Remains of Palms (Palmae) at Archaeological Sites in the New World: A Review

Remains of Palms (Palmae) at Archaeological Sites in the New World: A ReviewAuthor(s): Gaspar Morcote-Ríos and Rodrigo BernalSource: Botanical Review, Vol. 67, No. 3 (Jul. - Sep., 2001), pp. 309-350Published by: Springer on behalf of New York Botanical Garden PressStable URL: http://www.jstor.org/stable/4354394 .

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THE BOTANICAL REVIEW

VOL. 67 JULY-SEPTEMBER 2001 No. 3

Remains of Palms (Palmae) at Archaeological Sites in the New World: A Review

GASPAR MORCOTE-RioS AND RODRIGO BERNAL

Instituto de Ciencias Naturales Universidad Nacional de Colombia Apartado 7495, Bogota, Colombia

I. Abstract/Resumen .................... 309 II. Introduction .................... 310

III. Materials and Methods ........ ............ 311 IV. Results ........... .............. 311 V. Discussion ........ ............ 335

A. Acrocomia aculeata .................... 337 B. Bactris gasipaes .................... 338 C. Oenocarpus bataua .................... 340 D. Elaeis oleifera .................... 341 E. Attalea butyracea .................... 342 F. Mauritiaflexuosa .................... 342 G. Cocos nucifera .................... 342 H. Parajubaea cocoides .................... 342

VI. Acknowledgments ........ ............. 343 VII. Literature Cited ..................... 343

I. Abstract

A review of palm remains recorded at archaeological sites throughout the New World is presented. Remains have been found at 130 sites from the southern United States to southern Uruguay. They are of four kinds: carbonized or dry endocarps or seeds, phytoliths, pollen, and implements. Twenty-nine genera and at least 50 species of palms (i.e., about 9% of all Ameri- can species) have been recorded. The oldest record dates back to 14,700 B.P. for carbonized endocarp fragments of an unidentified palm in Rond6nia, Brazil. The use of palms, as recorded from remains, was particularly widespread after 9000 B.P. The predominant remains are endocarps of Acrocomia, Attalea s.l., Astrocaryum, Bactris, Syagrus, Elaeis, and Oeno- carpus, all of which are important sources of edible oils or edible fruits and are still widely used by aboriginal peoples. The review supports the hypothesis that human groups have

Copies of this issue [67(3)] may be purchased from the NYBG Press, The New York Botanical Garden, Bronx, NY 10458-5125, U.S.A. Please inquire as to prices.

The Botanical Review 67(3): 309-350, July-September 2001 (D 2002 The New York Botanical Garden 309



310 THE BOTANICAL REVIEW

played an important role in the dispersal of some palm species in the neotropics. Human- aided dispersal of Acrocomia aculeata from South America into Central America, and of Oe- nocarpus bataua from northwestern Amazonia to other areas, is postulated. Archaeological remains support the hypothesis that pejibaye (Bactris gasipaes) was domesticated in the inter- Andean valleys or on the adjacent Pacific lowlands of Colombia and later introduced into the Amazon Basin.

Resumen

Se presenta una revision de los registros de vestigios de palmas en yacimientos ar- queol6gicos en el Nuevo Mundo. Se han encontrado vestigios en 130 yacimientos, desde el sur de los Estados Unidos hasta el sur de Uruguay. Los vestigios son de cuatro tipos: endocar- pos quemados o secos, fitolitos, polen e implementos. Representan 29 generos y por lo menos 50 especies de palmas (es decir, cerca del 9% de todas las especies americanas). El registro mas antiguo data de 14,700 A.P., para fragmentos de endocarpo carbonizados de una palma no identificada en Rond6nia, Brasil. El uso de las palmas, tal como lo evidencian los vestigios, estuvo particularmente extendido despues del 9000 A.P. Los vestigios predominantes son en- docarpos de los generos Acrocomia, Attalea, Astrocaryum, Bactris, Syagrus, Elaeis, y Oeno- carpus, los cuales son fuentes importantes de aceites o frutos comestibles, todavia usados por pueblos abori genes. La revision apoya las hipotesis segu'n las cuales los grupos humanos han jugado un importante papel en la dispersion de algunas palmas en el neotropico. Se postula la dispersion antropogenica de Acrocomia aculeata desde Suramerica hacia Centroamerica y de Oenocarpus bataua desde el noroeste de la Amazonia hacia otras areas. Los vestigios ar- queologicos apoyan la hipotesis de que el chontaduro (Bactris gasipaes) fue domesticado en los valles interandinos o en las tierras bajas aledafias del Pacifico en Colombia, y despues in- troducido a la Amazonia.

II. Introduction

The widespread use of palms by humans has received much attention worldwide, and many studies have focused on the present relationship of people with palms (see references in Balick & Beck, 1990). In contrast, few studies focus on the prehistoric use of palms. How- ever, it has been suggested that such prehistoric use has been the base of dispersion, selection, domestication, and extinction (Patifno, 1963; Dransfield et al., 1984; Balee, 1988, 1989; Clement, 1988, 1992; Kahn & Moussa, 1995; McKillop, 1996).

Palm remains are common at archaeological sites, but they seldom receive much attention, and they are often cited in studies only incidentally. The few studies in which palm remains have been discussed in depth have shown the importance of this family for human groups in the past (Lentz, 1990a; Romero, 1995; McKillop, 1996; Morcote et al., 1996, 1998; Roosevelt et al., 1996). These findings have moved us to review the occurrence of palm remains at archaeological sites, as recorded in the literature. This review seeks to compile whatever is known of the past use of palms in America, as documented by remains, and to call the attention of archaeobotanists and archaeologists to this important family, which was perhaps as important as corn and cassava in the development of prehispanic societies in the Americas.

III. Materials and Methods

This paper is based on two kinds of references: published papers or books and unpublished reports, mostly for Colombian sites. Additionally, in two cases we have cited personal com-



REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 311

munications relating to ongoing research. We have had only limited access to unpublished reports or university theses outside Colombia. Published references often mention palms just in passing, so they are sometimes difficult to track; because of this, we are aware that we may have missed some records. On the other hand, except for one case (Gnecco & Mora, 1997), we have been unable to check the accuracy of identifications provided in the cited references; thus, unless there are obvious incongruities of distribution, we have not challenged identifications and have only made the appropriate nomenclatural changes. In contrast, the numerous unpublished reports from Colombia are based on studies carried out by or in collaboration with one of us (GM), so we have been able to verify identifications.

For our review we have included Easter Island, which, though not strictly a part of the New World, is politically dependent on Chile and includes a palm species related to the South American genus Jubaea (Dransfield, 1991). Moreover this island exhibits one of the most fas- cinating cases of palm remains. Finally, we have included pollen records only when the pollen grains were found in an archaeological context, not those found in palaeoecological studies.

IV. Results

Results of the review are presented in Tables I and II. We found references to 130 archaeological sites ranging from the southern United States to southern Uruguay (Figs. 1-2); that is, most of the area of distribution of palms in America. Elevation of the sites ranges from sea level to 2600 m, but most sites (68%) are located below 500 m. The remains found are of four main kinds: carbonized or dry endocarps or seeds, phytoliths, pollen, and implements. Addi- tionally, fragments of wood and leaves have been found at a few sites. There were 137 records of remains at the 130 sites. The kind of remains most frequently recorded are endocarps and seeds, which occurred at 71 % of the sites. These remains are often referred to by authors as "fruits." They are found carbonized, dried, or slightly mineralized. Most endocarps are found fragmented; this can be the result of intentional cracking for extracting the seed (as in A ttalea) or of a relatively thin endocarp (as in Elaeis oleifera or Bactris gasipaes).

Pollen has been found at 8.4% of the sites, and records are available for at least 21 species. For four species pollen is the only kind of remains known. This pollen may indicate that a par- ticular species was being used by people or that the grains were airborne to the site.

Phytoliths have been found at 12.3% of the sites. These structures are the archaeological form of the silica bodies, which occur in all palm organs, except roots (Tomlinson, 1990). Phytoliths remain in the soil at archaeological sites and can be an indicator of plant organs that were used at those sites. With some exceptions, phytoliths are diagnostic of palm subfamilies (Piperno, 1988).

Implements have been found at 2.3% of the sites; they include objects made from stems (like spears), spines (like darts), or leaves (like mats). In most cases they are associated with tombs and funerary clothes. The botanical identification of palm implements, as given in the references, is usually incomplete.

The age of the remains recorded ranges from 14,700 years B.P. up to the time of the Euro- pean conquest. Between 9000 and 5000 B.P. (Fig. 3) there is an increase both in the number of remains and in the diversity of species being used. Twenty-nine genera and at least 50 species of palms (i.e., about 9% of all palms known in America) have been identified at the sites. The most abundant remains belong to the genera Acrocomia, Attalea s.I., Bactris, Syagrus, Elaeis, Astrocaryum, and Oenocarpus. The species used in these genera have a high contents of edible oil, and most of them are widespread or locally abundant or highly productive. The five

best-represented palms at archaeological sites are Acrocomia aculeata, Attalea butyracea,

(Text continues on p. 335)



Table I List of palms represented by remains at archaeological sites in the New World. Identified species are arranged alphabetically, followed by

doubtful specific identifications; these are followed by identifications to genus, and then by doubtful generic identifications; unidentified remains come last. Within each category, records are arranged from the oldest to the most recent date.

Species Remains Age Site Altitude Reference Notes (B.P.) (m)

Acrocomia aculeata Carbonized endocarps 9530 ? 100 Colombia: Cauca: Morales: 1690 Gnecco & Mora, 1997 Misidentifications (Jacq.) Lodd. ex San Isidro Mart.

