seasonally foresta panama

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Prehistoric Human Adaptations to the Seasonally Dry Forests of PanamaAuthor(s): Richard Cooke and Anthony J. RanereSource: World Archaeology, Vol. 24, No. 1, The Humid Tropics (Jun., 1992), pp. 114-133Published by: Taylor & Francis, Ltd.Stable URL: http://www.jstor.org/stable/124901

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rehistoric h u r n n d pt tions tthe seasonally dry forests ofPanamaRichard Cooke and Anthony Ranere

IntroductionThe debate about when and how humans colonized tropical forests has been rekindled by abeguilingly straightforward null hypothesis that proposes that this bioome cannot sustainhuman hunters and gatherers (Bailey et al. 1989). Currently, it can be countered moreeffectively with theoretical ecological and substantive paleoecological data than withinformation from the field archaeological record (Colinvaux and Bush 1991;Piperno et al.1991b: 224). This is attributable to two factors. Firstly, many tropical regions that are orwere forested have excavation and survey records that are deficient with respect totemporal continuity, spatial integrity and cultural diversity. -lence, it is difficult to identifywhen humans first appeared in tropical forests and whether occupation prior to theappearance of easily detectable farming settlements was intermittent or uninterrupted,dispersed or localized, the result of endogenous growth or influenced by migrations.Secondly, differentiating between remains of wild plants and early cultivars, taxingenough biologically (Rindos 1984), is complicated in tropical forests by poor preservationof all but the most indestructible of diagnostic carbonized materials. In other words,archaeological inferences about the history and nature of human occupation of tropicalforests (especially before 3000--2000 BP) remain, in many regions, subservient todefinitions, points of view and even guesses.Long-term multidisciplinary projects that focus upon particular regions are anappropriate framework for identifying the arrival of human groups in tropical forests andfor tracing their short- and long-term impacts upon them. In this paper, we refer to such aproject - the 'Proyecto Santa Maria' (henceforth, PSM) - located in a small watershed ofthe same name on the Pacific slopes of central Panama (Fig. 1). The PSM combinedrandomly selected, 0.5 km-wide transects, traversed at intervals of 25m, with purposivesurveys, mapping and test excavations in rockshelters, shellmounds and nucleatedsettlements in riverine areas. For field methodology see Weiland (1984) and Hansell(1988).We originally (1981) proposed eight 'multiple working hypotheses', which highlightedour major research interests and served to orientate fieldwork and data recovery (Cookeand Ranere 1984). They are relevant to arguments stimulated by Bailey et al.s nullhypothesis. We reevaluate them in the light of currently analysed data. Additional detail

World Archaeology Volume 24 No. 1 The Humid 'Tropicsco Routledge 1i992 0043-8243/92/2401/114 $3.00/1


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Prehistoric human adaptationsto the seasonally dryforests of Panama 115

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116 Richard Cooke and Anthony J. Ranerecan be found in the following references that incorporate data generated by the PSM orrefer specifically to the study watershed: Cedefno1985; Claryet al. 1984; Cooke 1984b, inpress; Cooke and Ranere 1989, 1992;Hansell 1987, 1988;Norr 1990;Piperno 1988, 1989, inpress; Piperno and Clary 1984;Piperno et al. 1985, 1990, 1991a, 1991b; Ranere in press a,b; Ranere and Cooke in press a, b and c; Valerio 1985, 1987. Since our evaluations areessentially syntheses of these sources we keep subsequent references to them to aminimum. For information on sites in and adjacent to the watershed that were excavatedprior to the PSM, whose materials are being reanalysed, see: Cooke 1979, 1984a;McGimsey 1956; McGimsey et al. 1986-87; Ranere and Hansell 1978; Willey andMcGimsey 1954. All 14C dates are uncalibrated. Some of those run on marine shell havebeen corrected for 13C/12C ractionation (marked * in Tables 2 and 4). The reservoir effectwas not calculated.

The study watershedBefore addressing the multiple working hypotheses in turn, we briefly describe the studywatershed (catchment = 3,315 km2). We append a vignette of the state of vegetation whenthe Spanish interrupted indigenous development between AD 1501 and 1522.The 145km-long Santa Maria river rises in the cordillera (marked 1 in Figure 1) atc. 1,000m elevation, 22km from the Caribbean Sea. Initially, it descends southwards as astream, joined by affluents, through broken hilly terrain-- the 'foothills' (2). It then turnseastwards over an eroded Tertiary substrate, the 'Santiago Plain' (3). After brieflymeandering through Holocene colluvial deposits, the 'old and new deltas' (4 and 5) itenters the Pacific at Parita Bay, a recent marine extension of Panama Bay.During the rainy season (on average, late April to early December) the river runsorange-brown with suspended sediments. In some years, flood waters overflow seawardsalong features of earlier stages in the delta's evolution - meander scars, elbows of capture,swamps and larger, probably fossil, main channels (i.e. the Estero Salado and Escota'rivers'). During dry months, flow diminishes considerably, exposing dykes and sand barsthat hinder upriver travel by watercraft.The delta's position and dynamics are conditioned by the complex interplay of sea levelchange, surface topography, sediment discharge, tidal energy and the effects of past andpresent human activities. For the last 3,000 years the delta has been prograding, in thecentral section c. lkm per 1,000 years (Barber 1981; Clary et al. 1984; Dere 1981). Todayits largely mangrove-fringed coastline lies seaward of a patchwork of estuarine channels,salt marshes (alvinas) and low promontories capped with xerophile vegetation. The hightidal range (max. c. 6m) is conducive to the formation of mudflats, havens for estuarinefish, intertidal macroinvertebrates and shorebirds. Largerrocky 'inselbergs' that protrudethrough the deltaic sediments offer well-drained locales for human settlements. Theseretain remnants of tropical dry forest (Murphy and Lugo 1986).Precipitation is influenced primarily by orography, and wind direction and velocity.Annual rainfall averages 4,000-3,000mm in the cordillera and foothills above 300m,depending upon local topography, and 1,800-1,000mm in the plains adjacent to ParitaBay. The lowest figures occur right on the coast between Guarard and Nata (Fig. 1).


