Coconuts in The Americas

Bot. Rev.
DOI 10.1007/s12229-013-9121-z
Coconuts in the Americas
Charles R. Clement1,8 & Daniel Zizumbo-Villarreal2 &

Cecil H. Brown3,4 & R. Gerard Ward5 &
Alessandro Alves-Pereira6 & Hugh C. Harries7,8
1
Instituto Nacional de Pesquisas da Amaznia, Av. Andr Arajo, 2936, 69067-375 Manaus, AM, Brazil
2
Unidad de Recursos Naturales, Centro de Investigacin Cientfica de Yucatn A.C., Mrida, Yucatn, Mexico
3
Northern Illinois University, DeKalb, IL 60115, USA
4
University of West Florida, Pensacola, FL 32514, USA
5
Research School of Asia and the Pacific, Australian National University, Canberra, Australia
6
Genetics and Plant Breeding Post-graduate Program, Universidade de So Paulo, Escola Superior de
Agricultura Luiz de Queiroz, Piracicaba, SP, Brazil
7
Coconut Time Line, Weymouth, Dorset DT3 5NP, UK
8
Author for Correspondence; e-mail: cclement@inpa.gov.br; hugh.harries@gmail.com
# The New York Botanical Garden 2013
Abstract It has been clearly established that the Portuguese introduced coconuts to
the Cape Verde islands in 1499, and these supplied the Atlantic coasts and the
Caribbean in the 1500s. By contrast, early 16th century reports of coconuts on the
Pacific coast of Panama are controversial. Recent DNA analysis of modern coconut
populations there shows them to be similar to Philippine varieties, agreeing with
morphometric analysis. Hence, coconuts must have been brought by boat from the
western Pacific, but no archaeological, ethnobotanical or linguistic evidence for pre-
Columbian coconuts has been found. Thus, the most parsimonious explanation is that
coconuts were introduced to Panama after Spanish conquest, as supported by DNA
analysis and historical records of Spanish voyages. New collections along the Pacific
coast, from Mexico to Colombia, are increasing the sampling for genetic analysis, and
further work in the Philippines is suggested to test probable origins. Unless new
archaeological discoveries prove otherwise, the strong hypothesis of Philippine origin
should direct future research on the sources of American Pacific coast coconuts.
Resumen Los portugueses introdujeron el cocotero a las islas de Cabo Verde en 1499, y
este se distribuy a las costas del Atlntico y el Caribe. Sin embargo los registros del
cocotero en siglo XVI en la costa del Pacfico de Panam son polmicos. Los anlisis
recientes de ADN de poblaciones modernas de coco muestran que son similares a las
variedades Filipinas, lo que est de acuerdo con los anlisis morfo-mtricos previos. Por
lo tanto, el cocotero debe haber sido llevado en barco desde el Pacfico Occidental, pero
no hay evidencias arqueolgicas, etnobotnicas o lingsticas precolombinas. La
explicacin ms parsimoniosa es que fue introducido despus de la conquista espaola.
C.R. Clement et al.
Esto es apoyado por los anlisis de ADN y los registros histricos de los viajes espaoles
con cocos. Nuevas colectas a lo largo de las costas del Pacfico desde Mxico hasta
Colombia mejorarn el muestreo para el anlisis gentico, y se sugieren nuevos trabajos
en Filipinas para confirmar los orgenes precisos. A menos de que nuevos restos
arqueolgicos se encuentren que demuestren lo contrario, esta hiptesis puede orientar
nuevas investigaciones sobre los orgenes del cocotero en la Costa del Pacfico
americano.
Keywords Cocos nucifera . Molecular genetics . History . Archaeology . Linguistics .

Ethnobotany
Palabras claves Cocos nucifera . Gentica molecular . Historia . Arqueologa .

Lingstica . Etnobotnica
Introduction
Coconut is the iconic palm of beaches throughout the tropics. It was a major plantation
crop for much of the 19th and 20th centuries, and still provides an income to millions of
small farmers. It is an invaluable plant providing many of the basic necessities for
survival in traditional societies, especially in the southern and western Pacific, where it
will have a 21st-century role as a biofuel. It was possibly the first pan-tropical crop plant,
yet where it came from and how it was dispersed has long fascinated scholars. The most
hotly contested part of this discussion concerns the coconut reported by the first
European explorers on the Pacific coast of Panama in the early 16th century. Over
the past century the discussion has waxed and waned, and now new genetic
evidence suggests a relationship between the current tall population on the Pacific
coast of Panama and coconuts in the Philippines (Baudouin & Lebrun, 2009; Gunn
et al., 2011). The genetic analyses are compelling, but a single line of evidence is
seldom sufficient to convince the gamut of scholars. Thus it is worthwhile to review
all available evidence to try to answer the questions of how coconut traveled to
Panama, where it came from and when it arrived, or to suggest new hypotheses for
future research.
The renowned 19th century Swiss botanist, Alphonse de Candolle, is considered
the founder of modern crop biogeography because he proposed a multidisciplinary
methodology to identify crop origins and trace crop diffusions (1883). At a minimum,
this requires information from botany, especially patterns of variation, history, lin-
guistics and archaeology. During the early 20th century, the famous Russian genet-
icist, Nicolay I. Vavilov, expanded the patterns of variation to include genetics
(Vavilov, 1951), initially Mendelian and quantitative, and more recently molecular
genetics (Zeder et al., 2006).
This methodology, or parts of it, has been used to examine other candidates for
trans-Pacific human dispersal, notably sweet potato (Ipomoea batatas) and more
recently chicken (Gallus gallus). Several lines of evidence suggest that Polynesians
made voyages to and from the west coast of the Americas, carrying sweet potato
westwards to western Polynesia and chickens eastwards to coastal Chile (Jones
et al., 2011).