A. aculeata Carbonized endocarp 8040 +390 Panama: Veraguas: Carabali 300 Cooke, 1992 fragments

A. aculeata Carbonized endocarp 7000-4500 Panama: Coclk: Aguadulce <50 Cooke & Ranere, 1992 _ fragments

A. aculeata Carbonized endocarp 6750-2450 Panama: Chiriqui: Casita de 700 Ranere, 1980; Smith, 1980 As A. cf. vinifera fragments Piedra (QS-1) O

A. aculeata Carbonized endocarp 6750-2450 Panama: Chiriqui: Trapiche 700 Ranere, 1980; Smith, 1980 As A. cf vinifera fragments (QS-2) >z

A. aculeata Dry endocarps 6750 Mexico: Puebla: Coxcatlan 1200 Smith, 1965; Lenz, 1990a A. aculeata Carbonized endocarp 5630 + 180 Panama: Veraguas: Vaca del 800 Cooke, 1992; Cooke & Ranere, >

fragments Monte 1992 A. aculeata Endocarp fragments 4210 ? 90; Panama: Coclk: Aguadulce <50 Cooke & Ranere, 1992

2960 ?80 A. aculeata Pollen 3150 Dominican Republic: Mouth of <200 Vega, 1995 As A. quisqueyana

Rio Iguamo A. aculeata Carbonized endocarp 2920 t 180 Panama: Veraguas: Carabali 300 Cooke & Ranere, 1992

fragments A. aculeata Carbonized endocarps & 2500-1000 Honduras: El Cajon: Rio Saluco:- 300-600 Hirth & Coskren, 1989; Lentz,

seeds Salitrbn Viejo (PC-1) 1989 A. aculeata Carbonized endocarps & 2500-1000 Honduras: Yoro: El Cajon: 300-600 Hirth & Coskren, 1989; Lentz,

seeds Guarabuqui (PC-15) 1989 A. aculeata Carbonized endocarps & 2500-1000 Honduras: El Cajon: PC-22 300-600 Hirth & Coskren, 1989; Lentz,

seeds 1989 A. aculeata Pollen 2250 Dominican Republic: Santo <50 Vega, 1995 As A. quisqueyana

Domingo A. aculeata Endocarp fragments 2190+ 80; Panama: Coclk: Sitio Sierra <200 Cooke, 1992

1475 ? 100 A. aculeata Fruits 2150-1850 Belize: Northem Belize: Cerros <50 Crane, 1986; Lentz, 1990a



A. aculeata Carbonized endocarp 1750-1550 Panama: Chiriqui: Cerro Punta: 2000 Ranere, 1980; Smith, 1980 fragments Piti (BU-17)

A. aculeata Carbonized endocarps 1650-450 Belize: South coast: Wild Cane <50 McKillop, 1996 Cay

A. aculeata Carbonized endocarps 1650-450 Belize: South coast: Frenchman's <50 McKillop, 1996 Cay

A. aculeata Carbonized endocarps 1650-450 Belize: South coast: Orlando's <50 McKillop, 1996 Jewfish

A. aculeata Carbonized endocarps 1650-450 Belize: South coast: Pelican One <50 McKillop, 1996 Pot

A. aculeata Carbonized endocarps 1650-450 Belize: South coast: Tiger Mound <50 McKillop, 1996 > A. aculeata Endocarps 1550-1050 Honduras: Copan: Ostuman: Los 500-100 Lentz, 1990b t

Mangos 0 O A. aculeata Carbonized endocarps 1550-850 Brazil: Para: Ilha do Marajo: Teso <30 Roosevelt, 1991, 1999 t

dos Bichos >

A. aculeata Carbonized endocarp 1500 ? 40; Colombia: Casanare: Aguazul: 525 Morcote, 1998 >

fragments 1080 + 40 Cuarto Unete: Finca Santa Marta

A. aculeata Carbonized endocarp 1350 Panama: Chiriqui: Barriles 1320-14 Ranere, 1980; Smith, 1980 >

fragments (BU-24) 00 > A. aculeata Endocarps 1350-1050 Honduras: Ulua: Cerro Palenque 500-100 Joyce, 1985; Lentz, 1990a

0

A. aculeata Carbonized endocarp 1150-950 Panama: Chiriqui: La Pitahaya <100 Ranere, 1980; Smith, 1980 > fragments (IS-3) 0

A. aculeata Endocarps 1050-750 Belize: Northern Belize: Colha <50 Caldwell, 1980; Turner & Mik- O sicek, 1984; Lentz, 1990a C)

A. aculeata Carbonized endocarps 1050-750 Guatemala: Tikal 50-200 Caldwell, 1980 A. aculeata Carbonized endocarp 680 + 60 Brazil: Minas Gerais: Rio Que- 770 Anonymous, 1995 As A. sclerocarpa >

fragments bra-Anzol (MG 30) A. aculeata Pollen 458 Dominican Republic: Boca de <50 Vega, 1995

Yuma 1

A. aculeata Endocarps Not given Costa Rica: No site given Not given Smith, 1987, cited in McClung As A. mexicana de Tapia, 1992

Acrocomia hassleri Carbonized endocarps 1550-885 Brazil: Para: Ilha do Marajv: <30 Roosevelt, 1991, 1999 (Barb. Rodr.) Hahn Teso dos Bichos

Acrocomia Carbonized endocarps 11,200-9800 Brazil: Para: Santarem: Monte 80 Roosevelt, 1999 Most probably A. Alegre: Caverna da Pedra aculeata, based Pintada on present dis-

tribution



Table I (continued)

Age Altitude Species Remains (B.P.) Site (m) Reference Notes

Acrocomia Carbonized fruits 1960 ? 70 Brazil: Rio de Janeiro: Sernam- <200 Heredia & Beltrao, 1980 betiba

Aiphanes aff. aculeata Carbonized endocarp 2830 * 70- Colombia: Santander: Cimitarra: 158 Piazzinni, 1995 2740 70 Piamonte

A. cf. aculeata Willd. Carbonized endocarp 590 ? 60 Colombia: Cesar: Aguachica: San 260 Bernal, 1996; Morcote, 1996 fragments Francisco

A. cf. aculeata Dry endocarp fragments 360 + 70 Colombia: Santander: Santa Rosa 1476 Romero, 1999 del Espejo

Aiphanes Seeds & pollen 1350-915 Colombia: Antioquia: San 1000 Rodriguez, 1997c Older date is rela- Rafael: El Bizcocho tive chronology

cf. Allagoptera Carbonized fruits 5520 + 120; Brazil: Rio de Janeiro: Cabo Frio: 9 Scheel-Ybert, 1998 H

3940 ? 140- Forte l 2240 70

cf. Allagoptera Carbonized fruits 4340 70; Brazil: Rio de Janeiro: Cabo Frio: 12 Scheel-Ybert, 1998 0 1800 Salinas Peroano >

cf. Allagoptera Carbonized fruits 4160 ? 190 Brazil: Rio de Janeiro: 5 Scheel-Ybert, 1998 Saquarema: Beirada

cf. Allagoptera Carbonized fruits 3760; 1430 Brazil: Rio de Janeiro: Cabo Frio: 10 Scheel-Ybert, 1998 Boca da Barra

cf Allagoptera Carbonized fruits & 2080 + 70 Brazil: Rio de Janeiro: Arraial do 40 Scheel-Ybert, 1998 wood Cabo: Ponta da Cabeqa

Astrocarywn cham- Carbonized seed frag- 9250 ? 140- Colombia: Caqueta: Araracuara: 200 Cavelier et al., 1995; Morcote As A. aculeatum bira Burret ments 9125 ? 250 Penia Roja et al., 1996, 1998

A. chambira Carbonized & dry seed 1190-880 Colombia: Guaviare-Caqueta: 500 Van der Hammen & As A. aculeatum fragments Serrania de Chiribiquete: Castafno, 1995; Morcote,

Abrigo Selva unpubl. data Astrocaryumjauari Carbonized whole & 9250 ? 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier et

Mart. fragmented seeds 9125 + 250; Pefia Roja al., 1995; Morcote et al., 1900? 30- 1996, 1998 385 + 50

Astrocarywm malybo Carbonized endocarp 2830+ 70- Colombia: Santander: Cimitarra: 158 Piazzinni, 1995 H. Karst. fragments 2740? 70 Piamonte

A. malybo Carbonized endocarp 850 + 50 Colombia: Santander: Cimitarra: 150 Rodriguez & Restrepo, 1997 fragments San Juan: Finca La Marina

A. cf malybo Carbonized endocarp 1120 + 60- Colombia: Cesar: Curumani: 62 Bemal, 1995 fragments 790 + 60 Los Serenos



A. cf malybo Carbonized whole & 850 + 80; Colombia: Antioquia: Puerto 130 Penia & Morcote, 1997a fragmented endocarps 700 + 70 Nare: La Sierra: Finca Angos-

tura A. malybo Carbonized endocarp 730 + 50; Colombia: Caldas: La Dorada: 176 Pefia & Morcote, 1997b

fragments 420 + 70 La Juana A. malybo Carbonized endocarps 565 ? 115 Colombia: Caldas: La Dorada: 249 Morcote & Cavelier, 1999;

Los Achiles Mendez & Escobar, 2000 Astrocarywn muru- Carbonized endocarps 9250 + 140- Colombia: Caqueta: Araracuara: 200 Cavelier et al., 1995; Morcote As A. sciophilum

muru var. ciliatum 9125 + 250 Pefia Roja et al., 1996, 1998 (Kahn) Henderson

Astrocaywn muru- Pollen 1565 t 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 As A. sciophilum muru var. ciliatum 775 ? 25 Abeja c

Astrocarywn standley- Carbonized endocarps 2300-1600 Colombia: Nariiio: Tumaco: 5 Patiuno, 1999 0 anum L. H. Bailey Las Marias

A. standleyanum Carbonized endocarps 1105 + 50 Colombia: Valle del Cauca: <50 Romero, 1994, 1996; Salgado > Calima: Ord6fiez I & Stemper, 1995

Astrocaryun aff. tri- Carbonized endocarps 2830 70- Colombia: Santander: Cimitarra: 158 Piazzinni, 1995 andrum Galeano, 2740 70 Piamonte > R. Bemal & Kahn q

A. cf. triandrum Carbonized endocarp 730 + 50; Colombia: Caldas: La Dorada: 176 Peiia & Morcote, 1997b > fragments 420 ? 70 La Juana )

A. cf. triandrum Carbonized endocarps 565 4 115 Colombia: Caldas: La Dorada: 249 Morcote & Cavelier, 1999; > Los Achiles Mendez & Escobar, 2000 m

