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

Institubo de Ciencias Naturales

UniversidadNacional de Colombia
Apartado 7495, Bogota, Colombia

1. AbstractiResumen ................. 309

11. Introduction ................. 310
111. Materials and Methods ................. 311
1\'. Results .......... .................. 311
V. D iscussion ........................................................... 335
A. Acroconmia aculeata ................. 337
B. Bactris gasipaes ................. 338
C. Oenocarpus bataua ................. 340
D. Elaeis oleifera ................. 341
E. Attalea butyracea ................. 342
F. Mauritiaflex.uosa................. 342
G. Cocos nucifera ................................................. 342
H. Parajr,baeacocoides ............................................... 342
VI. Acknowledgments ................... 343
VIL. 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 re-
corded from remains, was particularly widespread after 9000 B.P. The predominant remains
are endocarps of Acrocomia, Attalea s.l., Astrocarunum, 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

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 THE BOTANICAL REVIEW

played an important role in the dispersal of some palm species in the neotropics. Human-
aided dispersal of Acrocomia aculeatafrom South America into Central America, and of Oe-
nocarpus bataua from northwestern Amazonia to other areas, is postulated. Archaeological
remains support the hypothesis thatpejibaye (Bactrisgasipaes)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 revisi6n 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 Rondonia, 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 aborigenes. La revisi6n apoya las hip6tesis segun las cuales los grupos humanos han
jugado un importante papel en la dispersi6n de algunas palmas en el neotr6pico. Se postula la
dispersi6n antropogenica de Acrocomia aculeatadesde Suramerica hacia Centroamerica y de
Oenocarpus bataua desde el noroeste de la Amazonia hacia otras areas. Los vestigios ar-
queol6gicos apoyan la hip6tesis de que el chontaduro (Bactrisgasipaes) fue domesticado en
los valles interandinos o en las tierras bajas aledanias 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 (Patifio, 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 ar-
chaeological 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 atten-
tion 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 re-
ports or university theses outside Colombia. Published references often mention palmsjust 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 identifica-
tions 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 ofthe 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 archaeo-
logical sites ranging from the southern United States to southem 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 Attalea)
or of a relatively thin endocarp (as in Elaeisoleifera or Bactrisgasipaes).
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.
Phytohiths 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 usec 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 ancd 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 generaAcrocomia,Attalea s.l., Bactris,Syagrus, Elaeis,
Astrocaryvni, and Oenocarpus.The species used in these genera have a high contents of edi-
ble oil, and most of them are widespread or locally abundant or highly productive. The five
best-represented palms at archaeological sites are Acroconiia aculeata, Attalea butyracea,

(Text continues on p. 335)

312 THE BOTANICAL REVIEW

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m O
+ v++- +* + - + +* +* *8 H+* * * * * oH * + o v
OC Co_
*, *
vo > vz oO^-
> uo Cr_ C C)')
CC0
v - _ CY -so - 6' a, -

C.) C

C, C ,C C C C ~~~ CO ~ ~~C
C C oC 5 -C-C:

j zi Q o

a ^Z O E t
C.) C.) C)Y C.)'CC -O C.i
324 THE BOTANICAL REVIEW

COCA 00

C.0'CC 0' - C-~~~~~~~~~~-

.0 0 C) C) a. -. C ' '0 M~

--- .0' 0 0 - -~~~~~~~~~~C

- v c r4cq
CA ..
~~ ~ ~ ~ ~ ~ ~
v
00c.o
0 'C'C
C

0'o.o~~~ ~ 0' '-

C)2 0' Os - 'A -~~~~~~~~~I

0' C.,~ ~ ~ ~ ~ ~ ~ ~~I

0.
LO. 0. 0R.u o o

o 0~~~~~~~~~~~~~
0 00 o A A' ' 0

+ CD0 ~ C

a, cq
~~ U
CA ~~
a,
~ . 2 ~~ ~ -
-
O
a,
o. o C-

In 0 .~~* C

CA
CA CA~qt CA z-g . 0 o-

0~~~~~~~~~~~~~

0
Cl~~~~~~~~~~~~u Co I., 9
'A ~~ ~ ~ ~ 4 0 ' AO Ct
0
C t
Cl'A00
C -
e~~~~~C- -l CA-
CA
- z ~~~~~~
C
0''0'A~r

CA
CA CA AC~~~~~~~~~~~~~~~~~~~~
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 325

N~~~~~~~~~~~~~~~~~~

1C
a

>] <
< ~ ~ ~ ~ ~ C C) N ._C)

afi E a; a; ~a c - ,,, XXXo ~~~~~~~~~~~~~

E C)b

_ ,- , ,a _ a a .a
S .G _ C X -
-D ~ ~ _

t-E4 <~~~~~~~~
E' _,U: Q > a' E - -

,~C ., ^ o~ o .
a - v n -= ,n a- C) C: a
~ _ 6 a e 6 2 ~ 2 ea2Om
..
=-.2,
0
c:_>c^=>n;r;
a cza' : a 3 <1: a Oa t e oU o CaE