The evidence for sweet potatoes includes archaeology, ethnobotany, linguistics

and genetics, and seems reasonably complete. The sweet potatoes in Oceania appear
to have originated from two separate introductions from America (Ballard et al.,
2005). The earliest was to Yens ellipse area of eastern Polynesia by 900
1,000 years before present (BP) (Yen, 1974; Green, 2005: 467; Lebot, 2009: 94)
and thence to New Zealand and Hawaii. Linguistic and other evidence is cited by
Scaglion (2005: 3541) for the Gulf of Guayaquil, Ecuador, as the origin of these
sweet potatoes, which corresponds with the South American center of sweet potato
domestication (Roullier et al., 2011). The second wave of introductions came to
Papua New Guinea and western Melanesia from Indonesia after the Spanish voyages
from Mexico; these introductions came from the Mesoamerican center of sweet
potato domestication (Roullier et al., 2011). A study by Zhang et al. (2004)
questioned whether the Ecuadorian origin was not representative of both origins
and dispersals, and concluded that Mesoamerican sweet potatoes dominate Oceania.
Roullier et al. (2013) have now confirmed Yens (1974) hypothesis, making this the
first plant evidence for a Neotropical to Oceania dispersal.
A combination of archaeological, genetic and some linguistic evidence was re-
cently used by Storey et al. (2007, 2011a) to propose that the chicken had been
introduced into El Arenal, Chile, from Polynesia a century before European conquest.
They counter earlier questioning of the validity of the archaeological results of the
chicken remains in Chile (e.g., Gongora et al., 2008a, b). Although Acosta (1940)
mentions them in the Caribbean and Mexico, Patio (1970: 34, 35) considers that no
reliable historical record exists before Columbus took chickens on his second expe-
dition and disembarked them in 1493 as presents for local chiefs, which thus explains
Acostas record, but is post-conquest in that region. Storey et al. (2007) affirm that
Francisco Pizarro found chickens in Peru, but Patios (1970) documental analysis
does not agree.
Skeletal, artifact and non-material cultural evidence also suggests Polynesian
contacts with the Chilean coast (Matisoo-Smith & Ramrez, 2010; Ramrez-Aliaga,
2011). Sewn plank canoe-building technology, characteristic of Polynesia and Mi-
cronesia, found in small areas in southern California and southern Chile, also
suggests contact with the Pacific Islands (Jones et al., 2011; Ramrez-Aliaga, 2011).
The information available for coconut is much less complete, with new genetic
information and an historical record that has been questioned and defended by
different authors. No information on the archaeology and linguistics of coconut in
Panama has yet been published. Only a fraction of the coconut genetic variation in the
Philippines is present in Panama, suggesting a single small founder event (Baudouin
& Lebrun, 2009), while populations on the Pacific coast of Mexico contain consid-
erable Philippine variation, but also only a fraction of the Panama genetic variation
(Gunn et al., 2011). This lack of variation raises two possibilities: a direct introduc-
tion from the Philippines to Panama (Baudouin & Lebrun, 2009), or an introduction
via Mexico after European conquest (Harries, 1978). Previous work has already
shown that this founder event could not be explained by coconuts drifting on ocean
currents (Ward & Brookfield, 1992). We thus seem to have a reasonable answer to the
where it came from question, but the how it got there remains an open question,
although a trans-Pacific route is clear. An associated question is when, which raises
contrasting possibilities: a very early direct introduction from the Philippines to
C.R. Clement et al.
Panama, e.g., 2,250 BP, as postulated by Baudouin and Lebrun (2009); at a date
immediately before European contact (suggested by the extremely limited endemic
distribution reported at the time of European conquest); or over the Manila-Acapulco
route less than 450 years ago (Small, 1929; Hill, 1929; Merrill, 1954; Harries, 1971,
1978). These hypotheses can be examined with linguistic and archaeological evi-
dence, as well as with evidence from ethnobotany and history, which might help with
the how question.
We apply de Candolles and Vavilovs multidisciplinary methodology to the
question of coconut on the Pacific coast of Panama at the time of European conquest.
As our starting point, we summarize in greater detail the new genetic information
concerning its origin. We then examine the historical record for the period between
the discovery of the Pacific Ocean by Spaniards and the trans-Pacific trade organized
by the Spanish crown, and investigate the archaeological record, linguistic and
ethnobotanical evidence. Finding that these latter lines of investigation are not
fruitful, we re-examine the genetic and historical evidence, as well as five hypotheses
concerning the origin of coconut in the Americas that have appeared in this century or
so of debate: coconuts 1) originated in the Americas; 2) floated on ocean currents
from Oceania; 3) were carried by Polynesians; 4) were carried by unknown mariners;
and 5) were carried by Spanish galleons returning from the Philippines.
The Genetic Evidence
Two recent studies based on DNA evidence suggest that the origin of the reputedly pre-
Columbian coconuts in Panama is in Southeast Asia, most probably in the Philippines
(Baudouin & Lebrun, 2009; Gunn et al., 2011). Although the coconuts of the Pacific
coast of Mexico and of Panama came from the same general region, they are readily
distinguished using microsatellite markers (Baudouin & Lebrun, 2009). The origin of
the Mexican coconuts is known: They were available directly from the Philippines (as
well as some from other locations) starting in 1565 for a period of 250 years, when the
Spanish crown inaugurated and managed the Manila-Acapulco trade route (Fig. 1).