Astrocarywn vulgare Carbonized endocarps 11,110 ? 310; Brazil: Para: Santarem: Monte 80 Roosevelt et al., 1998; ? Mart. 10,655 +285 Alegre: Caverna da Pedra Roosevelt, 1999 O

Pintada C) A. vulgare Carbonized endocarps 1335 ? 185 Brazil: Para: Ilha do Marajo: <30 Roosevelt, 1991, 1999

Guajara Astrocaryum Pollen 6000-5000 Colombia: Caldas: Villamaria: 1900 Rodriguez, 1997a High elevation for c'

Los Arrayanes an Astrocarym um Astrocaryum Carbonized endocarp 56304 180 Panama: Veraguas: Vaca del 800 Cooke, 1992; Cooke & Ranere, cn

fragments Monte 1992 Astrocaryum Carbonized endocarps 4260 ? 75; Brazil: Rio de Janeiro: Corondo <200 Carvalho, 1984

3010( 80 Astrocaryum Carbonized endocarp 2830 ? 70 Colombia: Santander: Cimitarra: 158 Piazzinni, 1995

fragments 2740? 70 Piamonte Astrocaryum Carbonized endocarps 2800 Brazil: Minas Gerais: Santana do <800 Resende & Prous, 1991

Riocho



Table I (continued)


Astrocaryum Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 249 Mora et al., 1991 775 25 Abeja

Astrocaryum Carbonized endocarp 1120 ? 60- Colombia: Cesar: Curumani: Los 62 Bemal, 1995 fragments 790 ? 60 Serenos

Astrocaryum Carbonized endocarp 850 ? 50 Colombia: Santander: Cimitarra: 150 Rodriguez & Restrepo, 1997 San Juan: Finca La Marina

Astrocarywm Carbonized endocarp 730 ? 50; Colombia: Caldas: La Dorada: 176 Pefia & Morcote, 1997b fragments 420 1 70 La Juana

cf. Astrocaryum Carbonized fruits 5520 ? 120; Brazil: Rio de Janeiro: Cabo Frio: 9 Scheel-Ybert, 1998 3940 140- Forte H

2240 70 cf. Astrocaryum Carbonized fruits 4340 ? 70; Brazil: Rio de Janeiro: Cabo Frio: 12 Scheel-Ybert, 1998

1800 Salinas Peroano 0 cf. Astrocaryum Carbonized fruits 4160 ? 190 Brazil: Rio de Janeiro: 5 Scheel-Ybert, 1998 H

Saquarema: Beirada z cf. Astrocaryum Carbonized fruits 3760; 1430 Brazil: Rio de Janeiro: Cabo Frio: 10 Scheel-Ybert, 1998

Boca da Barra >

cf. Astrocaryum Carbonized fruits & 2080 ? 70 Brazil: Rio de Janeiro: Arraial do 40 Scheel-Ybert, 1998 wood Cabo: Ponta da Cabeqa

Attalea butyracea Carbonized endocarps 7370 ? 130; Colombia: Tolima: Chaparral: 1800 Rodriguez, 1995 (Mutis ex L. f.) 5600 ? 90 El Limon: El Prodigio Wess. Boer

A. butyracea Carbonized endocarp & 6750-2450 Panama: Chiriqui: Casita de 700 Ranere, 1980; Smith, 1980 As Sheelea zonen- seed fragments Piedra sis

A. butyracea Carbonized endocarp & 6750-2450 Panama: Chiriqui: Trapiche 700 Ranere, 1980; Smith, 1980 As Sheelea zonen- seed fragments sis

A. butyracea Carbonized endocarp 3880? 80 Colombia: Boyaca: Puerto 150 Romero, 1996 fragments Boyaca: Marafial: Hacienda

Valparaiso A. butyracea Carbonized endocarp 2830 ? 70; Colombia: Santander: Cimitarra: 158 Piazzinni, 1995

fragments 2740 ? 70 Piamonte A. butyracea Carbonized endocarp 1550 Colombia: Tolima: Guamo: 402 Cifuentes, 1994 Relative cronology

fragments El Guamo with ceramic A. butyracea Carbonized endocarp 1150-950 Panama: Chiriqui: La Pitahaya <100 Ranere, 1980; Smith, 1980 As Sheeleazonen-

fragments (IS-3) sis



A. butyracea Carbonized endocarp 1120 + 60- Colombia: Cesar: Curumani: 62 Bernal, 1995 fragments 790 + 60 Los Serenos

A. butyracea Carbonized endocarp 850 ? 50 Colombia: Santander: Cimitarra: 150 Rodriguez & Restrepo, 1997 fragments San Juan: Finca La Marina

A. butyracea Carbonized endocarp 730 + 50; Colombia: Caldas: La Dorada: 176 Pefia & Morcote, 1997b fragments 420 ? 70 La Juana

A. butyracea Carbonized endocarp 590 60 Colombia: Cesar: Aguachica: 260 Bernal, 1996; Morcote, 1996 fragments San Francisco

A. butyracea Dry endocarps 330 A 60 Colombia: Tolima: Guamo: 402 Cifuentes, 1994 Colonial period La Chamba >

A. cf. butyracea Carbonized endocarps 565 + 115 Colombia: Caldas: La Dorada: 249 Morcote & Cavelier, 1999; As A. butyracea 0

Los Achiles Mendez & Escobar, 2000 In

cf. A. butyracea Phytoliths 1750-1150 Costa Rica: Puntarenas: La Pur- <100 Hoopes, 2000 As cf. Scheelea 0 ruja rostrata

Attalea cohune Mart. Carbonized endocarps 4450-1750 Belize: North Belize: Pulltrouser <50 Miksicek, 1983 As Orbignia > swamp cohune

A. cohune Endocarps 1650-1050; Belize: South coast: Wild Cane <50 McKillop, 1996 1050-450 Cay >

A. cohune Endocarps 1650-1050; Belize: South coast: Tiger Mound <50 McKillop, 1996 3 1050-450 >

A. cohune Endocarps 1650-1050; Belize: South coast: Frenchman's <50 McKillop, 1996 1050-450 Cay

A. cohune Endocarps 1650-1050; Belize: South coast: Pelican One <50 McKillop, 1996 t> 1050-450 Pot 0r

A. cohune Carbonized endocarp 730 A 50; Colombia: Caldas: La Dorada: La 176 Pefia & Morcote, 1997b 0 fragments 420 4- 70 Juana

A. cohune Carbonized endocarps 565 A 115 Colombia: Caldas: La Dorada: 249 Morcote & Cavelier, 1999; Los Achiles Mendez & Escobar, 2000

cf. A. cohune Phytoliths 2850-1050 Belize: North-central Belize: <50 Scott & Magennis, 1997 Phytoliths in dental "

Kichpanha calculus 1 Attalea cuatrecasana Carbonized endocarps 2190 60 Colombia: Choc6: Palestina I 25 Romero, 1994, 1996; Salgado

(Dugand) Hender- & Stemper, 1995 son, Galeano & R. Bemal

A. cuatrecasana Carbonized endocarps 1105 A 50 Colombia: Valle del Cauca: <50 Romero, 1994, 1996: Salgado Calima: Ordofiez I & Stemper, 1995

A. cuatrecasana Carbonized endocarps 990-885 Colombia: Choco: Palestina VIII 25 Romero, 1994, 1996 A. cuatrecasana Carbonized endocarps 815 A 35 Colombia: Choco: Palestina III 65 Romero, 1994, 1996



Table I (continued)

Age Altitude ?? Species Remains (B.P.) Site (m) Reference Notes

A. cuatrecasana Carbonized endocarps 665 + 30 Colombia: Choco: Palestina IV 25 Romero, 1994, 1996 A. cuatrecasana Carbonized endocarps 525 ? 35 Colombia: Choco: Palestina 25 Romero, 1994, 1996

vIII-C Attalea insignis Carbonized endocarp 9250 ? 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier

(Mart.) Drude fragments 9125 + 250; Pefia Roja et al., 1995; Morcote, 1998 1900+30- 385 + 50

A. insi,gnis Carbonized endocarp 1500 + 40; Colombia: Casanare: Aguazul: 525 Morcote, 1998 fragments 1080 + 40 Cuarto Unete: Finca Santa

Marta A. cf. insignis Carbonized endocarp 1560 + 100 Colombia: Casanare: Tauramena: 475 Penia, 1992; Lvpez et al., 1993;

fragments La Maporita Morcote, unpubl. data Attalea maripa Carbonized whole & 9250 + 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier

(Aubl.) Mart. fragmented seeds 9125 + 250; Pefia Roja et al., 1995; Morcote et al., ? 1900+ 30- 1996, 1998 > 385+50 z

A. maripa Carbonized seeds 2235 + 20 Colombia: Guaviare: San Jose del 240 Correal et al., 1990; Morcote, Guaviare: Guayabero I unpubl. data

Attalea microcarpa Carbonized seeds 11,145 + 135; Brazil: Para: Santarem: Monte 80 Roosevelt et al., 1996; Roose- Mart. 10,110 + 60 Alegre: Caverna da Pedra velt, 1999

Pintada Attalea racemosa Carbonized endocarp 9250+ 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier

Spruce fragments 9125 + 250; Pefia Roja et al., 1995; Morcote et al., 1900+ 30- 1996, 1998 385+ 50

A. cf. racemosa Carbonized & dry seed 1190-880 Colombia: Guaviare-Caqueta: 500 Van der Hammen & Castanio, fragments Serrania de Chiribiquete: 1995, Morcote, unpubl. data

Abrigo Selva A. cf. racemosa Carbonized & dry endo- 1065-1005 Colombia: Guaviare-Caqueta: 500 Van der Hammen & Castafho,

carp fragments Serrania de Chiribiquete: 1995; Morcote, unpubl. data Abrigo de Bemardo

Attalea spectabilis Carbonized seeds 10450 + 60; Brazil: Para: Santarem: Monte 80 Roosevelt et al., 1996 Mart. 10210+ 60 Alegre: Cavema da Pedra

Pintada Attalea (Scheelea) Carbonized endocarp 8040 + 390 Panama: Veraguas: Carabali 300 Cooke, 1992

fragments



Attalea (Scheelea) Carbonized endocarp 7000-4500 Panama: Cocle: Aguadulce <50 Cooke & Ranere, 1992 fragments

Attalea Phytoliths 6470:) 60 Colombia: Antioquia: Gomez 900 Castillo, 1998; J. Juan- Plata: El Moffo (021) Tressarras, pers. comm.