;) ~ ~
_)C)C ma m mC) E m- ar: 3

>: a {w C)x,}a
^ -r
c e
~~Cia'~~~ a'c 4 ' C4
a.
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a
,

3
'C N B. b X~a'a< N -' Za' a'
C') + + + + + +;
= ='
-4
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+
= .
+ + --
.C a'.
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qNC . Z 25 00 -. 'I a' 'Ca'a

a o, a CCC~~~~~~~~~~~~~~~~~~~~~~lo
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Q~~~~~~~~~~~~~~~~~~~~~~~~~~t -8 E Ea :-=3ss
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C).
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C)r U
326 THE BOTANICAL REVIEW

C) C C )~~~~~~
C )C)~~

CIA
M P C?0 0C e a.
IC = C- C > - 0 C>

O' -0 -, N N C> Q

t~~~~~~~~~~~~~~~~~~~~~~' *

nE<m Vo . C )' ~
~~ o V
- O ~~V - VZ 00 v0 ' 0C
VV 0 v
>~ ~~~~0_% 0' '0.0 ,g= - C) C)

C) 0~~~~C
N O ON -t NI CI 0
E 0 0
>cLC 0I0.00
-c

.0JN
C) ,S )0 S C C) C >
C) XC)C UC X ; C) C) CZP
) C) C) CL

Mo CQ
no no ro > C) ) C)

S; . 1; C C) S;C G s, i;C) C) C) C; C C)

- VO
O-~~~~~~~~~~
CI
-O cO c0 O c O c O c0~~~~~~~~~~~r
O k.
k,
CI
r~~~~~~~t

~~~~CI 73
~~~~~~~~~~~~C)
C)
C)
72 72 -.
C)
M= c=ooo=
oocc

._ u n n r n 0. CL Un x rA CL cr

- C V V V V
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 327

:, ~~ ~ ~ aC,oZa ) -- c

x~~ - ~~~ a a t 0
0fl~

0~~~~

a' C' 00 ~~~~~~~~~~~~C

o~~~ CD -- _
0' a' a' ~00 rj~ s.. s-.V)

0) UL J Um C ) U C
F- E .22 01

wC~~

S Db~~~~C

O0 O O O - -- - '- O '3 -

-: a' c' a' 0)= u

15 t s C ;- r- 0'
72'2 72 -2'2 72'2 1*~ -2 .0

r C CC)
 328 THE BOTANICAL REVIEW

Table 11
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 11
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
Man icaria saccifera
Mauritiaflexuosa
Oenocarpus bacaba
0. bataua
0. mapora
Paschalococosdisperta
Phytelephas aequatorialis
Prestoeacf. acuminata
Socratea exorrhiza
Syagrus cf. orinocensis
S. romanzoffiana *
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 329

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

* *

* *

*
 330 THE BOTANICAL REVIEW

Table II (continued)
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
(Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col)
Species
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
Iriarteadeltoidea
Lepidocaryum tenue
Manicaria saccifera *
Mauritiaflexuosa
Oenocarpus bacaba
0. bataua *
0. mapora
Paschalococosdisperta
Phytelephas aequatorialis
Prestoea cf acuminata
Socrateaexorrhiza
Syagrus cf orinocensis * *
S. romanzoffiana
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 331

Table 11 (continued)
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)
*

* * * *

* *

* *
* * * * * *

* *

* *

* * * *
 332 THE BOTANICAL REVIEW

Table II (continued)
Archaeological site

Los Pale- Pale- Pale- Pale- Pale-

Los Faral- Los Ordofiez stina stina stina stina stina
Arral- lones Serenos I I III IV Vill VIIII-C
lanes (Col) (Col) (Col) (Col) (Col) (Col) (Col) (Col)
Species
Acromia aculeata
Acrocomia hassl,eri
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 *
ef Dictyocaryum lamarckianum
Elaeis oleifera
Euterpe oleracea
E. cf precatoria
* * *
Geonoma deversa
Iriarteadeltoidea
Lepidocaryum tenue
Manicariasaccifera
Mauritiaflexuosa
Oenocarpus bacaba
*
0. bataua
0. mapora
Paschalococos disperta
Phytelephas aequatorialis
Prestoeacf. acuminata * * *
Socrateaexorrhiza
Syagrus cf. orinocensis
S. romanzoffiana
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 333

Table 11 (continued)
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)

* * *
*
*

* *

*
334 THE BOTANICAL REVIEW

Table II (continued)
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. Dichyocaryum lamarekianum
Elaeis oleifera
Euterpe oleracea
E. cf precatoria
Geonoma deversa
Iriarteadeltoidea
Lepidocaryum tenue
Manicariasaccifera *
Mauritiaflexuosa
Oenocarpus bacaba
0. batasa *
0. mapora
Paschalococos disperta
Phytelephas aequatorialis *
Prestoeacf acuminata
Socrateaexorrhiza
Syagrus cf orinocensis
S. romanzoffiana
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 335

Fig. 1. Location of archaeological sites with palm remains in North and Central America. 1. Central
Florida (USA); 2. Southem Florida (USA); 3. Baja Califomia (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.