This trade route did not necessarily involve a large number of introductions nor a large
number of nuts at any one time, as the coconuts were carried for consumption by the
crew and passengers during a three-month journey. Any early germinators not used
could be planted, but once an initial planting came into bearing, it would soon supply
all future planting material on a year-round basis and no further introductions were
needed until commercial demand for planting material occurred at the start of the 20th
century. By contrast, the Panama Tall on the Pacific coast is characterized by markedly
reduced diversity (expected heterozygosity=0.324 in this Panama Tall versus 0.579 for
all Tall coconuts combined; observed heterozygosity=0.230 in this Panama Tall versus
0.480 for all Tall coconuts; Gunn et al., 2011). Such a reduction is the signature of a
bottleneck, i.e., a strong reduction of population size at some moment in the history of
a population, as is typical of a small founder event.
This founder event is clearly seen by examining individual microsatellite loci, such
as locus CnCir 2 (Fig. 2). Only two of the seven alleles found in Southeast Asia exist
in Panama and allele 222 represents 96 % of the total variation present in Panama
(Baudouin & Lebrun, 2009). Allele 222 is also the most frequent in Southeast Asia,
Fig. 1 Map of the Pacific basin with features relevant to coconut in the Americas. Insetslocations on the
Pacific coast of Panama where coconuts were reported shortly after European conquest; locations in the
Philippines mentioned in the text. Map prepared by the Cartographic & GIS Services, Australian National
University, Canberra, Australia
and its frequency is especially high in the Philippines (65 %). Its frequency decreases
progressively from Southeast Asia to Melanesia, Micronesia and Polynesia. Such a
tendency is observed at most of the 30 loci studied (data not shown; see Baudouin &
Lebrun, 2009; Gunn et al., 2011).
Baudouin and Lebrun (2009) created a similarity index to integrate the allelic
information across 30 microsatellite loci to compare different sets of possible source
regions for the alleles found in the Pacific coast Panama Tall (Fig. 2). The Philippines
variety is most similar to both this Panama Tall and to the Mexican Pacific coast
1 1
0.9 CnCir2-222 0.9
0.8 Similarity 0.8
Similarity Index
0.7 0.7
Allele frequency
0.6 0.6
0.5 0.5
0.4 0.4
0.3 0.3
0.2 0.2
0.1 0.1
0 0
Pan Mex Phi SEA Ind GuN Mar GuS NBr Van Mic Pol
Fig. 2 Allele frequency of microsatellite locus CnCir2, allele 222, and similarity index across 30
microsatellites to compare allelic diversity of the Panama Tall with possible source regions in the Pacific
Ocean basin. Pan-Panama Tall, Mex-Mexico, Phi-Philippines, SEA-Continental Southeast Asia, Ind-
Indonesia, GuN-North coast of New Guinea, Mar-Markham Valley in New Guinea, GuS-South coast of
New Guinea, NBr-New Britain, Van-Vanuatu, Solomon Islands, New Caledonia, Mic-Micronesia, Pol-
Polynesia. Redrawn from Baudouin and Lebrun (2009)
C.R. Clement et al.
coconuts, while the Polynesian island sources are the least similar, although they are
geographically closer. This is the genetic basis for the affirmation that both the
Panama and Mexican Pacific coast coconuts were introduced from the Philippines.
The limited similarity between the Philippines and this Panama Tall (0.475) is due to the
extremely small founder population of the Panama coconut, and possibly different selec-
tion pressures since introduction (Baudouin & Lebrun, 2009). However, the initial selection
pressures may have been uniform and closely related to trans-Pacific travel. The reduction
in population size probably occurred because most of the nuts were consumed before
arrival and the selection pressure was for the early germinators that survived to be planted,
since these would have been less preferred for consumption. Thus, only a small fraction of
the allelic diversity found in the Philippines is found in this Panama Tall, whereas the allelic
diversity found in Mexico is quite similar to that found in the Philippines (Fig. 3). The
similarity of Mexican and Philippine varieties is probably due to several introductions after
1565, especially to supply the toddy market in the 17th century (Zizumbo-Villarreal, 1996).
A Time Line of Spanish Contact with Coconut
The journal of Christopher Columbus (Cristobal Colombo) records that, on the 17th
of November 1492, when sailing near Puerto del Principe on the north coast of Cuba:
There was a beautiful meadow and many very tall palms. They found a very large
nut of the kind belonging to India, (Columbus, 1893: 80). Columbus thought he
had reached Asia and found coconuts, but he had misidentified the Royal palm
(Roystonia spp). Some 30 years later, palms also thought to be coconuts were
reported on the Pacific coast of Panama by Gonzalo Fernndez de Oviedo y Valds,
who was appointed as one of the official historians of the Indies (Spanish American
colonies) in 1523, after his return from Santo Domingo (todays Dominican Repub-
lic). He published a first Summary entitled Natural History of the Indies in 1526,
before returning to the Americas. Part of his General and Natural History of the
Indies was published in 1535, but the whole General History was only published in
18511855, edited by J.A. de los Rios y Serrano.
Although Oviedo was not a naturalist, his Natural History is considered an
important reference about the natural resources of the Americas. Throughout he makes
Fig. 3 Multi-locus allelic composition of the Philippine, Mexican and Panamanian varieties studied by
Gunn et al. (2011). The colors in the pie charts represent the groups identified by Structure analysis at K=5
and the size of the slice is the proportion of each group in the region. The admixed group represents hybrids
between the Indo-Atlantic origin and Pacific origin coconuts that may have resulted from hybridization on
the Caribbean coast of Panama with coconuts brought by the Spanish across the Atlantic Ocean. Redrawn
from Gunn et al. (2011)
clear that only part of the information was obtained personally, and part was obtained
by interviews and correspondence, as the Spanish crown demanded information from
all explorers who received crown support. Before and during the period that Oviedo
was active as historian, information was flowing into the Iberian Peninsula at a great
rate, as Spanish and Portuguese explorers visited new parts of the New and Old
Worlds. It is important to remember that they did not explore the same areas because
of the Treaty of Tordesillas, which effectively divided the non-European world
between Spain and Portugal.