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Prehistoric human adaptationsto theseasonally dry forests of Panama 117During the dry season the north-easterly trade winds that blow perpendicular to thecordillera become katabatic as they cross the Pacific slopes, so that their strength andconstancy have a profound desiccating effect on the landscape and vegetation. Below1,500-500m, dry seasons are arid (3.5-5 completely dry months), but above theseelevations they are mitigated by mists and light rain (bajareque). Rainy seasonprecipitation is highest in the foothills and cordillera north-west of a line between Chitraand Santiago. Along the coastal plain and at higher elevations towards the eastern marginof the watershed rainless periods of up to two months can occur, especially when thetropical convergence zone is weakly developed over the Isthmus.

Vegetation at first contact with the SpanishThe vegetation of the Santa Mariawatershed and others immediately adjacent to it at theend of the Precolumbian era is described grosso modo in eye-witness documents that referto the years between Columbus' settlement of Santa Mariade Bel6n on the Caribbean sea(AD 1502-3) and the establishment of Nata, the firstpermanent town in the central Pacificlowlands (AD 1522) (Fig. 1). Documentation for the following summary can be found in:Toribio-Medina 1913: 154-83,191-207, 272-304; and Columbus 1959.

(1) The Caribbean slopes north of the Santa Maria headwaters were heavily forested,probably, as today, with evergreen rain forest. Amerind settlements were small andscattered, but there was greater nucleation near auriferous streams.(2) Over most of the watershed itself, vegetation is described generally as clear land andsavanna (tierraclaray sabanas), without deep forests (montes), in some places treeless (sinarcabucos), but with woods (arboledas) along rivers and on hill tops. A landscape akin tothe sabanas arboladas of Guanacaste, Costa Rica, is a reasonable reconstruction(Elizondo and Jimenez 1988). The riverine woods had trees large enough to makesea-going war canoes. Specific sections of the watershed, identified in Figure 1, aredescribed as follows:

(a) between Nata and Guarare: 'all savanna, without forest except for woods on thebanks of rivers and on hill tops',(b) between the Santiago Plain and 'Esquegua' (probably modern Chitra, elevation500m): 'savannas and open land',(c) the political unit controlled by Natd: 'clear land and savannas and cultivated (orshaded) riverbanks [our translation for vegas]'. As Piperno et al. (1991b) suggest, LaYeguada may have been situated between the territories of Nata and Esquegua. Localvegetation immediately before the period of post-conquest forest recovery consisted ofanthropogenic wooded savannas.(3) Some large villages were encountered. Espinosa estimated Nata's population at> 1500in AD 1516-17. Nata is an extensive archaeological site (Cooke 1979), but existingfield data do not permit verification of Espinosa's figure. If it is an ambitious lieutenant's

hyperbole, it may be compensated by the negative demographic effects of Europeandiseases, which were surely felt here before AD 1516. It is consistent with the'repartimiento' figures for 3 November 1519 and May 1522 (Romoli 1987: table 2), and


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118 Richard Cooke and Anthony J. Ranerewith population estimates inferred from artifact distributions at older prehistoricsettlements in the region, i.e., La Mula-Sariguaand Sitio Sierra (Table 5).Stands of fruit trees, i.e. 'mameyes' (Sapotaceae), and defences constructed with thornybushes and spiny bromeliads, would have given these indigenous villages a more coppice-or orchard-like appearance than rural communities today.

(4) The consumption of ground and fermented maize was widespread in a region whichquickly became the granary for the Spanish administrative capital at Panama Viejo.Extensive maize cultivation is described for the lower courses of the rivers that flow intoParita Bay. Cassava (Manihot esculenta) and cucurbits ('melones') were also planted infields near the coast. Spanish references to fauna emphasize white-tailed deer (Odocoileusvirginianus), 'partridges' (undoubtedly quails, Colinus cristatus), and pigeons and doves(Columbidae). Deer meat was commonly preserved in storehouses. Vertebrate archaeo-faunas from sites located within 60km of ParitaBay, which date from 5000 to c. 500 BP, arenotable for the absence or extreme rarityof taxa that prefer continuous or old forest.

Evaluation of the multiple working hypotheses(1) A dry episode did (did not) mark the end of the Pleistocene, resultingin anopen or savanna setting(2) Paleoindian populations were (werenot) adapted to tropicalforest biomesEvidence from sediment cores raised from lake La Yeguada, at 650m elevation within thewatershed, and from the extinct caldera lake at El Valle, 90km to the east at 500melevation (Bush and Colinvaux 1990), suggests that late glacial (14,000-10,000 BP)vegetation at these altitudes consisted of montane forests dominated by oaks (Quercus),which today thrive 1,000m higher. A temperature depression of 40-60C is inferred.Lowered lake levels should correspond to drier conditions, particularly between16,000-11,000 BP, although exactly how much drierremains moot. Diatom analysis showsthat La Yeguada did not become a shallow-water environment. The area was not as aridoropen as the late glacial Peten (Guatemala) or Lake Valencia (Venezuela) (Bradburyet al.1981; Leyden 1984; 1985).Projections made on the basis of these data and the present-day distribution of life zonessuggest that the watershed below 500m would have been dominated by formations of lowtrees, shrubs and succulents, rather than by grassland savannas. Hence, while there is noevidence for late glacial savannas above 500m, the connection between dryness andopenness below this altitude cannot be demonstrated.PSM surveys did not locate archaeological materialswhich can be considered anterior tothe late glacial Paleoindian occupation. The oldest in situ 14C-dated occupation wasidentified at a small rockshelter (Corona): 10,440 ? 650 BP (Beta-19,105). It is associatedwith abifacial stone technology, but lacks diagnostic artifacts. At La Mula-West, a shallowlocus that was disturbed in antiquity, stone projectile points, scapers and gravers are sosimilar to those of the Clovis horizon, dated in North America to c. 11,200-10,900 BP(Haynes 1991), that their contemporaneity is likely. The first indications of burningsuggestive of constant human activity are detected at La Yeguada c. 11,050 BP.