Attalea Phytoliths 3450-1950 Colombia Valle del Cauca: 1650 Piperno, 1985; As Scheelea; eleva- Calima: El Topacio Bray et al., 1988 tion too high for

an Attalea Attalea Carbonized endocarps 2380 + 60; Panama: Chiriqui: Baru: Cerro 2000 Linares et al., 1975 As Orbignya

1750-1550 Punta: Piti (BU-17) Attalea Endocarp fragments 2190 + 80; Panama: Cocle: Sitio Sierra <200 Cooke, 1992 Scheelea

1475 100 Attalea Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991

775+25 Abeja o Attalea Pollen 1500-1515 Colombia: Tolima: Coyaima: 350 Rodriguez, 1997b

Buenavista Attalea Carbonized endocarp 1120 + 60- Colombia: Cesar: Curumani: 62 Bernal, 1995

fragments 790+ 60 Los Serenos rd Bactris barronis L. H. Carbonized endocarps 525 + 35 Colombia: Choco: Palestina 25 Romero, 1994, 1996 >

Bailey VIII-C -H

Bactris coloradonis Carbonized endocarps 665 + 30 Colombia: Choco: Palestina IV 25 Romero, 1994, 1996 > L. H. Bailey

Bactris gasipaes Carbonized endocarps 2250-1650 Costa Rica: Puntarenas: Quebrada 20 Corrales & Mora Urpi, 1990 Kunth Seca >1

B. gasipaes Carbonized endocarps 2250-1650 Costa Rica: Lim6n: Severo 75 Corrales & Mora Urpi, 1990 r Ledezma o

B. gasipaes Carbonized endocarps 2190 + 60 Colombia: Choco: Palestina I 25 Romero, 1994, 1996; Salgado & Stemper, 1995

B. gasipaes Carbonized endocarps 1650-1050 Costa Rica: Alajuela: La Fabrica 1000 Corrales & Mora Urpi, 1990 r B. gasipaes Carbonized endocarps 1650-1050 Costa Rica: Limon: Bremen 40 Corrales & Mora Urpi, 1990 B. gasipaes Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 1

775 +25 Abeja B. gasipaes Carbonized endocarp 1080 + 40 Colombia:Casanare: Aguazul: 504 Morcote, 1998

fragments Finca Santa Marta B. gasipaes Carbonized endocarps 990-885 Colombia: Choco: Palestina VIII 25 Romero, 1994, 1996 B. gasipaes Carbonized endocarps 525+ 35 Colombia: Choco: Palestina 25 Romero, 1994, 1996

VIII-C B. gasipaes Endocarps 405 + 25 Colombia: Huila: San Agustin: 1730 Duque G6mez & Cubillos,

La Estaci6n 1981



Table I (continued)


B. gasipaes Carbonized endocarps 380 ? 50 Colombia: Meta: Acacias 523 Mora & Cavelier, 1987 cf B. gasipaes Wooden comb Not given Peru: Lima: Valle de Ancon <100 Rochebrune, 1879, cited in As Guilielma spe-

Towle, 1961 ciosa Bactris guineensis Carbonized endocarp 1120 + 60- Colombia: Cesar: Curumani: 62 Bernal, 1995

(L.) H. E. Moore fragments 790 + 60 Los Serenos Bactris major Jacq. Carbonized endocarps 3950-1650 Belize: North Belize: Cuello <50 Turner & Miksicek, 1984; Mik-

sicek, 1991 B. major Endocarps 1650-450 Belize: South coast: Wild Cane <50 McKillop, 1996

Cay B. major Endocarps 1650-450 Belize: South coast: Tiger Mound <50 McKillop, 1996 B. major Endocarps 1650-450 Belize: South coast: Pelican One <50 McKillop, 1996

Pot B. cf. major Carbonized endocarp 8040 + 390 Panama: Veraguas: Carabali 300 Cooke, 1992 O

fragments 0 Bactris setosa Mart. Carbonized endocarps 2260 ? 60- Brazil: Rio de Janeiro: Ze <200 Kneip & Pallestrini, 1987 >

1180 170 Espinho Bactris Pollen 10,000-5000 Colombia: Caldas: Villamaria: 1900 Rodriguez, 1997a >

Los Arrayanes Bactris Carbonized endocarps 4260 + 75- Brazil: Rio de Janeiro: Corondo <200 Carvalho, 1984

3010 + 80 Bactris Carbonized endocarps 4450-1750 Belize: Northem Belize: Colha <50 Miksicek, 1983 Bactris Wood charcoal 4450-1750 Belize: Northern Belize: San <50 Miksicek, 1983

Antonio: Albion Island Bactris Carbonized endocarps 4450-1750 Honduras: Copan 500-100 Miksicek, 1983 Bactris Seeds & pollen 1350-915 Colombia: Antioquia: San Rafael: 1000 Rodriguez, 1997c Relative chronol-

El Bizcocho ogy Bactris Carbonized endocarp 1120 ? 60- Colombia: Cesar: Curumani: 62 Bernal, 1995

fragments 790 + 60 Los Serenos cf. Bactris Carbonized fruits 5520 ? 120; Brazil: Rio de Janeiro: Cabo Frio: 9 Scheel-Ybert, 1998

3940 + 140- Forte 2240 + 70

cf. Bactris Carbonized fruits 4340 + 70, Brazil: Rio de Janeiro: Cabo Frio: 12 Scheel-Ybert, 1998 1800 Salinas Peroano

cf. Bactris Carbonized fruits 4160+ 190 Brazil: Rio de Janeiro: S Scheel-Ybert, 1998 Saquarema: Beirada



cf. Bactris Carbonized fruits 3760; 1430 Brazil: Rio de Janeiro: Cabo Frio: 10 Scheel-Ybert, 1998 Boca da Barra

cf. Bactris Carbonized fruits & 2080 ? 70 Brazil: Rio de Janeiro: Arraial do 40 Scheel-Ybert, 1998 wood Cabo: Ponta da Cabesa

Brahea dulcis (Kunth) Leaf 11,950-9150 Mexico: Puebla: Tehuacan 1200 Smith, 1965 Mart.

Butia capitata Carbonized endocarps, 5000 Uruguay: Rocha: Puntas de San <100 Del Puerto & Campos, 1998 (Mart.) Becc. phytoliths Luis

Chamaedorea Carbonized wood 4450-1750 Belize: North Belize: San Anto- <50 Miksicek, 1983 nio-: Albion Island

Chamaedorea Pollen 1565 ? 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 775 ? 25 Abeja m

Cryosophila staura- Wood charcoal 4450-1750 Belize: North Belize: Pulltrouser <50 Miksicek, 1983 As Cryosophila ar- o cantha (Heynhold) Swamp gentea T

Linden cf. Dictyocarywn Pollen 590 ? 160 Colombia: Magdalena: Santa 1300 Herrera de Turbay, 1985

lamarckianwn Marta: Buritaca 200 Elaeis oleifera Carbonized endocarp 6180 ?120; Panama: Cole: Aguadulce <50 Cooke & Ranere, 1992 >

(Kunth) Cortes fragments 5840 ? 100 3 E. oleifera Endocarp fragments 4210 + 90; Panama: Coclk: Aguadulce <50 Cooke & Ranere, 1992 >

2960 ? 80 - E. oleifera Carbonized endocarp 3880 ? 80 Colombia: Boyaca: Puerto 150 Romero, 1996

fragments Boyaca: Marafial: Hacienda >

Valparaiso 0 E. oleifera Carbonized endocarp 2920 ? 180 Panama: Veraguas: Carabali 300 Cooke & Ranere, 1992 0

fragments C

E. oleifera Carbonized endocarps 1550 Colombia: Tolima: Guamo: 402 Cifuentes, 1994 Relative chronol- > El Guamo ogy with ce- r

ramic cn2

E. oleifera Carbonized endocarps 1340 ? 85 Colombia: Sucre: San Marcos: 28 Rojas & Montejo, 1999 rm

Viloria E. oleifera Carbonized endocarp 1120 ? 60- Colombia: Cesar: Curumani: 62 Bernal, 1995

fragments 790 60 Los Serenos E. oleifera Carbonized whole & 850 80; Colombia: Antioquia: Puerto 130 Pefia & Morcote, 1997a

fragmented endocarps 700 ? 70 Nare: La Sierra: La Coquera: Finca Angostura

E. oleifera Carbonized endocarp 730 ? 50; Colombia:Caldas: La Dorada: 176 Pefia & Morcote, 1997b fragments 420 70 La Juana



Table I (continued)


E. oleifera Carbonized endocarps 565 + 115 Colombia: Caldas: La Dorada: 249 Morcote & Cavelier, 1999; Los Achiles Mendez & Escobar, 2000

Elaeis Phytoliths 3450-1950 Colombia: Valle del Cauca: 1650 Piperno, 1985a; Calima: El Topacio Bray et al., 1988

Euterpe oleracea Carbonized seeds 1000 ? 90 Brazil: Para: Ilha do Marajo: <30 Roosevelt, 1991, 1999 Mart. Teso dos Bichos

Euterpe cf. preca- Carbonized seeds 780 Colombia: Risaralda: Santa Rosa 1700 Rojas, 1997 toria Mart. de Cabal: El Jazmin

Euterpe Pollen 10,000-5500 Colombia: Caldas: Villamaria: 1900 Rodriguez, 1997a Los Arrayanes

Euterpe Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 t1

775 + 25 Abeja 0

cf Gastrococos Endocarps Not given Cuba: Santiago de Cuba: Cueva Probably Roumain, 1942 Cited as "corojo." i crispa (Kunth) del Purial <50 Identification >

H. E. Moore based on common name & > distribution of species, as given ? by Saakov (1970). Might be instead Acro- comia aculeata.