The case of coconut is emblematic of this flow of information, as Vasco de Gama
was the first to bring coconut to Portugal in 1499, after leaving seedlings in the Cape
Verde islands, where they grew to reproductive age and were distributed to the
Americas (Harries, 1977). The conquistadors who sailed to the New World in the
first decades of the 16th century knew that coconuts were special because in 1501
King Manuel of Portugal had written a letter to Ferdinand and Isabella of Spain,
extolling the prime value of coconuts as a source of drinking water and cordage for
sailing ships (cited by Harries, 1977; Patio, 1963, 2002). However, the first com-
plete information available in Iberia about coconut uses in Asia was published by
Ludovico di Varthema (1510), based on his travels in Asia between 1501 and 1508.
His report would have been read by Magellan before setting out in 1519 to navigate
the globe and by Oviedo before going to Darien (Caribbean coast of Panama) in
1521. Hence, as Patio (1963, 2002) emphasizes, Oviedo and others of the period
used information from Asia and the Americas in their reports without specifying the
origin of each detail, a common occurrence before modern citation methods were
developed. What has not previously been well recognized is that no Spanish explorers
or historians had ever seen growing and fruiting coconut palms because the Treaty of
Tordesillas prevented them from sailing to Asia, although some may have seen the
nuts in Lisbon or other Iberian cities.
Zizumbo-Villarreal & Quero (1998) analyzed Oviedos Summary and General
History to determine if coconut was present in Panama at the time of European
conquest. Patio (1963; 2002) analyzed both of these documents and the origins of
the information that Oviedo used to prepare his Summary and General History,
although he recognized that many documents have been lost and some were
unavailable to him even after 40 years of effort. Both Patio and Zizumbo & Quero
cautiously accept that Oviedos reports are correct. Based on Patios analysis, this is
the time line for the early Spanish references to coconut in Panama.
1492Discovery of New World

1494Treaty of Tordesillas divided the world outside Europe between Portugal
(eastwards) and Spain (westwards), which effectively prevented Spanish mari-
ners from sailing to Asia (until 1580).
1501King Manuel of Portugal wrote about coconut in detail in a letter to
Queen Isabella and King Ferdinand of Spain.
1513Vasco Nez de Balboa crossed the Isthmus of Panama and discovered the
Pacific Ocean. The expedition explored only a small area and returned without
mentioning coconut, although numerous documents of this expedition have been lost.
15141515Alonso de la Puente (royal treasurer in Darien, on Panamas Carib-
bean coastheard of coconut) and Rodrigo Colmenares (ship pilot and interpreter
C.R. Clement et al.
saw coconut) visited the Pacific coast of Panama and were interviewed by Pedro
Mrtir de Anglera, another official crown historian, in late 1516 in Spain.
Angleras report was first published in 1530 (translated in 1944), but Oviedo knew
Anglera and had access to his report, which explains why this information is in the
Summary. The explorers Puente and Colmenares reported on the coconut (that they
thought to be the same fruit as cultivated in Calicutthe common term for India in
the first decades after European contact) and go on to state that it was cultivated on
some of the islands in the Gulf of Panama. The fact that they thought it to be the
same as the coconut from India is significant, but does not mean that it was indeed
coconut.
Anglera states that coconuts were observed in three places along the Pacific
coast: Chimn, to the southeast of the current City of Panama; Nat, to the west of
the City of Panama; and Burica, further to the west at the frontier with modern Costa
Rica (Fig. 1, Panama insert). Chimn is thought to be the first place that Europeans
saw coconut and Patio even suggests that Balboas expedition may have seen it
there also. Nat was reputed to have an abundance of coconuts, reported by the
natives to germinate and grow spontaneously along the shore, although they also
reported young plants were transplanted. Anglera also reports that Some think that
sea currents bring the seeds of these trees from unknown regions.
1519Gaspar de Espinosa y Luna (explorer, participated in the founding of the
City of Panama in 1519) explored west of the City of Panama as far as the point
of Burica, where he reported Many beautiful and large mameys (Pouteria
zapota) and many palms with the large coconuts, It is not clear exactly when
Anglera obtained this information from Espinosa. Patio suggests that Espinosa
may only have realized the importance of his observation after learning about
coconut in Asia.
1524Francisco Pizarro Gonzlez (mayor of the City of Panama 15191524)
led an expedition that explored the South American coast from the City of
Panama to the northern limits of Colombia. Near the Cape of Corrientes
(Fig. 1), they reported a large quantity of coconuts at La Candelaria Bay,
which are reported in Oviedos General History, but not in the Summary.
1526Juan de Cabezas (pilot) discovered Cocos Island (Fig. 1) and provided
information used by Oviedo in the full General History.
1539Alvaro de Guijo (resident in the City of Panama) sent nuts to Hernn
Corts de Monroy y Pizarro (conqueror of Mexico), along with advice about how
to plant them. In his letter that accompanied them, Guijo wrote: I have heard
that you do not have this fruit we call cocos, so I sent a boat to a place along the
coast to collect some, and I send you two dozens of ripe fruits. Some of the riper
ones can be sown by placing half the fruit in the ground. If you like them, I can
send more another time, as well as some already germinated. This letter is cited
in Bruman (1947), but Zizumbo-Villarreal & Quero (1998) caution that this
possible introduction to Mexico may not have been successful. In fact, Hernn
Corts left for Spain in 1541 and never returned to Mexico.
1549Coconuts from the Cape Verde islands (Portuguese) planted in Puerto Rico
(Spanish), attributed to Diego Lorenzo, canon of Cape Verde (Bruman, 1944).
1553Coconuts from the Cape Verde islands planted at Bahia, Brazil (Bruman,
1944), possibly the first coconuts on continental America.