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Prehistorichuman adaptationsto theseasonally dry forests of Panama 119We propose that Paleoindian (late glacial) populations in the Santa Maria watershedwere moving in and out of both premontane oak forests and lowland woody scrub. Wedoubt whether the Clovis-like and the probably descendant fish-tail stemmed points,

found elsewhere in Panama (Bird and Cooke 1978), were manufactured for activities otherthan hunting large terrestrial mammals. The hypothesis that some units of Paleoindiansocieties were engaged primarily in gathering wild plant foods in forests (Piperno et al.1991a; cf. Ranere 1980a) can only be confirmed with dated archaeobotanical macro-remains (Dillehay 1990).(3) Changingrates in sea-level rise and sedimentation and tectonicmovementsresulted (did not result) in an oscillating coastline duringthe HoloceneTextural analysis of Vibracore and surface sediments permits preliminary correlationsbetween the evolution of the Santa Mariadelta, sea-level change and the local topographyof important archaeological sites. More detailed paleogeographical reconstructions willrequire the application of additional field and laboratory technologies.The Parita Bay coastline since human arrival has, indeed, oscillated. Barber (1981)proposed a facies change model that relates Cerro Mangote (7000-5000 BP) to coastlinemovements (Table 1).La Mula-Sarigua, firstoccupied duringlate glacial and early Holocene times, became animportant village between 2390 ? 70 BP (Beta-12,931) and 1970 + 45 BP (SI-5689), whenits refuse covered 58ha (Table 5). Today the alvina that stretches c. 2km in front of the sitedries out in the windy months. Clouds of saline dust driven inland curtail human activities.Bare alvina surfaces did not form between the site and the coastline until c. 1600 BP, bywhich time prehistoric occupation had declined noticeably. Hence La Mula-Sarigua'sapogee as a regional centre occurred at a time when the site was adjacent to, or evensurrounded by, mangroves and when dry seasons were free of saline dust storms.The occupation of two earlier coastal settlements, Monagrillo (4405 ? 75 BP (SI-2842) -3245 ? 100 BP (SI-2843)) and Zapotal (4010 ? 100 BP (Beta-21,389) - 3500 ? 80 BP(Beta-9574)) can be related to short-term features of the coastal landscape, in the formercase, a shallow coastal embayment (Willey and McGimsey 1954) and in the latter, theactive delta of the Santa Maria river.

Table Therelationship etween hearchaeologicalite of CerroMangoteandcoastlinedynamicsbetween10,000BPand thepresent afterBarber1981).DateBP Maxlmin Rateofsea SedimentationCoastlinedistance f levelrise vssealevel mechanicsmarine etting rise10,000 >15 Rapid Slower Transgression10,000-8000 15-1.4 Very rapid Slower Transgression8000-7000 1.4-1.2 Slower Stillstand?7000-5000 1.2-5.5 Slow Faster Progradation5000-3000 5.5-4.2 Faster Slower Transgression3000-Present 4.2-8.0 Gradual Faster Progradation


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Table 2 1C dates (uncalibrated) for pre-2500 BP occupations in the Santa Maria drainage, with brief subotanical remains. Only dates considered non-controversial by the excavators have been included. Saassociated with ceramics in the Monagrillo and subsequent styles.Site Site '4Cdate Lab. no. Mat, Lithics2type (BP) Bif EGC Mic AxeCoronaAG-123VampirosCarabaliCoronaLos SantanasCueva LadronesCerro MangoteCerro MangoteCerro MangoteAguadulceCerro MangoteCoronaAguaduiceVaca del MonteCerro MangoteCerro MangoteCerro MangoteCUEVA LADRONESCUEVA LADRONESMONAGRILLOMONAGRILLOAGUADULCEMONAGRILLOMONAGRILLO

rS3 10440 ? 650sm 8795 ? 135rs 8560 ? 160rs 8040 ? 3907440 ? 280rs 7100 ? 230rs 6860 ? 90sm,cm 6810 ?110

6710 ? 1706670 ? 215rs 6180 ? 1205990 ? 1805980 ? 1005840 ? 100rs 5630 ? 1805440 ? 1305140 ? 1205055 ? 1504800 ? 1004520 ? 1004405 ? 754350 ? 1604210 ? 904135 ? 804090 + 70

Beta-19105SI-6223Beta-5101Beta-9575Beta-19411Beta-9576TEM-123Y-458DTEM-206Beta-1219TEM-131TEM-174Beta-27529TEM-106Beta-12437TEM-176TEM-175TEM-207TEM-119TEM-124SI-2842TEM-208TEM-130SI-2844?-585

char Xshellchar Xchar Xchar Xchar Xcharcharshell*charshellshellshell*shellcharshellshellshell*SHELLCHARCHARSHELL*SHELLCHARCHAR

XXXXXyNN37/3N

NN37/3

N

N

XX

NX


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ZAPOTALAG-66CUEVA LADRONESCUEVA LADRONESZAPOTALVAMPIROSCUEVA LADRONESMONAGRILLOZAPOTALRio CobreZAPOTALZAPOTALMONAGRILLOMONAGRILLOMONAGRILLOMONAGRILLOVAMPIROSAGUADULCECARABALISE-111VAMPIROSVAMPIROSLA MULAAG-88LA MULAAGUADULCEAGUADULCEVAMPIROS

SMSM

rs

HA

SMVIL

4010 ? 1003945 ?+1153880 ? 803860 ? 903850 ? 703800 ? 1203770 ? 803615 ? 803610 ? 703580 ? 1503520 ? 803500 ? 803485 ? 1003385 ? 753325 ? 853245 ? 1003100 ? 602960 ? 802920 ? 1802855 ? 952840 ? 702820 ? 652820 ? 502685 ? 1052640 ? 602570 ? 952540 ? 702540 + 90

BETA-21,389SI-6238TEM-122TEM-121BETA-20,849BETA-5870TEM-120SI-2840BETA-21,388Beta-19,106BETA-20850BETA-9574SI-28391-28381-9384SI-2843SI-5687TEM-126BETA-19101BETA-19497BETA-27591SI-5682BETA-6006SI-6237BETA-21898TEM-107TEM-125BETA-27592

x

X

xx

SHELL*SHELLCHARSHELLSHELL*CHARSHELLCHARSHELL*charSHELL*SHELL*CHARCHARCHARCHARCHARSHELLCHARCHARSHELL*CHARSHELL*SHELLSHELL*SHELLSHELLSHELL*

xxx

xxxxxxxX XX X X

x xx x1:char = charcoal. *:corrected for 12C/13C. 2: bif = bifacial reduction, egc = edgeground cobbles, mic =axe = polished axes, palm:ac: Acrocomia, as: Astracaryum, e: Elaeis, mph = maize phytoliths and/or pfruit = b: Byrsonima, c: Curatella, m: cf Manilkaria, sP: Spondias, s: Sapotaceae, 3: rs = rockshelter,cm = cemetery.