Geonoma deversa Carbonized seeds 2190 + 60 Colombia: Choco: Palestina I 25 Romero, 1994, 1996; Salgado (Poit.) Kunth & Stemper, 1995

Geonoma deversa Carbonized seeds 1105 + 50 Colombia: Valle del Cauca: <50 Romero, 1994, 1996; Salgado Calima: Ordoniez I & Stemper, 1995

Geonoma deversa Carbonized seeds 525 + 35 Colombia: Choco: Palestina 25 Romero, 1994, 1996 VIII-C

Geonoma Pollen 6000-5000 Colombia: Caldas: Villamaria: 1900 Rodriguez, 1997a Los Arrayanes

Geonoma Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 775 +25 Abeja

Geonoma Seeds & pollen 1350-915 Colombia: Antioquia: San Rafael: 1000 Rodriguez, 1997c Older date is rela- El Bizcocho tive chronology



Iriartea deltoidea Pollen 1565 4 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 As Iriartea

Ruiz & Pav. 775 ? 25 Abeja Lepidocarywn tenue Pollen 1565 + 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 As Lepidocarywn

Mart. 775 + 25 Abeja Manicaria saccifera Carbonized seeds 3550 + 65 Guyana: Hosororo Creek <30 Denis, 1992

Gaertn. M saccifera Pollen 1565 ? 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991

775 +25 Abeja Mauritiaflexuosa L. f Carbonized seeds 9250 ? 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier

9000; Peia Roja et al., 1995; Morcote et al., 1900 ?t30- 1996, 1998 385?50 cn

M.flexuosa Pollen 1565 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 O 775 25 Abeja

Mflexuosa Carbonized seeds 1560 100 Colombia: Casanare: Tauramena: 475 Peia, 1992; Lopez et al., 1993; La Maporita Morcote, unpubl. data

Oenocarpus bacaba Carbonized seeds 9250 140- Colombia: Caqueta: Araracuara: 200 Herrera et al., 1987; Cavelier

Mart. 9000; Pefla Roja et al., 1995; Morcote et al., 1900 ? 30- 1996, 1998 385 ?50 >

Oenocarpus batauw Carbonized seeds 9250 ? 140- Colombia: Caqueta: Araracuara: 200 Herrera, 1987; Cavelier et al.,

Mart. 9125 ?250; Pefa Roja 1995; Morcote et al., 1998 1900?30- > 385 50 0

0. bataua Carbonized seeds 3550 ? 65 Guyana: Hosororo Creek <30 Denis, 1992 O

0. bataua Carbonized endocarps 2800 Brazil: Minas Gerais: Santana <800 Resende & Prous, 1991 As Jessenia C

do Riocho bataua; proba- Cl bly a misidenti- r- fication.

0. bataua Seeds 1565 ? 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991 As Jessenia sp. 0 775 25 Abeja

0. bataua Carbonized & dry seed 1190-880 Colombia: Guaviare-Caqueta: 500 Van der Hammen & Castaino, fragments Serrania de Chiribiquete: 1995; Morcote, unpubl. data

Abrigo Selva 0. bataua Carbonized endocarps 665 + 30 Colombia: Choco: Palestina IV 25 Romero, 1994, 1996

Oenocarpus Carbonized seeds 9250 + 140- Colombia: Caqueta: Araracuara: 200 Cavelier et al., 1995; Morcote

mapora 9125 20+ ;0 Pefia Roja et al., 1998; Herrera et al., H. Karst. 1900?+ 30- 1987

385?+50



Table I (continued)


0. mapora Carbonized endocarps 2190 60 Colombia: Chocv: Palestina I 25 Romero, 1994,1996; Salgado & Stemper, 1995

0. mapora Carbonized endocarps 525 +35 Colombia: Choc6: Palestina 25 Romero, 1994, 1996 VIII-C

Paschalococos dis- Endocarps 820 + 40 Chile: Easter Island: Ana Okeke: <100 Flenley & King, 1984; Drans- perta J. Drans- Paike Peninsula field et al., 1984; Drans- field field, 1991

Phytelephas cf. ae- Phytoliths 3200 150- Ecuador: Esmeraldas: Cayapas: <200 Pearsall & DeBoer, 1996 As Phytelephas quatorialis Spruce 1540 + 80 C69

P. aequatorialis Phytoliths 1390-1030 Ecuador- Esmeraldas Cayapas: <200 Pearsall & DeBoer, 1996 R30

Phytelephas cf. tuma- Endocarp fragments 2300-1600 Colombia: Narino: Tumaco: 5 Patifio, 1999 As P. seemannii cana 0. F. Cook Las Marias 0

Prestoea cf. acumi- Carbonized seeds 1050-400 Colombia: Huila: Isnos: SA202 1800 Romano, 1997 nata (Willd.) z H. E. Moore

Pseudophoenix vini- Pollen 2130-1825 Dominican Republic: Rio <200 Vega, 1995 > fera (Mart.) Becc. Iguamo: El Caimito

Sabal Wood 4450-1750 Belize: Northern Belize: Pulltrou- <50 Miksicek, 1983 ser Swamp

Sabal Seeds 2150-1850 Belize: Northern Belize: Cerros <50 Cliff& Crane, 1989 Serenoa repens (Bar- Seeds Not given U.S.: Central & south Florida <200 Bennett & Hicklin, 1998

tram) Small cf. S. repens Mat 7290 U.S.: Florida: Windover <50 Field, 1996 Socratea exorrhiza Carbonized seeds 2190 + 60 Colombia: Choco: Palestina I 25 Romero, 1994, 1996

(Mart.) H. Wendl. S. exhorriza Carbonized seeds 990-885 Colombia: Choco: Palestina VIII 25 Romero, 1994, 1996 S. exhorriza Carbonized seeds 665 ?30 Colombia: Choco: Palestina IV 25 Romero, 1994, 1996 S. exhorriza Carbonized seeds 525 ?35 Colombia: Choco: Palestina 25 Romero, 1994, 1996

VIII-C Syagrus coronata Carbonized endocarps 11,000-8000; Brazil: Minas Gerais: Santana do <800 Resende & Prous, 1991

(Mart.) Becc. 2800 Riocho Syagrus cf. orino- Carbonized endocarps 1435 Colombia: Guaviare-Caqueta: 500 Van der Hammen & As Syagrus sp.

censis (Spruce) Serrania de Chiribiquete: Castano, 1995; Morcote, Burret Abrigo del Arco unpubl. data



Syagrus cf. orinocen- Dry seed fragments 520 Colombia: Guaviare-Caqueta: 500 Van der Hammen & As Syagrs sp. sis Serrania de Chiribiquete: Ab- Castanio, 1995; Morcote,

rigo del Valle de las Piramides unpubl. data Syagrus romanzoffi- Carbonized endocarps 680 _ 60 Brazil: Minas Gerais: Rio Que- 770 Anonymous, 1995

ana (Cham.) bra-Anzol (MG 30) Glassman

cf. S. romanzoffiana Carbonized endocarps 720 + 70 Brazil: Minas Gerais: Cobra 760 Anonymous, 1995 Cited as "sementes Coral (MG 29) de palmeira

jeriva" Syagrus Pollen 1565 _ 35; Colombia: Caqueta: Araracuara: 245 Mora et al., 1991

775 + 25 Abeja cf. Syagrus Carbonized fruits 5520 + 120; Brazil: Rio de Janeiro: Cabo Frio: 9 Scheel-Ybert, 1998 ZCt

3940 +140- Forte O 2240 + 70 T

cf. Syagrus Carbonized fruits 4340 + 70, Brazil: Rio de Janeiro: Cabo Frio: 12 Scheel-Ybert, 1998 '

1800 Salinas Peroano r

cf. Syagrus Carbonized fruits 4160 + 190 Brazil: Rio de Janeiro: 5 Scheel-Ybert, 1998 4 Saquarema: Beirada

cf. Syagrus Carbonized fruits 3760; 1430 Brazil: Rio de Janeiro: Cabo Frio: 10 Scheel-Ybert, 1998 > Boca da Barra >

cf. Syagrus Carbonized fruits & 2080 + 70 Brazil: Rio de Janeiro: Arraial do 40 Scheel-Ybert, 1998 wood Cabo: Ponta da Cabeca )

cf. Washingtonia ro- Containers from wood Not given Mexico: California: Baja Not given Lopez, 1996 > busta H. Wendl. California O

Unidentified Carbonized endocarp 14,700+ 195- Brazil: Rondonia: Chapada dos 500-1000 Miller, 1987 O 0 fragments 8930 + 100 Parecis: Abrigo do Sol C

(MT-GU-1) Unidentified Carbonized endocarps 11,000-8000 Brazil: Minas Gerais: Santana <800 Resende & Prous, 1991 >

do Riocho Unidentified Fruits 10,740 + 85- Brazil: Goias: Serranopolis: 500-700 Schmitz, 1987; Barbosa, 1992

5720+ 50 GO-JA-01O Unidentified Phytoliths 10,000-7000 Panama: Cocle: Cueva de Los 500-1000 Pipemo, 1985b

Vampiros Unidentified Carbonized endocarps 10,050 + 100 Colombia: Cauca: Morales: 1690 Gnecco & Mora, 1997 Misidentifications;

San Isidro not palms Unidentified Endocarp fragments 9670 + 150- Colombia: Valle del Cauca: 1600 Bray et al., 1988; Herrera

9300 + 100 Calima: Sauzalito et al., 1992 Unidentified Carbonized endocarp 9210+ 120 Venezuela: Amazonas: Puerto 51 Barse, 1990, 1995

fragments Ayacucho: Provincial



Table I (continued)


Unidentified Carbonized endocarp 9210 + 120; Venezuela: Amazonas: Puerto 51 Barse, 1990, 1995 fragments 9020 + 100- Ayacucho: Pozo Azul Sur 2

Unidentified Endocarp fragments 8750 + 160- Colombia: Valle del Cauca: 1650 Bray et al., 1988; Herrera 7830 + 140 Calima: El Recreo et al., 1992