1565Philippines to New Spain (Mexico) return trip was accomplished inde-

pendently by Alonso de Arellano and Andrs Urdaneta, and the Manila-
Acapulco commercial route was established by 1571 and continued until 1815
(Fig. 1). Coconuts, carried for consumption by passengers and crew, became a
regular item but were not always mentioned in the ships manifests.
1569Alvaro de Mendaa introduced coconuts from the Solomon Islands to
Colima, Mexico (Sevilla de Rio, 1974).
1580Treaty of Tordesillas became invalid when Portugal was ruled by Philip II
of Spain.
The Archaeological Evidence
The central Pacific coast of Panama has a long cultural history, with human activities
present in the archaeological record since 11,000 years before the present (BP)
(Piperno & Pearsall, 1998). By the late Pre-Ceramic Period (70005000 BP), horti-
culture started to become important, expanding rapidly through the Early Ceramic
Period (50002500 BP) until most subsistence was horticultural by the Middle
Ceramic Period (25001500 BP). The Ceramic Period extended until European
Conquest (500 BP). Nonetheless, subsistence still included gathering, especially of
palm fruits and other useful plants.
The macro-and micro-archaeological records show that palms were very important
to the subsistence of the Native Americans in the Gran Chocl (central Panama) and
Gran Chiriqu (western Panama) regions. The macro-archaeobotanic remains from
these regions include Acrocomia aculeata (Jacq.) Lodd. ex Mart., Attalea butyracea
(Mutis ex L.f.) Wess. Boer, Bactris major Jacq., Elaeis oleifera (Kunth) Corts and
Astrocaryum sp. (Dickau, 2010). All of these species are members of the Cocoseae
tribe within the Arecaceae, but Acrocomia, Astrocaryum and Bactris are spiny and
like Elaeis have fruits that are morphologically clearly distinct from coconut. The
Attalea butyracea is especially important to our discussion because it has relatively
large fruits (4.58.5 cm long by 34.5 cm wide) that look somewhat like very small
dried coconuts, and the palm has a similar stature to that of coconut (Henderson et al.,
1995), permitting the supposition that some of the first reports may have confused
this Attalea with coconut. A. butyracea grows on the slopes above beaches and river
courses along the Pacific coast of Panama in many places today.
Large numbers of endocarp fragments of Attalea butyracea are reported at numer-
ous locations in the Gran Chocl and Gran Chiriqu regions of western Panama from
7000 BP to European conquest (Smith, 1980; Cooke and Ranere, 1992; Dickau,
2010). Phytoliths, which are microscopic silica bodies found within and between cells
in many plants, of palms are also abundant in the same region during the same period
(Dickau, 2010).
It is important to mention that most of this work was carried out in the central and
western parts of Panama. In terms of the Pacific coast, the Gran Chocl and Gran
Chiriqu regions of Panama extend from near Nata on the coast of Parita Bay west of
Panama City to near Burica on the coast of the Gulf of Chiriqui, at the frontier with
Costa Rica (Fig. 1, Panama insert). Specifically, Dickau (2010) mentions finds at La
Pitaya, a small coastal island about 50 km east of Point Burica, and several coastal
C.R. Clement et al.
sites about 50 km southwest of Nata (Vampiros, La Mula Sarigua, Monagrillo, Cerro

Juan Diaz), but she does not report coconut. In comparison, the Gran Darien region of
eastern Panama is much less well studied. Hence, none of the archaeological sites
reported are close to Chimn or La Candelria Bay.
In summary, no archaeological evidence exists for coconut in Panama before
European conquest, as observed by Morcote-Rios and Bernal (2001: 342), even
though the woody endocarp of the coconut is an appropriate material to be preserved
at archaeological sites in wet environments. However, the absence of evidence is not
necessarily proof of absence, as the archaeological record tends to be determined by
sampling intensity, artifact preservation, and luck. Several archaeological sites are
close to historically mentioned sites, but none are exactly at those historically
mentioned sites, so future work may reveal new information.
The Ethnobotanical Evidence
Although many large palms have been important in Native American subsistence in
Panama since well before European Conquest, there is no record of use of coconut by
native Panamanians at the time of conquest (Patio, 2002), although the native
Panamanians apparently were familiar with its propagation (see Time Line 1514
1515 above). Patio (2002: 253) writes: There is no doubt that the inhabitants lacked
the tradition about the use of this plant; Would one have to conclude that these
Nations met this palm at least four generations ago, but did not use it? Patio states
that there would have been no reason for coconut not to be culturally assimilated by
the native population quite rapidly, even if it were growing in relatively remote
locations. This is a remarkable anomaly given Native American uses of a wide
variety of palm species, including endosperm, and even the domestication of peach
palm (Bactris gasipaes Kunth; Clement, 1995). This dramatic lack of evidence is a
strong reason to doubt the accuracy of the first Spanish reports, or at least to assume
coconuts could not have arrived in Pacific Panama more than a few years before
European Conquest.
The Linguistic Evidence
Paleobiolinguistics uses the comparative method of historical linguistics to recon-

struct the biodiversity known to human groups of the unrecorded past (Brown, 2006a,
b, 2010). By comparing words for biological species in languages of the same
language family, paleobiolinguistics facilitates reconstruction of terms for plants
and animals in the vocabularies of their ancestral or proto-language. Reconstructed
words for species are indicative of their substantial significance to speakers of proto-
languages. The approach, then, has the potential to contribute to the discussion of the
antiquity of coconut in Panama by comparing words for the species in genetically
related modern languages spoken there and in adjacent areas.