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122 Richard Cooke and Anthony J. RanereThe lake coring programme, conducted after the coastal Vibracoring, revealed theSanta Maria watershed has been subject to tectonic activity during the Holocene. Basalsediments of a probable lateral explosion crater at El Valle date to 2700 ? 90 BP

(Beta- 11,637). A small lake near Chitra (basal date: 360 ? 80 BP [Beta- 12,795]) wasdammed by ejecta from a nearbycrater(Media Luna). It is possible that volcanic eruptionsand earthquakes influenced prehistoric settlement patterns in this watershed as they didelsewhere in Central America (Linares et alo1975; Sheets 1983; Sheets (ed.) 1984).(4) The watershed was (was not) inhabited10,000-7000 BPArchaeological sites that antedate 7000 BP have been identified in the watershed's fivephysiographic zones. Their small size and the very low artifact density at stratified locirefer to small, probably quite mobile populations. Human occupational debris assignableto this period is present in five rockshelters: Aguadulce, Carabali, Corona, Los Santanasand Vampiros (Table 2).The bifacial stone technology introduced or developed during the late glacial byPaleoindians continued. Surface-collected materials from a 0.7ha quarry-workshop, LaMula-Central, represent the only evidence in Panama for projectile point types seeminglyderived from Paleoindian models. These are similar to materials described as 'archaic'elsewhere in Central and North America. Bifacial technology gradually became de-pauperate: for example, at Carabali, where the pre-7000 BP record is longest, carefulplatform-preparation by grinding is not visible in deposits dated c. 8040 ? 390 BP(Beta-9575). We propose that the 'phasing-out' of bifacial chipped tools was anendogenous response to changing subsistence patterns, which we elucidate in the nextsection.

There is clear evidence for continuing human activities at La Yeguada across the ratherrapid (< 100 year) change from late glacial to Holocene climatic conditions and during theslower (c. 2,500 years) period of transition from montane to premontane evergreen forest.Charcoal and invasive weedy taxa reached very high levels by 8600 BP; over 90 per cent ofsedge (Cyperaceae) and Heliconia phytoliths continue to show evidence of burning. Thesepatterns are most parsimoniously attributed to the clearing of small swidden gardens bydry season slashing and burning.

(5) There were (were not) hunting-gatheringpopulations which did not utilizecoastal resources(6) Tropical forest agricultureof some kind did (did not) precede theintroductionof intensive maize agricultureEvaluation of these hypotheses is contingent upon the definition and identification ofstages of food production prior to agriculture ('a dependence on domesticated plants for asubstantialpartof the diet' (Rindos 1984:236)). This is a complex task, which is still a longway from being fulfilled. There are some uncomfortable paradoxes. On the one hand, wecannot guarantee that any Holocene populations in the drainage study subsisted withoutplants in initial stages of domestication, although we can probably do so safely for lateglacial Paleoindian populations. On the other hand, microbotanical sediment analyses and


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Prehistorichuman adaptationsto the seasonally dry forests of Panama 123inferences from the stone tool record offer more information on subsistence activities priorto 2500 BP than the macrobotanical record, which contains few taxa, none of theseindisputably a domesticate. lefining which segments of the population 'utilized' coastalresources, and when, is complicated by differential preservation of faunal remains and bythe fact that, in a narrowisthmus, Holocene communities were always less than 80km fromthe sea: marine foods can be readily distributed to quite 'inland' locations.In pre-2500 BP sites, the most abundant carbonized remains, other than wood, are palmfruits (identified taxa:Acrocomia mexicana, Astracaryumsp., Elaeis oleifera, and Scheeleasp.). These are typical of wet or disturbed habitats, i.e. swampy meadows, riverbanks andpaths, rather than of continuous forest cover. A. mexicana may have been introduced byhumans. Dicot fruits (Byrsonima crassifolia (Malphigiaceae), Hymenea courbaril (Legu-minoseae), andSapotaceae) and seeds of a fire-resistant savannatree (Curatellaamericana(Dillinaceae)) occur in smaller numbers (Table 2).

In all deposits where these plants have been identified, ground stone technologycomprises one-hand millingstones and edge-ground cobbles (Ranere 1980b, c). The latterare present by 8040 ? 390 BP at Carabali. It is conceivable that they were used to preparenon-dietary substances, such as fibresor bark. We continue to believe, however, that wearpatterns and their ubiquity and relative abundance before 2500 BP, in this watershed andelsewhere in Panama, reflect the preparationof root crops for human food. This inferenceis fortified by the fact that several wild species of tropical tubers and some domesticates(i.e. 'bitter' cassava) need to be traumatized in order to release or biochemically modifytoxins (Coursey 1967;Dufour 1988).Piperno identified phytoliths of a cultigen, arrowroot (Maranta arundinacea), in somerockshelter deposits. These occur at Vampiros in a stratum dated to 8560 ? 160 BP(Beta-5101). Alvina and coastal scrub, adjacent to the site at this time, is not arrowroot'snatural habitat. These data and the La Yeguada paleovegetational record suggest thatforest agrilocality (Rindos 1984: 176), which emphasized native tuberous taxa and treecrops, appeared in the study watershed not long after the inception of the Holoceneperiod.After c. 7000 BP there are sufficient changes in the paleovegetational and fieldarchaeological records to indicate that human impacts upon the watershed landscapeintensified. At La Yeguada, disturbance indicators become steadily more apparent. By3000 BP, the local landscape shows categoric effects of clearance and burning. It is possiblethat a mid-Holocene drier period (c. 7000-5000 BP) accelerated these processes byenhancing the effects of burning and retardingforest regrowth during fallow periods.

During the period 7000-4500 BP, maize (Zea mays) - a domesticated and introducedplant in Panama - appears as pollen and phytoliths in lake sediments in the Santa Mariawatershed and nearby (Piperno 1984; 1988) and in refuse lenses at small rockshelters in thefoothills (Ladrones, Los Santanas). Macroremainsof maize, however, are not identified inculturaldeposits screened over > 1mm mesh until much later: 2015 ? 80 BP (1-9702) is theearliest 4Cdate that can be associated stratigraphicallywith carbonized maize fragments.1560 ? 60 BP (Beta-46,402) is the earliest date run on maize remains themselves(compensated for 12C/13C fractionation) (both are from Sitio Sierra). Stone toolstraditionally associated with the grinding of maize for flour, i.e. carefully shaped metatesand two-handed cylindricalmanos, and ceramic vessels of sufficient size and thickness to