Unidentified Carbonized endocarp 8240 + 160- Brazil: Minas Gerais: Lapa <800 Brian & Gruhn, 1978 fragments 7590 + 100 Pequena

Unidentified Pollen 8000-5500 Colombia: Caldas: Villamaria: 1900 Rodriguez, 1997a Los Arrayanes

Unidentified Seeds 6000-4000 Brazil: Para: Caverna dos <100 Roosevelt, 1998 Gavives H

Unidentified Seeds 6000-4000 Brazil: Rio Grande do Sul 500-1000 Rodriguez, 1992 Unidentified Palm mat 3490 + 120 Brazil: Minas Gerais: Sao Fran- <800 Machado, 1992 w

cisco: Gruta do Gentio II 0

Unidentified Phytoliths 3215;850 Ecuador: Napo: Lumu 450 J. Juan-Tresserras & A. H z

Sanchez, pers. comm. Unidentified Phytoliths 2950-418 Ecuador: Manabi: Valle del Rio 135 Pearsall, 1994b, 1994c

Jama: San Isidro Unidentified Carbonized endocarps 2800 Brazil: Minas Gerais: Santana <800 Resende & Prous, 1991

do Riocho Unidentified Pollen 2490 + 50 Colombia: Cauca: Timbiqui: 50 Patinio, 1988, 1992

La Cocotera Unidentified Seeds 2100-1600 Venezuela: Middle Orinoco: ca. 200 Roosevelt, 1998 Chronology of this

La Gruta site taken from Roosevelt (1980)

Unidentified Phytoliths 1750-1500 Ecuador: Esmeraldas: Cayapas: <200 Pearsall, 1994a Perdomo: C36

Unidentified Carbonized endocarp 1750-1150 Costa Rica: Puntarenas: La <100 Hoopes, 2000 Bactris sp. or fragments Purruja Elaeis sp.

Unidentified Phytoliths 1750-1150 Costa Rica: Puntarenas: La <100 Hoopes, 2000 Bactris gasipaes or Purruja Acrocomia acu-

leata Unidentified Phytoliths 1700-1450 Ecuador: Esmeraldas: Santiago: <200 Pearsall, 1994a

Selva Alegre: RIO



Unidentified Seeds 1600-1100 Venezuela: Middle Orinoco: Ron- ca. 200 Roosevelt, 1998 Chronology of this quin site taken from

Roosevelt (1980)

Unidentified Fruit fragment 1550-700 Ecuador: Manabi: Valle del Rio 135 Pearsall, 1994b, 1994c Jama: San Isidro

Unidentified Phytoliths 1550-418 Ecuador: Manabi: Valle del Rio 100 Pearsall, 1994b, 1994c Jama: El Tape

Unidentified Phytoliths 1450-950 Ecuador: Esmeraldas: Herradura: <200 Pearsall, 1994a C55

Unidentified Carbonized endocarps 1400-1150 Mexico: Tabasco: La Venta Rust & Sharer, 1998 z Unidentified Seeds & pollen 1350-915 Colombia: Antioquia: San Rafael: 1000 Rodriguez, 1997c Older date is rela- u

El Bizcocho tive chronology 0 'TI

Unidentified Implements for weaving 1250-1150 Colombia: Narifio: Guaitarilla: ca. 2600 Cardale de Schrimpff& Fal- Date estimated by I San Isidro chetti de Saenz, 1980 designs of asso- >

ciated ceramics & gold En

Unidentified Phytoliths 1150-650 Ecuador: Napo: Yurulpa 450 J. Juan-Tresserras & A. San- > chez, pers. comm. ,

Unidentified Phytoliths 950; 400 Colombia: Valle del Cauca: 1400 Piperno, 1985; Bray et al., >

Restrepo: Valle del Dorado 1988 Unidentified Implements for weaving 960 + 70 Colombia: Santander: Los Santos: 1310 Cardale de Schrimpff, Date associated >

Cueva de El Duende 1987 with funerary m cloth r-1

Unidentified Carbonized endocarp 730 + 50 Colombia: Caldas: La Dorada: 176 Peiia & Morcote, 1997b c0 fragments 420 +70 La Juana

Unidentified Spear thrower & darts 715 t 45 Colombia: Valle del Cauca: 1600 Von Schuler-Schomig, 1981 Probably Weutinia > Calima: Samaria

Unidentified Pollen 530 Colombia: Antioquia: San Rafael: 1000 Rodriguez, 1997c Los Farallones

. . _~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~rr




Table II A summary of palm species represented by remains at each of the known sites

in the New World

Archaeological site

Abrigo do Gruta Sol Boca C. da do

(MT- da Pedra Cobra Gentio GU-1) Beirada Barra Pintada Coral Corond6 Forte Guajara II

Species (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra)

Acromia aculeata Acrocomia hassleri Aiphanes cf. aculeata Astrocaryum chambira A. jauari A. malybo A. murumuru var. ciliatum A. standleyanum A. aff. triandrum A. vulgare * * Attalea cohune A. butyracea A. cuatrecasana A. insignis A. maripa A. microcarpa *

A. racemosa A. spectabilis *

Bactris barronis B. coloradonis B. gasipaes B. guinnensis B. setosa Butia capitata cf. Dictyocaryum lamarckianum Elaeis oleifera Euterpe oleracea E. cf. precatoria Geonoma deversa Iriartea deltoidea Lepidocaryum tenue Manicaria saccifera Mauritiaflexuosa Oenocarpus bacaba 0. bataua 0. mapora Paschalococos disperta Phytelephas aequatorialis Prestoea cf. acuminata Socratea exorrhiza Syagrus cf. orinocensis S. romanzoffiana *




Table II (continued)

Archaeological site

R. Que- Serra- Lapa Ponta brada Santana n6polis Teso Pe- da Arzol Salinas do Sernam- (GO- Dos Ze Easter

quena Cabe9a (MG 30) Peroano Riocho betiba JA-01) Bichos Espinho Island Acacias Abeja (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Bra) (Chi) (Col) (Col)

*

* *

*

* *





Archaeological site

Abrigo del

Abrigo Abrigo Valle de Buri- de del Las Pira- Abrigo Buena- taca El El El

Bernardo Arco mides Selva vista 200 Bizcocho Guamo Jazmin Species (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col)

Acromia aculeata Acrocomia hassleri Aiphanes cf. aculeata Astrocaryum chambira * A. jauari A. malybo A. murumuru var. ciliatum A. standleyanum A. aff. triandrum A. vulgare Attalea cohune A. butyracea * A. cuatrecasana A. insignis A. maripa A. microcarpa A. racemosa * * A. spectabilis Bactris barronis B. coloradonis B. gasipaes B. guinnensis B. setosa Butia capitata cf. Dictyocaryum lamarckianum * Elaeis oleifera * Euterpe oleracea E. cf. precatoria * Geonoma deversa Iriartea deltoidea Lepidocaryum tenue Manicaria saccifera * Mauritiaflexuosa Oenocarpus bacaba 0. bataua * 0. mapora Paschalococos disperta Phytelephas aequatorialis Prestoea cf. acuminata Socratea exorrhiza Syagrus cf. orinocensis * * S. romanzoffiana





Archaeological site

Finca El Finca Finca Santa Hacienda

Morro El Angos- La Marta Val- La La La La La Los (021) Prodigio tura Marina (CC2) paraiso Chamba Cocotera Estaci6n Juana Maporita Achiles (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col)

*

* * * *

* *

* *

* * * * * *

* *





Archaeological site

Los Pale- Pale- Pale- Pale- Pale- Los Faral- Los Ordofiez stina stina stina stina stina

Arral- lones Serenos I I III IV VIII VIIII-C Species lanes (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col)

Acromia aculeata Acrocomia hassleri Aiphanes cf. aculeata Astrocaryum chambira A. jauari A. malybo *

A. murumuru var. ciliatum A. standleyanum *

A. aff. triandrum A. vulgare Attalea cohune A. butyracea * A. cuatrecasana * * * * * *

A. insignis A. maripa A. microcarpa A. racemosa A. spectabilis Bactris barronis *

B. coloradonis *

B. gasipaes * * *

B. guinnensis * B. setosa Butia capitata cf. Dictyocaryum lamarckianum Elaeis oleifera *

Euterpe oleracea E. cf. precatoria Geonoma deversa * * *

Iriartea deltoidea Lepidocaryum tenue Manicaria saccifera Mauritia flexuosa Oenocarpus bacaba 0. bataua *

0. mapora Paschalococos disperta Phytelephas aequatorialis Prestoea cf. acuminata Socratea exorrhiza * * *

Syagrus cf. orinocensis S. romanzoffiana





Archaeological site

Santa San Rosa

Pefia Pia- Fran- del El Roja monte cisco Espejo SA200 Viloria C 36 C 55 C 69 Tape Lumu R 10 (Col) (Col) (Col) (Col) (Col) (Col) (Ecu) (Ecu) (Ecu) (Ecu) (Ecu) (Ecu)

* * *

*

*

*

*

*

* *

*

*~~~~~~~~~~~





Archaeological site

Hoso- Valle Pozo San roro de Azul

R 30 Isidro Yuralpa Creek Anc6n Rocha Sur 2 Species (Ecu) (Ecu) (Ecu) (Guy) (Per) (Uru) (Ven)

Acromia aculeata Acrocomia hassleri Aiphanes cf. aculeata Astrocaryum chambira A. jauari A. malybo A. murumuru var. ciliatum A. standleyanum A. aff. triandrum A. vulgare Attalea cohune A. butvracea A. cuatrecasana A. insignis A. maripa A. microcarpa A. racemosa A. spectabilis Bactris barronis B. coloradonis B. gasipaes * B. guinnensis B. setosa Butia capitata * cf. Dictyocaryum lamarckianum Elaeis oleifera Euterpe oleracea E. cf. precatoria Geonoma deversa Iriartea deltoidea Lepidocaryum tenue Manicaria saccifera * Mauritia flexuosa Oenocarpus bacaba 0. bataua * 0. mapora Paschalococos disperta Phytelephas aequatorialis * Prestoea cf. acuminata Socratea exorrhiza Syagrus cf. orinocensis S. romanzoffiana