The modern languages of Panama include Spanish, Creole, and Amerindian
languages affiliated with two language families, Chibchan and Choco (Fig. 4). The
Chibchan languages of Panama include (from west to east) Teribe, Ngbare, Buglere,
Fig. 4 Map of the distribution of modern languages of Panama created with Ethnologue (Lewis, 2009)
and Kuna. Other Chibchan languages are spoken in Honduras, Nicaragua, Costa
Rica, and Colombia. The smaller Choco language family includes Ember and Woun
Weu, both having dialects spoken in Panama and Colombia.
A large portion of Panama is empty of contemporary Native American languages
(white areas of Fig. 4), notably the western Pacific coastal area and central parts of the
country. This does not mean that no indigenous languages were spoken there in the past.
Archaeological sites in parts of the area attest to Native American occupation, perhaps as
late as the 19th century (Locascio, 2010). However, very little, if any, evidence bears on
the language or languages of these archaeological populations. Kaufman (1994) pub-
lished a time-of-contact language-distribution map for the Caribbean region that in-
cludes southern Central America. On that map much of the contemporary empty regions
of Panama (Fig. 4) is identified as being filled by speakers of Mobe, Doraske, and
Bokota (in the west) and Kuna (in the east), all Chibchan languages.
For this study, we undertook a paleobiolinguistic investigation of domesticated and
useful plants, including coconut, in both Proto-Chibchan and Proto-Choco. Plants
that reconstruct for Proto-Chibchan, spoken at the latest around 4400 BP (Holman
et al., 2011), include cacao (Theobroma cacao), cotton (Gossypium hirsutum), hog
plum (Spondias spp.), maize (Zea mays), manioc (Manihot esculenta), sweet potato
(Ipomoea batatas), and tobacco (Nicotiana tabacum) (cf. Constenla Umaa, 1981).
Those reconstructed for Proto-Choco (c. 2258 BP) include Banisteriopsis caapi,
bottle gourd (Lagenaria siceraria), guava (Psidium guajava), maize, manioc, and
sweet potato. No words for coconut reconstruct for these prehistoric proto-languages,
suggesting that the species was not known to their speakers.
Comparative evidence is such that a word for coconut is not reconstructable for
any chronological stages of Chibchan and Choco language families, including those
closest to the time of the European conquest. For example, a term for coconut cannot
C.R. Clement et al.
be retrieved for Proto-Ember, a daughter language of Proto-Choco spoken at the

latest around 875 BP. This is due to the fact that all contemporary offspring languages
of Proto-Chibchan and Proto-Choco for which lexical sources are available fail to
show native terms for coconut that are cognate. In fact, all of the terms for the referent
in available lexicons are of non-native origin. All languages except one or possibly
two have borrowed words for the plant and its fruit from a European language, either
Spanish (coco) or English (coconut).
Twelve terms for coconut, presented in original orthography, were extracted from
lexical sources available to us for Chibchan and Choco languages (Table 1). All but two
terms in the 12 languages are unambiguous loans from European languages. The two
exceptions are Boruca siahu and Kuna goba, both of which are so phonologically
dissimilar to one another as to exclude the possibility of cognation and, thus, the
possibility that a word for coconut with such reflexes pertained to Proto-Chibchan. In
fact, the phonology of the Kuna word is such that its status as a loan based on Spanish
coco cannot be ruled out as a strong possibility. Of the remaining 10 terms, nine are
loans based on coco, and one (in Rama) is a loan from English (coconut).
Words for coconut in the 12 languages robustly suggest that the plant was
introduced by Europeans into southern Central America. Speakers of native lan-
guages of Latin America typically have named newly encountered items, including
plants, animals, and artifacts, by borrowing words for these things from languages of
the people that introduced them, in this case from Spanish and Portuguese (Brown,
1994, 1999). On the other hand, only rarely have native terms for indigenous things
been replaced by Spanish or Portuguese loanwords (Brown, 1999: 92104).
However, occasionally Latin America Indians have coined words for introduced
items by using the lexical resources of their native languages rather than by borrowing
a term from a European language. This practice has been very common in languages
spoken by Amerindians influenced by English, French, and Russian intruders, but
comparatively rare among native languages of Latin America (Brown, 1994, 1999).
For example, in the Bachajn dialect of Tzeltal, a Mayan language of southern Mexico,
Table 1 Terms for coconut in Chibchan and Choco languages
Coconut Term Language Family Location
siahu Boruca Chibchan Costa Rica

koko Bribri Costa Rica
k ko Chimila Colombia
koko Dorasque (extinct) Panama
coco Guatuso Costa Rica
goba Kuna Panama
kko Ngbare Panama
kukunp Rama Nicaragua
kokoha Paya (Pech) Honduras
kko Northern Ember Choco Panama
koko Epena Panama
kk Woun Weu Panama
the introduced sheep is tumin ix, literally cotton deer, a usage almost certainly
motivated by the resemblance of the European sheep to the native deer, the most salient
mammalian herbivore known to Tzeltal speakers (Witkowski & Brown, 1983).
The only word for coconut of the 12 languages that clearly is not a European loan
is Boruca siahu. Plausibly, this term originally denoted a native palm similar to the
introduced coconut. Evidence for this comes from Cabcar, which, like Boruca, is a
Chibchan language of Costa Rica. Cabcar contains a word similar to the Boruca
term, i.e., ser, designating Acrocomia aculeata, a palm whose fruit is fed to cattle
and occasionally consumed by humans in the Cabcar region. The phonological
similarity of Boruca and Cabcar words may be due to cognation or, if not, to
borrowing. In either case, this suggests the original referent of the Boruca term was
A. aculeata, a word that later became referentially extended to the introduced
coconut. No term for coconut is listed in Margerys (1989) exceptionally thorough
dictionary of Cabcar, and no term for A. aculeata is found in Quesada Pacheco and
Rojas Chavess (1999) comprehensive dictionary of Boruca.