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124 Richard Cooke and Anthony J. Ranerecontain large quantities of fermented maize liquor make their firstappearance in domesticcontexts dated to 2500-2000 BP. Polished axes and adzes are found in association withthem. Dental caries, which reflects the intensity of carbohydrate consumption, isnoticeably less prevalent at Cerro Mangote c. 6000 BP than at Sitio Sierra c. 1600BP. The3C/12Csignal at the latter site predictably suggests a higher input of 14C plants in the diet.As soon as maize fragments displace palm fruits as the most abundant carbonizedremains, they occur at an evolutionary stage and in abundance levels that indicate that thiscultigen was an important crop in regional production systems, perhaps the mostimportant (Bird 1980; 1984; see also Galinat 1980; Pearsall and Piperno 1990). Thediscrepancy between the micro- and macrorecords suggests either that maize was used inways that were not conducive to its carbonization (i.e. as a green vegetable) or that itsphysical properties (i.e. 'popping') prevented its survivingburnt in a recognizable state.In summary, several lines of evidence indicate that the indigenous peoples of the SantaMaria watershed adopted maize as a staple crop quite rapidly between c. 2500 and 1500BP. The PSM data suggest strongly that there was a positive feedback between increasingkernel size and 'flouriness', the development of efficient polished stone tools for forestclearance, degradation of hillslope soils and the nucleation of settlements in lowlandcolluvial areas and the watershed's few broad highland valleys. Earlier regional projectsare consistent with this interpretation (Linares et al. 1975; Linares 1980). Even so, maizecultivation in the Santa Mariawatershed appears to have remained extensive even thoughthis crop was the major staple; as yet there is no archaeological or remote sensinginformation to suggest the presence of raised fields, terraces or other intensive cultivationtechniques.

We assume that Paleoindian (late glacial) populations were hunters and gatherers, butdo not know whether they utilized coastal resources. Remains of fish (including sea catfish(Ariidae) and mullet (Mugil)) and marine molluscs are present at Vampiros in the ninthmillennium BP stratum. At Corona (35km from the coast), a sample of Anadaratuberculosa, Protothaca sp., and Thais sp. shells was directly dated to 5980 ? 100 BP(Beta-27,529). The first two taxa are found buried in intertidal sand/mud substrates andthe last on rocky shores and mangrove roots. Seven mangrove-estuary mollusc species arepresent at Carabali (55km inland), along with sea catfish vertebrae, in contexts datedstratigraphicallybefore 4500 BP.The shells could be trinkets, rather than dietary resources. However, at Ladrones (justeast of the watershed, and > 20km inland 4000 BP) bones of thread-herring(Opisthonemalibertate) and a croaker (Ophioscion typicus) and several species of mangrove-estuaryshells and crabs occur in deposits dated to between 6860 ? 90 BP (TEM-123) and3770 ? 80 BP (TEM-120). The transportinland of sun-dried and salted nearshore fish likethese two small species (average adult weight < 300g), still an important activity in PacificPanama, is surely very ancient.Coastal resources had become important elements in the diet of coastal sites by7000-5000 BP. At Cerro Mangote, 25 families, 48 genera and 74 species of marine fishhave been identified. Mangrove-estuary crabs and molluscs and shorebirds were alsoconsumed. 14N/15Nvalues for human bone from this site give a more 'terrestrial' signalthan individuals from a cemetery at Sitio Sierra, a maize-farming village located 12kminland, where three AMS dates on human bone range from 1680 ? 60 BP to 1880 ? 95 BP


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Prehistoric human adaptationsto the seasonally dry forests of Panama 125[AA-3240-42]). Two explanations are possible: (1) Cerro Mangote was utilized seasonallyor occasionally by a population that spent most of the year away from the coast, or (2)euryhaline estuarine organisms prevailed in the diet. Archaeofaunal data support thelatter.A change in coastal fishingstrategies occurred after the abandonment of Cerro Mangoteand before the occupation of Monagrillo at c. 4400 BP. At the former site, fishingconcentrated on small taxa that can be caught with simple techniques in drying alvinapools, such as the eleotrine goby Dormitator latifrons and species that attain large sizes(> lkg) and can be abundant close to mangrove swamps and in tidal channels, such as thesea catfish (Ariidae spp.), corvinas (Sciaenidae: Cynoscion), snapper (Lutjanidae) andsnook (Centropomus). After 4400 BP, smaller shoaling taxa, i.e. herrings (Clupeidae:Opisthonema libertate), lookdowns (Carangidae: Selene peruviana) and small grunts(Haemulidae: Orthopristis chalceus), dominate the samples. The use of fine-meshed netsand watercraft is inferred.(7) Sedentary agricultural villages first appeared (a) before 4500 BP, (b)before 2300 BP or (c) after2300 BP and before 1500 BP(8) Population increasepreceded (resulted rom, was systematicallylinked to)intensive maize agricultureClarification of these hypotheses depends upon the final statistical and geographicalevaluation of transect and purposive survey data, which is not complete. We offer onlygeneralized statements that refer to major demographic processes on the understandingthat we might have to modify our stances in the near future.Pre-2500 BP cultural deposits are most visible in the cordillera, foothills and SantiagoPlain. Except for quarry-workshopslocated near chalcedony sources, where pre-2500 BPstone tool types can be found dispersed over > lha, these are small (< O.lha) loci, oftensituated on flattish spurs that overlook water courses. Some loci form clusters that suggestgroups of dwellings arranged into small hamlets. Circular spaces devoid of artifacts mayrepresent individual houses. Some rockshelters (i.e. Los Santanas) probably alsofunctioned as dwellings or storehouses in a hamlet or extended family unit.Dating the small, largely eroded or partially buried open sites is difficult. Three usefulfeatures of lithic technology are: (1) bifacial reduction techniques, which the rockshelterrecord suggests had disappeared c. 7000 BP; (2) a microtool industry, which employedbipolar reduction of agate and chalcedony nodules (this appeared after 7000 BP and is notpresent at post-2500 BP maize-farming villages); and (3) pointed blade-flakes andprismatic blades with flat, unmodified striking platforms and polished cutting tools madeof igneous materials, which are present only after 2500 BP.If we use these criteria, twenty-six sites in the watershed are pre-7000 BP in age. Tentimes this number can be assigned to the period 7000-2500 BP. Five out of the thirteentested rockshelters have pre-7000 BP deposits. All but one (Rio Bermejito) exhibit majorperiods of activity between 7000 and 2500 BP. The intensification of rockshelter use isepitomized by the increasing numbers and functional diversity of stone tools. Table 3illustrates this trend by reference to Carabali.It seems clear that the cordillera, foothill and Santiago Plain zones of the Santa Maria