Fig. 1. Location of archaeological sites with palm remains in North and Central America. 1. Central Florida (USA); 2. Southern Florida (USA); 3. Baja California (Mex); 4. Bocas del Purial (Cuba); 5. Rio Iguamo, El Caimito (DR); 6. Boca de Yuma (DR); 7. Santo Domingo (DR); 8. Tehuacan (Mex); 9. Cerros (Bel); 10. La Venta (Mex); I1. Pulltrouser Swamp, Albion Island, Cuello (Bel); 12. Coxcactlan (Mex); 13. Colha (Bel); 14. Kichpanha (Bel); 15. Tikal (Gua); 16. Orlando's Jewfish (Bel); 17. Wild Cane Cay, Pelican One Pot, Tiger Mound, Frenchman's Cay (Bel); 18. Copan, Los Mangos (Hon); 19. Salitr6n Viejo, Guarabuqui, PC-22 (Hon); 20. Cerro Palenque (Hon); 21. Severo Ledezma (CR); 22. La Fabrica (CR); 23. Bremen (CR); 24. Quebrada Seca (CR); 25. Casita de Piedra, Trapiche, Barriles, Piti (Pan); 26. Carabali, Aguadulce, Vaca de Monte (Pan); 27. La Pitahaya (Pan); 28. Sitio Sierra, Cueva de Los Vampi- ros (Pan). Bel = Belize; Chi = Chile; Col = Colombia; CR= Costa Rica; DR= Dominican Republic; Ecu = Ecuador; Gua = Guatemala; Guy = Guyana; Hon = Honduras; Mex = Mexico; USA = United States of America; Pan = Panama; Uru = Uruguay; Ven = Venezuela.

Bactris gasipaes, Elaeis oleifera, and Oenocarpus bataua. These species meet all the above- mentioned criteria. Some of these species, such as Acrocomia aculeata, have been used since 11,200 B.P., and all of them continue to be intensively used today.

V. Discussion

For most of the references studied it is not clear whether dates cited were based on the palm remains themselves or on other kind of remains at the site. Except for a few studies (e.g., Roosevelt et al., 1996; Morcote & Cavelier, 1999) in which dates were based on the palm remains themselves, it is possible that chronology, as cited in the papers reviewed, do not repre- sent the exact age of palm use. The error, however, is probably only of the order of a few decades.

The predominance of endocarps at the studied archaeological sites is probably the result of both cultural and environmental conditions. Cultural conditions include the selection, proc- essing, and consumption of oil-rich fruits and the subsequent disposal of endocarps. Some In- dian groups still use these endocarps as fuel in stoves or roast them for facilitating the extraction of the nuts or of the weevil larvae that develop inside (Romero, 1994; Cabrera et

al., 1999). Environmental conditions include the anaerobic environment of submerged ar-




0 O

29

31~~~~~~~17

4 56

51 *53

056 5 8~~~~~~~~~~~~~~~~~~~~~5

60

61 ~~~~~~~~~62

63

1~~~~~~~~' ~~~~~~~64 65.*

66 6766

/ 0~~~~~~~~7

Ea1er slan . 7

Fig. 2. Location of archaeological sites with palm remains in South America. 29. Buritaca 200 (Col); 30. Los Serenos (Col); 31. Viloria (Col); 32. San Francisco (Col); 33. Hosororo Creek (Guy); 34. El Morro (021) (Col); 35. Finca Angostura, Hacienda Valparaiso (Col); 36. Piamonte, Finca La Marina (Col); 37. Los Farallones, El Bizcocho (Col); 38. Santa Rosa del Espejo (Col); 39. La Juana, Los Achiles (Col); 40. Los Arrallanes (Col); 41. Pozo Azul Sur 2, Provincial (Ven); 42. Finca Santa Marta (CC2), La Maporita (Col); 43. El Jazmin (Col); 44. Palestina (Col); 45. Ord6ffez I (Col); 46. El Guamo, La Chamba, Buenavista (Col). 47. El Prodigio (Col). 48. Acacias (Col); 49. San Isidro, Samaria, Valle del Dorado, El Recreo, Sauzalito, El Topacio (Col); 50. Guayabero I (Col); 51. La Cocotera (Col); 52. La Estaci6n (Col); 53. Abrigo Selva, Abrigo de Bernardo, Abrigo del Valle de Las Piramides, Abrigo del Arco (Col); 54. Perdomo, Herradura, C69, Selva Alegre, R30 (Ecu); 55. San Isidro, El Tape (Ecu); 56. Abeja (Col); 57. Pefia Roja (Col); 58. Lumu, Yuralpa (Ecu); 59. Teso dos Bichos, Guajara (Bra); 60. Caverna da Pedra Pintada (Bra); 61. Valle de Anc6n (Peru); 62. Abrigo do Sol (Bra); 63. Serran6polis (Bra); 64. Gruta do Gentio II, Santana do Riocho, Lapa Pequena (Bra); 65. Cobra Coral, Rio Quebrada Arzol (Bra); 66. Co- rond6, Forte, Salinas Peroano, Boca da Barra, Ponta da Cabeca (Bra); 67. Sernambetiba (Bra); 68. Ze Es- pinho (Bra); 69. Beirada (Bra); 70. Rio Grande do Sul (Bra); 71. Easter Island (Chi); 72. La Gruta, Ronquin (Ven); 73. Las Marias (Col); 74. Rocha (Uru). See abbreviations for countries under Fig. 1.




Geonoma deversa Attalea cuatrecasana Bactris gasipaes Astrocaryum cf. triandrum Aiphanes cf. aculeata Astrocaryum malybo

Attalea cohune Manicaria saccifera

- _ _ _ - ~~~~~~~~Elaeis oleifera Attalea butyracea Bactris major Oenocarpus mapora Oenocarpus bataua Mauritia flexuosa Attalea racemosa

Attalea maripa --?? Attalea insignis

?-- U Astrocaryum murumuru -? Astrocaryum jauari

?- - - . Astrocatyum chambira _?_ Astrocaryum vulgare

?- Acrocomia aculeata

12,000 10,000 8,000 6,000 4,000 2,000 0

Years before present

Fig. 3. Chronology of records of palm remains in the Americas. Only species found at two or more sites are graphed. Gaps in the chronological record have been disregarded. The dashed line of some species corresponds to remains most likely assignable to those species.

chaeological sites (McKillop, 1996) and the dry environment of other sites (e.g., Morcote, unpubl. data). Furthermore, the hard nature of the endocarps and seeds favors their preservation.

For some of the most productive and widespread palms, archaeological data suggest that their distribution and abundance may have been the result of human use and dispersal. For other useful species, the lack of remains is intriguing-or just underlines the lack of appropriate studies. Below we discuss in detail some of the most interesting species for which remains have been recorded, or those for which their absence is remarkable.

A. ACROCOMIA ACULEATA

This palm is the most remarkable example of use, as represented by remains. It grows in the dry areas of the New World, from Mexico and the West Indies to northem Argentina (Henderson, 1995; Henderson et al., 1995), and produces fruits with oil-rich mesocarp and edible seeds (Galeano & Bemal, 1987; Lleras & Coradin, 1988) and sugar-rich sap (R. Ber- nal, pers. obs.; Janzen, 1983; Balick, 1990). It is used locally throughout its range today, and has been found, at least, at 29 sites from Mexico to Brazil. The oldest sites for Acrocomia are found in Santarem, northeastem Brazil (11,200 B.P.) (as Acrocomia sp., but most likely A. aculeata), Panama (8040 B.P.), and Mexico (6750 B.P.) (Fig. 4). Although there are more re-




15 * 7

19 X :0

67

Fig. 4. Location of sites where remains of Acrocomia aculeata have been found. Also included is the Cuban locality of Bocas del Purial, where remains could be Gastrococcos crispa or Acrocomia aculeata. See names of localities under Figs. 1-2.

cent sites in the intervening areas, this decline in the age of the oldest dates known suggests a northward migration in the use of this species, from some dry area in South America. There are even good reasons to think that the palm itself, not just its use, was dispersed during that time by humans: First, the fruits of Acrocomia aculeata, unlike those of other oil-producing palms, have an abundant mesocarp that can be consumed directly, without a long and time- consuming oil-extraction process. Second, the thick, brittle exocarp offers good protection for the mesocarp and can be easily peeled off when necessary. Third, the fruits do not ferment quickly, thus allowing their consumption for a period of several weeks. Thus, it would seem a good choice for migrating people to take fresh fruits of Acrocomia for eating first the fleshy and oily mesocarp during travel and then the seed (Levi-Strauss, 1950; Janzen, 1983), or dis- carding the intact nut, if cracking the endocarp was not an easy option.

The ecology of this species, which thrives easily in disturbed areas and is favored by fires, would contribute to its wide dispersal by humans. Human dispersion of Acrocomia aculeata has been discussed by Janzen (1983), Scariot (1988), Kahn and Moussa (1995), and Pipemo and Pearsall (1998). Janzen (1983) suspected that the species (as A. vinifera) was introduced by Amerindians into Costa Rica; Pipemo and Pearsall (1998) have suggested that it (as




A. mexicana) was taken from Mexico to Panama. Dispersal from South America to Central America seems more likely, not only because of the northward decline in the oldest dates of remains but also because the genus itself is probably of South American origin, as suggested by the fact that its other known species, Acrocomia hassleri, is restricted to cerrado areas in Brazil (Henderson, 1995).

B. BACTRIS GASIPAES

Pejibaye, Bactris gasipaes, is the only palm domesticated in America (Clement, 1992). This species is found throughout the neotropics, from Nicaragua to the Amazon Basin, and its large, mealy, protein-rich fruits are an important foodstuff for numerous aboriginal peoples throughout its range (Anonymous, 1975; Mora Urpi, 1984; Clement, 1988). Ethnohistorical data indicate that its use was widespread at the time of the European conquest, and it was so important to aboriginal peoples that the Spanish conquerors often cut down the trees as a strat- egy to subdue Indian groups (Patifno, 1963).