In conclusion, a pre-Columbian presence of coconut in Panama and surrounding areas
is not attested by paleobiolinguistic evidence. Indeed, this evidence strongly suggests that
the modern occurrence of the plant in the region is accountable to European introduction
in historical times.
We have also undertaken a preliminary paleobiolinguistic survey of many language
families of Latin America for evidence of pre-Columbian coconut, mainly focusing on the
reconstructibility of terms for the plant in proto-languages. With only one possible
exception, no such terms are apparent. The possible exception is Proto-Chinantecan
(c. 1935 BP), whose contemporary offspring languages are spoken in northern Oaxaca
state, Mexico. For this ancestral language, Rensch (1989:78) reconstructs *h:H (H=high
tone), assigning to it the gloss coconut. However, a survey of reflexes of this hypothetical
word in various Chinantecan languages shows that some of these denote Acrocomia
aculeata rather than, or in addition to, coconut. This suggests that, like the Boruca term
discussed above, the Proto-Chinantecan word designated A. aculeata, and that its reflexes
were referentially extended to the introduced coconut. If so, paleobiolinguistics evidence
assembled to date fails to provide support for the prehistoric occurrence of coconut for any
region of Latin America. This finding concurs with the work of Merrill (1937), who
contrasted abundant linguistic evidence for a long human association with coconut in
Southeast Asia and Oceania with the lack of anything similar for the Americas.
A Preliminary Summary of the Evidence
The genetic evidence concerns modern coconuts, so says nothing about the historical
presence of coconut in Panama. The historical record had previously been accepted
with caution (Patio, 1963, 2002; Zizumbo-Villarreal & Quero 1998), but the clear
absence of archaeological, ethnobotanical or linguistic evidence suggests that the
caution was warranted. This lack of interacting evidence is an unexpected result when
applying de Candolles and Vavilovs methodology, and suggests either that coconut
arrived immediately before European Conquest, rather than 2,250 years BP as
suggested by Baudouin and Lebrun (2009), or it arrived after European Conquest.
Either way, both the genetic evidence and historical records need further study.
C.R. Clement et al.
Genetic Sampling and Expanded Analysis
In any study of the genetic relationships among populations, such as coconuts in

Panama, Mexico and the Philippines, the sample used will affect results. Hence,
we examine the samples used in the genetic studies, especially those from Mexico
and Panama. The coconut data set was developed within the framework of the
Generation Challenge Program (GCP) of the Consultative Group on International
Agricultural Research by Luc Baudouin and Patricia Lebrun, of CIRAD, the
French Agricultural Research Center for International Development, Montpellier,
France. This data set was not designed primarily to identify the origin of the
Pacific coast Panama Tall, although Baudouin and Lebrun (2009) used it this way
with interesting results. It is worth mentioning, however, that sampling in Oceania,
including the Philippines, does not yet represent the variability that is present in
the region either (Fig. 1, contrast between shaded areas and the Pacific Ocean).
Even the Philippines, where coconut is a major crop today, is not well represented
(Table 2).
Table 2 Coconut varieties included in the Generation Challenge Program data set that were analyzed with
microsatellite markers for the Baudouin and Lebrun (2009) and Gunn et al. (2011) studies. Variety names
follow GCP/CIRAD nomenclature
Variety n Comments
The Philippines 46
Ballesteros Tall Tarraq 7
Baybay Tall 8
Macapuno Tall 5
Pandan Tall 6
San Ramon Tall 6 Fruit similar to Panama Tall
Tagnanan Tall 14
Mexico 43
Pacific Tall Colima 14 11 plants possibly introduced from Rennell Island (Solomon Islands,
Melanesia) in 1569fruit similar to Rennell Tall; 3 plants with
fruit similar to Philippine varieties
Pacific Tall Guerrero 11 8 plants with fruit similar to San Ramon; 3 plants with fruit similar
to Baybay Tall
Pacific Tall Michoacn 14 Unnamed Philippine variety introduced in the 1930s
Pacific Tall Nuxco 4 Possibly introduced from the Philippines into Acapulco shortly after
1572
Panama 105
Panama Tall 44 Some introgression with Indo-Atlantic; includes 5 from
Oxtapacab, Yucatan, Mexico
Panama Tall Agua Dulce 13 West of Nata, inland, taken to Jamaica, then Ivory Coastconsiderable
introgression with Indo-Altantic
Panama Tall Bowden 10 Taken to Jamaica
Panama Tall Costa Rica 19 Costa Ricaminor introgression with Indo-Atlantic
Panama Tall Monagre 19 West of Nata along coastminor introgression with Indo-Atlantic
The varieties in the GCP data set that interest us most are those from the
Philippines, Mexico and Panama (Table 2). Dwarf varieties from the Philippines
are not included in the table, as these are unlikely to have contributed to this Panama
Talls genetic composition, nor is the Mexican Atlantic Tall, as this variety originated
in the Indo-Atlantic group of varieties (Harries, 1977; Gunn et al., 2011).
The sample of Mexican Pacific Tall coconut varieties used by Baudouin and
Lebrun (2009) and Gunn et al. (2011) includes several with known origins and dates
of introduction, but some plants have been joined into state-level varieties without
due consideration of their morphology (Table 2). The Pacific Tall of Colima is a
mixture of 11 plants that have Rennell Tall fruit morphology and three that appear to
be from the Philippines. The 11 Rennell Tall-type plants are derived from a plantation
established in 18901900 at the margins of the Coauhuayana River in Tecoman,
Colima state. It is represented in the vast majority of plantations in Colima, the
western portion of Michoacn and eastern Jalisco states. The seed originated in the
environs of the city of Colima, from orchards established in the 18th century. They
may be progeny of the introduction made by Alvaro de Mendaa from the Solomon
Islands in 1569 (Zizumbo-Villarreal & Colunga-GarcaMarn, 2001). The Pacific Tall
of Guerrero includes eight coconuts that are morphologically similar to the San
Ramon variety and three that are similar to the Baybay Tall variety. No historical
record of these introductions exists, but they agree with the Philippine location of
probable early introductions (see below). The Pacific Tall of Michoacn was intro-
duced by President Lzaro Crdenas, apparently from the Philippines, and was
initially established in plantations in 19371938. This is the most representative
variety between Acapulco and Lzaro Crdenas Port, Guerrero state (Zizumbo-
Villarreal & Colunga-GarcaMarn, 2001). The Nuxco plantation was established in
the 1950s, possibly from progeny of the early introductions to Acapulco from
Philippines. All of these varieties have clearly different microsatellite profiles when
compared to the Pacific coast Panama Tall (Gunn et al., 2011: Table S1).