Bactrisgasipaes, Elaeis oleifera, and Oenocarpusbataua. These species meet all the above-
mentioned criteria. Some of these species, such as Acrocontia 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 re-
mains 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-
336 THE BOTANICAL REVIEW

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. Ord6fdez 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. Perla Roja (Col); 58. Lumu, Yuralpa (Ecu); 59. Teso dos Bichos, Guajara (Bra); 60. Cavema da Pedra
Pintada (Bra); 61. Valle de Anc6n (Peru); 62. Abrigo do Sol (Bra); 63. Serran6polis (Bra); 64. Gruta do
Gentio 11, 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. Semambetiba (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. I.
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 337

Geonoma deversa
Attalea cuatrecasana
Bactris gasipaes
Astrocaryum cf. tnandrum
Aiphanes cf. aculeata
Astrocaryum malybo
Attalea cohune
Manicaria saccifera
Elaeis oleifera
Attalea butyracea
Bactms major
Oenocarpus mapora
Oenocarpus bataua
Mauritia flexuosa
Attalea racemosa
Attalea manpa
Attalea insignis
Astrocaryum murumuru
- --- -
Astrocaryum jauari
Astrocaryum 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 spe-
cies corresponds to remains most likely assignable to those species.

chaeological sites (McKillop, 1996) and the dry environment of other sites (e.g., Morcote, un-
publ. 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-orjust underlines the lack of appropri-
ate 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 northern Argentina
(Henderson, 1995; Henderson et al., 1995), and produces fruits with oil-rich mesocarp and
edible seeds (Galeano & Bernal, 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, northeastern Brazil (11,200 B.P.) (as Acrocomnia sp., but most likely
A. aculeata), Panama (8040 B.P.), and Mexico (6750 B.P.) (Fig. 4). Although there are more re-
338 THE BOTANICAL REVIEW

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 Gastrococcoscrispa 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 Piperno
and Pearsall (1998). Janzen (1983) suspected that the species (as A. vinifera) was introduced
by Amerindians into Costa Rica; Piperno and Pearsall (1998) have suggested that it (as
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 339

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, Acrocomnia hassleri, is restricted to cerradoareas 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 (Patiflo, 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 eastern 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 southwestern 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 sup-
port this view; instead, it seems to support a northern, Andean or trans-Andean place of do-
mestication. 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 eastern 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. 1100- 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 (Patino, 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 eastern 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 southwestern 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, Bactrisgasipaes 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. Bernal, pers. obs.). The
340 THE BOTANICAL REVIEW

Fig. 5. Location of sites where remains of Bactris gasipaeshave 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., 1981), as had the northem Pacific coast of Colombia (north of Cape Corri-
entes) with Panama (Reichel-Dolmatoff& Dussan, 1961; Bray, 1984). Detailed archaeologi-
cal 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, 1981; Balick, 1986). Furthermore, virtually all
parts of the palm are used for some purpose (Balick, 1986; Galeano, 1991). 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
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 341

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(Bemnal et al., 1991), and it is proba-
bly the center of diversification for the genus. Thus it seems reasonable to think that the use of
Oenocarpusbataua 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 (Steatorniscaripensis) (Snow &
Snow, 1978) and the capuchin monkey (Cebus apella) (Stevenson et al., 1991), the role of hu-
mans 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
342 THE BOTANICAL REVIEW

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 archaeo-
logical 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 ofMauritia have been recorded only at three ar-
chaeological 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 Mauritiaseeds 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 nu-
cifera), and the coco cumbe (Parajubaeacocoides).

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; Patino, 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 co-
conut 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, Parajubaeacocoides, is known only in cultivation in the highlands of

Ecuador and southern Colombia, whereas the two other species in the genus Parajubaeagrow
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
 REMAINS OF PALMS AT ARCHAEOLOGICAL SITES 343

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 low-
land sites. It is true that most Andean palms do not produce edible or oil-rich fruits. However,
many of them, like Weltinia, 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. Sanchez for their help with references and other in-
formation, S. Mora, D. Piperno and T. van der Hammen for their comments on the manu-
script, and J. C. Pinz6n for drawing the maps.

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