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126 Richard Cooke and Anthony J. RanereTable3 Distribution f stone toolsbyfunctional lass andpreparationechnologyatCarabaliby chronological eriods fromValerio 1987: ablas19 and20).Function Preparation Date (BP)

technology pre-7000 7000-4500 4500-2000'Cutting Chipped 12 17 26Abrasion Chipped 6 4 50Perforating Chipped 0 14 20Splitting Chipped 0 4 16Microliths Chipped 0 13 50Split/cutting Ground/polished 0 0 3Grinding Ground 10 20 57Multipleuse Ground 0 2 0Totals: 28 74 2221Thisfigureprobablyncludesmaterials hatrepresenthesporadicuse of theshelterafterthis date.

drainage basin slowly accommodated more people and larger settlements during thepre-2500 BP Holocene. Hence, population increase occurred before the establishment ofvillages that cultivated maize extensively. We consider this pattern to be characteristic ofsmall, predominantly hillside communities that, lacking stone technologies appropriatefor the efficient removal of riverine gallery forests, practised a mixed economy oflong-fallow cultivation, gathering and hunting.In the old and new deltas, the only preceramic (i.e. pre-4500 BP) sites that have beenpositively identified are the two loci at La Mula-Sarigua ('West' and 'Central'),Aguadulce, Vampiros and Cerro Mangote. It is likely that other contemporary sites lieburied under Holocene colluvium and marine deposits. After 4500 BP, when crude,poorly fired ceramics of the Monagrillo tradition were introduced (Willey and McGimsey1954), a trend towards larger site size is observed (compare figures for Cerro Mangote,Monagrillo and Zapotal in Table 5). Whether these sites were used seasonally orcontinually is not entirely clear. The presence of ninety burials at Cerro Mangote and thedepth of the refuse deposits at this site and Zapotal favour relatively long periods of use.Zapotal's size (3. iha) and structure (Giausserand recovered part of a single oval ?house)suggest that a village existed here by 4000-3500 BP. We referred earlier to a possibleconnection between the maximization of coastal resources via technological improve-ments c. 4000 BP and population increase and/or sedentism.A causal relationship between maize's use as a staple crop and accelerated populationgrowth is inferred by the coincidence of this cultigen's visibility in the archaeological andpaleovegetational record after 2500 BP with equally visible changes in settlement size anddistribution.In Table 4 we present 14C dates that can be stratigraphically associated either withcarbonized remains of this cultigen or with tools traditionallyused for grindingit into flour.Table 5 presents the sizes of sites in the old and new delta zones. These are inferred from


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Prehistorichuman adaptations to theseasonally dry forests of Panama 127Table 4 4C ates (uncalibrated) for post-2500 BP occupations in the SantaMaria drainage, which are associated stratigraphicallywith carbonizedmaize remains or with manos and metates used for grinding maize forflour.Site Site type '4C date (BP) Lab. no. MaterialLa Mula-S vil 2340 ? 70 Beta-12,931 shell*La Mula-S vil 2270 ? 90 Beta-12,729 shell*La Mula-S vil 2220 ? 70 Beta-12,728 shell*La Mula-S vil 2190 ? 90 Beta-18,863 shell*Sitio Sierra cem 2190 ? 80 1-9704 charSitio Sierra vil 2015 ? 80 1-9702 charSitio Sierra vil 1975 ? 80 1-9703 charSitio Sierra cem 1880 ? 95 AA-3241 bone*'aSitio Sierra vil 1835 ? 90 1-9701 charSitio Sierra cem 1835 ? 90 AA-3242 bone*'aSitio Sierra vil 1715 ? 90 1-8613 charSitio Sierra cem 1680 ? 60+ AA-3240 bone*'aSitio Sierra vil 1640 ? 90 Gif-2346 charSitio Sierra cem 1560 ? 60+ Beta-46,402 maize*Sitio Sierra vil 1475 ? 110 1-8556 charSitio Sierra vil 1425 ? 60 SI-6300 shellSitio Sierra vil 1405 ? 50 SI-6301 shellSitio Sierra vil 1370 ?+110 Beta-46,401 char*AG-73 vil 1290 ?+55 Beta-46,398 char*'aSitio Sierra vil 1260 ? 60 SI-6302 shellAG-73 vil 1035 ?+55 Beta-46,390 char*'aSitio Sierra vil 1000 ? 40 SI-6303 shellSitio Sierra cem 920 ? 80 1-8381 charSitio Sierra vil 900 ? 40 SI-6299 shellSF-25 vil? 700 + 60 SI-6218 charAG-220 vil 625 ? 65 SI-6219 char*: calibrated for 13C/'3Cractionation; a: calculated by the AMS method;+: dates refer to the same burial (AG-3-75-3), adult male, buried withpolished axes.

the maximum areal extent of artifactual materials deemed contemporary by stylistic ortechnological affiliation. In the absence of house remains at all sites except Zapotal andSitio Sierra, these figures are, at best, approximate.The clearest indication of increasing nucleation is given by La Mula-Sarigua. This site'sapogee coincides with the abandonment of rockshelters as regular dwelling places andepitomizes the ascendancy of colluvial riverine zones as the axes of cultural developmentin the region.Use of the term 'sedentary' requires caution. Only at Sitio Sierra, where maximum sitesize inferredfrom artifactdistributionsis 45ha (1500-1200 BP), were superimposed housesidentified (Cooke 1979). This feature, and the size and density of refuse lensesaccumulated alongside groups of houses, suggest multiannual periods of occupation.Contact-period ethnohistoric data, however, warn against assuming that even chiefs'


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128 Richard Cooke and Anthony J. RanereTable5 Important archaeological sites in the Santa Marfadrainage (old and new deltas), whose size has beenestimated from the maximum extent (ha) of artifactualmaterials.Site Age (BP) Max extent (ha) ofcultural materials

for timeperiodCerro Mangote 7000-5000 0.175Monagrillo 4400-3200 1.400La Mula 4400-3200 1.300Zapotal 4000-3500 3.100La Mula 3200-2400 4.400La Mula 2400-1500 58.000Sitio Sierra 2000-1000 45.000AG-73 1500-1000 38.500La Mula 1000-500 74.000

villages in the fertile colluvium were used for longer than decades at a time. Cycles ofabandonment and reoccupation would have been conditioned by geomorphology,sanitary conditions and the regional social environment (i.e. alliance formation anddefence requirements). Even the largest sites we have investigated show evidence ofhiatuses in their occupational record.

