of

L\;lPLlCATIONS FOR ARCHAEOLOGICM.

neither, perhaps, would recovered from cultural deposits

Procedures for tracing the sources of Tridacna materials to individual island groups, similar to those for sourcing volcanic glass and pottery tempers, have not been and may never be developed. Nonetheless, the formulation and testing of environmental-ecological correlates (such as Pl and P2) should contribute to our awareness of the ecology of these species. An improved understanding of the habits and habitats of the Tridacnidae would allow archaeologists to anticipate, within known geographical distributions, the presence or absence and relative abundance of these resources in' the concepts of marginal, and exceptional species to the conditions reef system.

Further, environmental-material corrclates (such as P3, P4, and P5) us to test and to build propositions through the and island ecosysterns excavation. New evidence, including in the biological or should stimulate further and help to modify this prelirninary model in useful ways.

A multidisciplinary approach of this kind suggests that it is not enough to examine the tools of a culture apart from the ecosystemic contexts in which they occur; the materials from which tools are made are as much components of an ecosystem as are the human beings who make them. The complex associations of the Tridacnidae with their natural habitats are directly related to their ecological success, and thus to their availability for human exploitation.

EDGMENTS

This paper has with many individuals, ful for their input. access to unpublished artifact collections. Melody Actouka, Tim Adams, Joyce Bath, Robert Gerald Heslinga, Jay Kay, Patrick Kirch, Munro, Richard Radtke. Donald Rubinstein, Ahm Sepedi, Matthew Spriggs, Charles Streck, and Steven Wickler for providing data, comments, and criticism and for tolerating persistent questions. Any errors of presentation or interpretation are solely my responsibility.

ATHENS, J. S. 1980 Archaeological

Monograph 1984 Jobs Bill

submitted to

submitted to

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