The place of origin of pejibaye has been the source of a long debate (Spruce, 1871; Huber, 1904; Burret, 1934; Mora Urpi, 1984, 1999; Clement, 1988, 1992; Clement et al., 1989), and both the westem and eastem foothills of the Andes, as well as the Cauca River Valley of Co- lombia, have been postulated as the place of domestication. Clement et al. (1989) and Clem- ent (1992) have favored southwestem Amazonia as the place where domestication probably took place, a hypothesis supported by a recent cladistic analysis of Bactris gasipaes and its closest relatives (Ferreira, 1999). However, the archaeological evidence does not seem to support this view; instead, it seems to support a northem, Andean or trans-Andean place of domestication. The oldest archaeological remains known for this species (carbonized endocarps) date back to 2250-1650 B.P. for two sites in the lowlands of Costa Rica (Corrales & Mora Urpi, 1990) and 2190 ? 60 B.P. for one site in the Pacific lowlands of Colombia (Ro- mero, 1995). The oldest records for the Amazon lowlands correspond to endocarp fragments found at Aguazul, in eastem Colombia, dated 1080 ? 40 B.P. (Morcote, 1998), and to pollen grains identified at Abeja (Araracuara), on the Caqueta River of Colombia, dated 775 ? 25 B.P.

(Mora et al., 1991). These records are ca. 1 100-1400 years more recent than the trans-Andean sites (Fig. 5). Thus, archaeological data, though scarce, suggest that introduction of pejibaye to Colombian Amazonia was a more recent event. On the other hand, the absence of this palm on the lower Amazon at the time of the European conquest (Patifno, 1963) and the absence of remains at the archaeological sites studied in detail in that area (Roosevelt, 1991, 1999; Roo- sevelt et al., 1996) do not support an origin in the upper Amazon Basin. Migration from the Andean foothills toward eastem Amazonia via the Amazon and its tributaries was intense (Meggers 1976; Lathrap, 1970, 1977), and it is difficult to believe that, if pejibaye originated in southwestem Amazonia earlier than 2250 B.P., it would not have descended the Amazon in the following 1700 years but instead crossed the Andes, moved along the Pacific coast up to Costa Rica, and then crossed the isthmus to the Atlantic lowlands.

If we accept that pejibaye was not domesticated in Amazonia, the most likely place is not Central America, even though the oldest archaeological remains (2250 B.P.) come from Costa Rica. The wild relative of pejibaye, Bactris gasipaes var. chichagui (= B. macana) (Hender- son, 2000) grows along the Andes from Venezuela to Bolivia and does not grow in Central America (Henderson et al. 1995, Henderson, 2000; but see Mora Urpi, 1999). The second- oldest site, Palestina, on the Pacific lowlands of Colombia (2190 ? 60 B.P.) makes better sense. This site is close to the Cauca River Valley, where the wild var. chichagui was once common (see, e.g., Karsten, 1857) and still grows in some places (R. Bemal, pers. obs.). The




35_i 21

45*

Fig. 5. Location of sites where remains of Bactris gasipaes have been found. See names of localities under Figs. 1-2.

Cauca Valley and the Pacific lowlands of Colombia had an intense trade in pre-Columbian times (Bray et al., 198 1), as had the northern Pacific coast of Colombia (north of Cape Corri- entes) with Panama (Reichel-Dolmatoff & Dussa'n, 196 1; Bray, 19 84). Detailed archaeological studies in the middle and upper Amazon Basin, as well as in other areas in the neotropics, are required to shed light on this issue.

C. OENOCARPUS BATAUA

This tall, oil-producing palm is widespread and very abundant throughout the Amazon Ba- sin, the Pacific lowlands, and the Magdalena River of Colombia and reaches eastern Panama. Its fruits are used to obtain a protein-rich beverage and a high-quality oil, much appreciated wherever the palm grows (Balick & Gershoff, 198 1; Balick, 1986). Furthermore, virtually all parts of the palm are used for some purpose (Balick, 1986; Galeano, 199 1). Studies of the Nu- kak, a nomadic group in the Colombian Amazon, have shown that human groups play a sig- nificant role in the dispersal of this species (Politis, 1996; Morcote et al., 1998; Cabrera et al., 1999). Archaeological evidence shows a strong chronological gradient in the use of this palm, from the middle Caqueta' River in Colombia (9250 B.P.), northeast to Guiana (3550 B.P.) and west to the Pacific lowlands of Colombia (665 B.P.) (Fig. 6). The middle Caqueta' River is the




.6. .13

45 4~~~~~~~~~~~~~~~~~~~~6

Fig. 6. Location of sites where remains of Elaeis oleifera (triangles) and Oenocarpus bataua (circles) have been found. See names of localities under Figs. 1-2.

area with the highest diversity of species of Oenocarpus (Bemal et al., 1991), and it is probably the center of diversification for the genus. Thus it seems reasonable to think that the use of Oenocarpus bataua began at a very early date in this area and spread from there in all direc- tions throughout the Amazon and other humid lowlands. Although dispersal of this palm is also effected by several animal species, including the oilbird (Steatornis caripensis) (Snow & Snow, 1978) and the capuchin monkey (Cebus apella) (Stevenson et al., 1991), the role of humans may have been decisive to its present distribution and overall abundance.

D. ELAEIS OLEIFERA

This oil-producing palm, closely related to the African oil palm, has a disjunct distribution in the neotropics, with isolated populations in Central America, the lower Atrato and the Mag- dalena River Valleys in Colombia, and several scattered sites in the Amazon Basin (Hender- son, 1995; Henderson et al., 1995). Both the mesocarp and the seed of this palm yield edible oil, and the species is believed to have been distributed in the Amazon by human groups (Balee, 1989). Although palm groves of this species have been associated with anthropogenic black soils (terra preta) in the Amazon Basin (Balee, 1988, 1989), archaeological remains have apparently not been found in that region. All known remains for this species come from




Panama and northern Colombia. The oldest site (6180 B.P. lies in Panama, and more recent sites are distributed along the Magdalena River Valley in Colombia (Fig. 6), suggesting a later introduction into this area.

E. ATTALEA BUTYRACEA

This variable and widespread species is one of the most massive palms in the neotropics. Distributed from Mexico to Brazil and Bolivia, it grows mostly in dry forests, sometimes reaching wet forests. Its mesocarp and seed produce edible oil, which is used in many areas where the palm grows (Standley & Steyermark, 1958; R. Bernal, pers. obs.). The palm is also used for its leaves, and wine is produced from its sweet sap (Bernal, 1992). Although the thick and hard endocarps of this species are easily preserved, they are not frequent at the archaeological sites reviewed. All of the records come from Panama and the Magdalena River Valley in Colombia, where the species has been used since the mid-Holocene.

F. MAURITIA FLEXUOSA

This palm, one of the most abundant and useful ones in South America today (Galeano, 1991; Henderson, 1995), was also extensively used in the past, as recorded by ethnohistorical data (e.g., Gumilla, 1745). However, remains of Mauritia have been recorded only at three archaeological sites, one of them represented only by pollen. This is probably due in part to the very specific ecological requirements of the palm, which grows in swamp areas; but in part it may be also due to the focus of the studies, which have not addressed plant remains in depth. As a matter of fact, the experience of one of us (GM) shows that remains of Mauritia seeds are easily identified even as minute fragments. Two useful palms with woody endocarps are note- worthy, because they have not been recorded at archaeological sites: the coconut (Cocos nucifera), and the coco cumbe (Parajubaea cocoides).

G. COCOS NUCIFERA

The origin of the coconut was the subject of a long debate (see the review in Child, 1964), although now it is mostly accepted that it originated in the western Pacific (Harries, 1978, 1995). Wherever it may have originated, there seems to be no doubt that the coconut grew wild and was cultivated on the Pacific coast of Panama and Colombia at the time of European contact (Cook, 1910; Patiinio, 1963; Zizumbo-Villarreal & Quero, 1998). When and how it may have arrived in this part of the continent is unknown, but Ward and Brookfield (1992) have shown that taken by sea currents, the coconut could hardly have reached America. Ar- chaeological evidence could be a vital element in this debate. The woody endocarp of the coconut is an appropriate material to be preserved at archaeological sites in wet environments. It is therefore of great importance to undertake archaeobotanical studies on the Pacific coast of Colombia and Panama, in order to look for direct evidence of past use of this species and of its possible anthropic dispersal.

H. PARAJUBAEA COCOIDES

The coco cumbe, Parajubaea cocoides, is known only in cultivation in the highlands of Ecuador and southern Colombia, whereas the two other species in the genus Parajubaea grow wild in the Andes of Bolivia (Moraes & Henderson, 1990; Moraes, 1996). The nuts of Para- jubaea cocoides are highly appreciated, and the use and distribution of the palm suggest a




long history of human management. Therefore, it is interesting that no remains of this species have been recorded so far. We suggest three possible explanations for this: first, the kind of human use might not have contributed to the preservation of remains; second, the remains may have been overlooked by researchers; or third, vulcanism in the Ecuadorian Andes has hindered site discovery (Piperno & Pearsall, 1998). A fact that still needs an explanation in connection with archaeological remains of palms is why, although in South America more studies have been made in the Andean highlands, most palm remains have been found at lowland sites. It is true that most Andean palms do not produce edible or oil-rich fruits. However, many of them, like Wettinia, Dictyocaryum, and Aiphanes, which have diversified mostly in the Andes (Moraes et al., 1995), have hard woods that could be represented as implements. But such implements are scarce at Andean sites.

It is evident, from this review, that archaeobotany can shed light on many questions related to the past use, distribution, and domestication of palms. A greater emphasis on plant remains at excavations is required, as well as archaeological exploration in critical areas.

VI. Acknowledgments

A preliminary version of this review was presented at the annual meeting of the Society for Economic Botany in Aarhus, Denmark, on July 13-17, 1998. We are grateful to H. Balslev for inviting us to that meeting and for encouraging this work. We also thank S. Cano, I. Cavelier, A. Henderson, J. Juan-Treserras, and A. S'anchez for their help with references and other information, S. Mora, D. Pipemo and T. van der Hammen for their comments on the manu- script, and J. C. Pinz6n for drawing the maps.

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