ConclusionsSome of the multiple working hypotheses presented in 1981 now seem naive or poorlydefined, while the uncertainties subsumed in others have not necessarily been mitigated bynew evidence. Notwithstanding these weaknesses, however, we believe that the Santa.Maria watershed provides a clear example of uninterrupted human occupation fromc. 11,000 BP until the present. It also offers insights about how a particularpopulation ofNew World indigenous peoples adapted to and partitioned an area of the seasonally drytropics that, on its arrival as hunters and gatherers, was forested, albeit with formationsthat have no direct modern analogues.In order to respond adequately to hypotheses that, on the one hand, question theappropriateness of tropical forests for hunting and gathering communities, and on theother, underline the inappropriateness of the biome for human cultural complexity(Meggers 1987), it behoves us to provide substantive data about very subtle aspects ofpre-agricultural human settlement and subsistence. Sceptics have probably formed theopinion that the PSM has gone no further than confirming a process that is apparent inmany New World tropical forests: that nucleated maize-dependent prehistoric settlementsbecome visible fairly recently in the Holocene archaeological record (i.e. after 2500 BP)and that their development is causally related to population increase and settlementnucleation (e.g. Roosevelt 1980). We accept that the pre-3000 BP record is conducive to


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Prehistorichuman adaptationsto theseasonally dry forests of Panama 129multiple interpretation. We hope, however, that most readers will concur that collabor-ation between archaeologists and specialists in forest history has established beyondreasonable doubt that, once humans arrived in the forested Santa Maria drainage, theyquickly began to modify it in ways that, not long after the inception of the Holoceneperiod, are highly suggestive of plant domestication, swidden farming and hamlet, if notvillage, life. Hence, this process is detectable several millennia before the introduction ofpolychrome pottery, metalwork, fine metates and the other artifacts for which centralPanama is justly famous.Elsewhere (Cooke and Ranere 1992; Ranere and Cooke in press b) we evaluatedarguments for and against inferring that this population was culturally and geneticallyhomogeneous during the Precolumbian era and that its direct descendants are the extantChibchan-speaking polities of central and western Panama (the Bugle and Ng6b6). Wepointed out that exogenous demic influences might be responsible, at least in part, forshifts in settlement and subsistence patterns, which seem quite rapidin the existing record,i.e. c. 7000 BP (when maize firstappears) and about 2500 BP (when maize reflects geneticvariability and sites become fewer and larger). These problems are by no means new(Pinart 1882; Linares and Ranere 1980). But they are complex. We will continue toexplore them with the aid of linguists, specialists in human microevolution and physicalanthropologists (Arias et al. 1988;Barrantes et al. 1990; Constenla 1990).27.ix.91 R.G.CSmithsonian Tropical Research InstituteAncon, Panama

A.J.RDepartmentof AnthropologyTemple UniversityPhiladelphia, USA 19122

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130 Richard Cooke and Anthony J. RanereBird, R. McK. 1980. Maize evolution from 500 BC to the present. Biotropica, 12: 30-41.Bird, R. McK. 1984. South American maize in central America? In Pre-Columbian Plant Migration(ed. D. Z. Stone). Papers of the Peabody Museum of Archaeology and Ethnology, 76: 39-65.Bradbury, J. P., Leyden, B., Salgado-Labouriau, M., Lewis, R. M. Jr., Schubert, C., Binford, M.W., Frey, D. G., Whitehead, D. R. and Weibezahn, F. H. 1981. Late Quaternary environmentalhistory of Lake Valencia, Venezuela. Science, 214: 1299-305.Bush, M. B. and Colinvaux, P. A. 1990. A pollen record of a complete glacial cycle from lowlandPanama. Journal of VegetationScience, 1: 105-18.Cedefio, H. 1985. El medio ffsico del area de Sarigua: base para una polftica de ordenamientoespacial. Graduation thesis, Department of Geography, Universidad de Panama.Clary, J., Hansell, P., Ranere, A. J. and Buggey, T. 1984. The Holocene geology of the westernParita Bay coastline of central Panama. In Lange (ed.) 1984: 55-83.Colinvaux, P. A. and Bush, M. B. 1991. The rain forest ecosystem as a resource for hunting andgathering. American Anthropologist, 93: 153-60.Columbus, F. 1959. The Life of the Admiral ChristopherColumbus by his Son Ferdinand (trans. andannotated by Benjamin Keen). New Brunswick NJ: Rutgers University Press.Constenla, A. 1990. Las Lenguas del Area Intermedia:una Introduccion a su Estudio Areal. San Jos6(Costa Rica): Universidad de Costa Rica.Cooke, R. G. 1979. Los impactos de las comunidades agricolas precolombinas sobre los ambientesdel Tr6pico estacional: datos del Panama prehist6rico. Actas del TVSimposio Internacional deEcologia Tropical, 3: 917-73. Panama: Instituto de Cultura and others.Cooke, R. G. 1984a. Archaeological research in central and eastern Panama: a review of someproblems. In The Archaeology of Lower Central America (eds F. W. Lange and D. Z. Stone).Albuquerque: University of New Mexico Press, pp. 263-302.Cooke, R. G. 1984b. Birds and men in prehistoric central Panama. In Lange (ed.) 1984: 243-81.Cooke, R. G. In press. The relation of estuarine fish resources to archaeological site history andlocation: a comparison between two prehistoric settlements of different ages on the Pacific coast ofPanama. Proceedings of the Circum-PacificPrehistory Conference, Seattle. Olympia: University ofWashington Press.Cooke, R. G. and Ranere, A. J. 1984. The 'Proyecto Santa Maria': a multi-disciplinary analysis ofprehistoric adaptations to a tropical watershed in Panama. In Lange (ed.) 1984: 3-30.Cooke, R. G. and Ranere, A. J. 1989. Hunting in Prehistoric Panama: a Diachronic Perspective. InThe Walking Larder: Patternsof Domestication, Pastoralism and Predation (ed. J. Clutton-Brock).London: Unwin Hyman, pp. 295-315.Cooke, R. G. and Ranere, A. J. 1992. The origins of wealth and hierarchy in the Central Region ofPanama, with observations on its relevance to the phylogeny of Chibchan-speaking polities inPanama and elsewhere. In Wealth and Hierarchy in the Intermediate Area (ed. F. W. Lange).Washington DC: Dumbarton Oaks pp. 243-316.Coursey, D. G. 1967. Yams: an account of thenature, origins, cultivation, and utilisationof the usefulmembersof the Dioscoreaceae. London: Longmans.Dere, C. 1981. The geological and paleographic setting of an archaeological site on the southwesterncoast of Parita Bay, Panama. Master's thesis, Department of Geology, Temple University.Dillehay, T. 1989. Monte Verde: a Late Pleistocene Settlement in Chile, Vol. 1. Washington DC:Smithsonian Institution Press.Dufour, D. L. 1988. Cyanide content of cassava (Manihot esculenta, Euphorbiaceae) cultivars usedby Tukanoan Indians in northwest Amazonia. Economic Botany, 42: 255-66.


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132 Richard Cooke and Anthony J. RanerePiperno, D. R. and Clary, K. H. 1984. Early plant use and cultivation in the Santa Maria Basin,Panama: data from phytoliths and pollen. In Lange (ed.) 1984: 85-121.Piperno, D. R., Bush, M. B. and Colinvaux, P. A. 1990. Paleoenvironments and human settlementin late-glacial Panama. QuaternaryResearch, 33: 108-16.Piperno, D. R., Bush, M. B. and Colinvaux, P. A. 1991a. Paleoecological perspectives on humanadaptation in Panama. I. The Pleistocene. Geoarchaeology, 6: 201-26.Piperno, D. R., Bush, M. B. and Colinvaux, P. A. 1991b. Paleoecological perspectives on humanadaptation in Panama. II. The Holocene. Geoarchaeology, 6: 227-50.Piperno, D. R., Clary, K. H., Cooke, R. G., Ranere, A. J. and Weiland, D. 1985. Preceramic maizein central Panama. American Anthropologist, 87: 871-8.Ranere, A. J. 1980a. Human movement into Tropical America at the end of the Pleistocene. InAnthropological Papers in Memory of Earl H. Swanson (eds L. B. Harten, C. N. Warren and D. R.Tuohy). Pocatello: Idaho Museum of Natural History, pp. 41-7.Ranere, A. J. 1980b. Preceramic shelters in the Talamancan range. In Linares and Ranere (eds)1980: 16-43.Ranere, A. J. 1980c. Stone tools from the rio Chiriquf shelters. In Linares and Ranere (eds)1980:316-53.Ranere, A. J. In press a. Implements of change in the Holocene of Central Panama. In Arqueologiay Medio Ambiente (eds T. van der Hammen and 0. Ortiz-Troncoso). Proceedings of the 50thInternational Congress of Americanists, Amsterdam.Ranere, A. J. In press b. Les cultures pal6oindiennes en Am6rique Centrale dans un contextepal6o6cologique. L'Anthropologie.Ranere, A. J. and Cooke, R. G. In press a. Early human migration through the Isthmus of Panama.Proceedings of the Circum-PacificPrehistory Conference. Olympia: Washington State University.Ranere, A. J. and Cooke, R. G. In press b. Identifying cultural boundaries in prehistoric Panama.Ranere, A. J. and Cooke, R. G. In press c. Paleoindian occupation in the Central American tropics.Clovis, Origins and Human Adaptation (eds R. Bonnichsen and K. Fladmark). Orono: Center forthe Study of the First Americans.Ranere, A. J. and Hansell, P. 1978. Early subsistence patterns along the Pacific coast of Panama. InPrehistoric Coastal Adaptations (eds B. L. Stark and B. Voorhies). New York: Academic Press,pp. 43-59.Rindos, D. 1984. The Origins of Agriculture: an Evolutionary Perspective. Orlando: AcademicPress.Romoli, K. 1987. Los de la Lengua Cueva: los Grupos Indigenas del Istmo Orientalen la Epoca de laConquista Espafola. Bogota: Instituto Colombiano de Antropologia and Instituto Colombiano deCultura.Roosevelt, A. C. 1980. Parmana:prehistoric Maize and Manioc Subsistence along the Amazon andOrinoco. New York: Academic Press.Sheets, P. D. 1983. Archaeology and Volcanism in Central America: the Zapotitldn Valley of ElSalvador. Austin: Texas University Press.Sheets, P. D. (ed.). 1984. Investigaciones arqueol6gicas en la Cordillera de Tilaran, Costa Rica.Vinculos 10.Toribio-Medina, J. 1913. El Descubrimiento del Oceano Pacifico: Vasco Nunfiez e Balboa, Fernandode Magallanes y sus Companferos.Tomo II: Documentos Relativos a Niunezde Balboa. Santiago deChile: Imprenta Universitaria.


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Prehistoric human adaptationsto theseasonally dry forests of Panama 133Valerio-Lobo, W. 1985. Investigaciones preliminares en dos abrigos rocosos en la Regi6n Central dePanama. Vinculos, 11: 17-29.Valerio-Lobo, W. 1987. Analisis functional y estratigraficode Sf-9 (Carabali), un abrigo rocoso en laregi6n central de Panama. Graduation thesis, Department of Anthropology, Universidad de CostaRica.Weiland, D. 1984. Prehistoric settlement patterns in the Santa Maria drainage of Panama: apreliminary analysis. In Lange (ed.) 1984: 31-54.Willey, G. R. and McGimsey, C. R., III. 1954. The Monagrillo Cultureof Panama. Papers of thePeabody Museum of Archaeology and Ethnology, 49(2). Cambridge: Harvard University Press.

Abstract

Cooke, R. and Ranere, A. J.Prehistoric human adaptations to the seasonally dry forests of PanamaSince 1981 research in the Santa Maria Basin, an area of seasonally dry forest on the Pacific coast ofPanama, has shown that collating environmental records with regional archaeological surveysenables us to identify small dispersed populations in New World tropical forests. Original multipleworking hypotheses are reevaluated. Data show that: 1) the watershed has been occupiedcontinuously since 11,000 BP; 2) population increases took place c. 7000 and 2500 BP; 3)domestication of some native tubers occurred before 7000 BP; 4) maize, introduced 7000-5000 BP,was extensively cultivated by sedentary communities by 2500 BP; 5) forest degradation took placeearlier on stony hillslopes than in flatter colluvial zones; and 6) coastal resources, first visible at 8600BP, became important between 7000 and 3500 BP.

seasonally foresta panama

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