Numerous samples of Panama Tall from the Pacific coast were taken into the
Caribbean during the 20th century and were then distributed elsewhere. When the
Maypan hybrid (a cross between the Malayan Dwarf variety and the Panama Tall)
was produced in Jamaica (Harries & Romney, 1974) it was resistant to lethal yellowing,
the most important coconut disease in the Americas, and the Panama Tall became very
important in the Caribbean (Harries, 1995). However, not all coconut populations along
the Pacific coast of Panama were sampled. For example, the sample used by Baudouin
and Lebrun (2009) and Gunn et al. (2011) did not include plants from the Gran Darien
nor the Gran Choc (present day Pacific coast of Colombia), where historic sources
place the pre-conquest presence of coconut at Chimn and Cape Corrientes, respective-
ly. Thus, the sample is biased towards the middle-western portion, covering the Gran
Chocl and Gran Chiriqu regions. Note also that these middle-western regions are
precisely those where Native American populations have disappeared (Fig. 4), so one
might expect coconut introductions from other localities during the colonial and modern
periods. Unfortunately, we have not found a history of these introductions.
Additionally, the various samples of Panama Tall from the Pacific coast listed
above (Table 2) were not collected directly in Panama for the GCP study. Rather, they
had been collected years earlier and taken to various countries before having their
DNA extracted (Luc Baudouin, pers. com., 2011), which explains part of the Indo-
C.R. Clement et al.
Atlantic alleles found in some plants that is evidently due to introgression (see Gunn
et al., 2011: Table S1).
The two genetic studies were both well executed for the questions that they asked, but
neither took the opportunity to look more closely at the variability within the countries
involved. Part of this is due to the very small sample sizes (Table 2), especially in the
Philippines, but also Mexico and to some extent even Panama, because this will determine
the reliability of the relationships found. Nonetheless, a look at these relationships can
offer ideas for new studies. Hence, we extracted the microsatellite genotypes for the
Philippines, Mexico and Panama from the Global Challenge Program dataset, and used
Nei et al.s (1983) genetic distance, the Neighbor Joining algorithm, and 1,000 bootstrap
iterations to obtain a preliminary idea of relationships. Because sample sizes are small and
all bootstrap confidence levels are weak, we do not present the dendrogram. However,
there seems to be a relationship between the Philippine San Ramon variety and the
Mexican Pacific Tall of Colima, which is curious because this Mexican variety may have
originated in the Solomon Islands (Table 2). All the other Mexican varieties appear to be
derived from this relationship and are a sister group to the Panamanian varieties. The
relationship with all the other Mexican varieties may be due to the close relationship of
San Ramon with a small set of Philippine varieties (Baybay, Pandan, Tagnanan), one of
which (Baybay) is morphologically similar to the Pacific Tall of Guerrero (Table 2). The
modern Pacific coast Panama Tall is also morphologically similar to San Ramon (Vargas
& Blanco, 2000). Hence, this preliminary analysis suggests an introduction of coconut to
Mexico that then influenced other introductions and was also introduced to Panama, with
the very small founder event detected by Baudouin and Lebrun (2009).
To look even more closely at how the Panama Tall may be related to the Philippine
and Mexican varieties, we extracted the Structure assignments at K=5 from Gunn
et al.s (2011) Table S1 (Fig. 5). Four Philippine varieties (San Ramon, Baybay,
Fig. 5 Group assignment at K=5 for the Philippines, Mexican and Panamanian coconut varieties studied
by Gunn et al. (2011). The colors represent the five groups identified by Structure analysis; the length of
each color in each bar is the proportion of each group in that plant; the size of the slice is the proportion of
each group in that country. Redrawn from Gunn et al. (2011) Table S1
Pandan, Tagnanan) have numerous plants with considerable proportions of Panama

assignment, so even though San Ramon is identified as the most probable ancestor of
the Pacific coast Panama Tall, other varieties could have contributed and would not
be easily detected because of the extremely reduced size of the Panama Tall founder
event. From the Structure analysis for the Mexico Pacific Tall of Colima, it is possible
to hypothesize that the proportions of Panama and Papua New Guinea assignments
explain its similarity with San Ramon, even though the high proportion of South
Pacific confirms the origin of Colima in the Solomon Islands. The Mexican varieties
also have considerable proportions of Panama assignment, although less than the
Philippines overall. Nonetheless, numerous plants in this small sample have enough
Panama assignment to suggest that the Panama Tall is derived from the Philippines
via Mexico. We then took the GCP dataset and analyzed the Philippine, Mexican and
Panama varieties with Structure 2.3.3 (Pritchard et al., 2000; Hubisz et al., 2009),
following Gunn et al.s (2011) parameters. This generated three groups (data not
shown), with all Philippine and Mexican varieties except the Mexican Atlantic Tall in
one group, all Panama varieties in one group, and the Mexican Atlantic Tall in the
third group. The San Ramon variety had some Atlantic assignment, as is also visible
in Fig. 5, and there is some admixture of Atlantic with Panama. This new analysis
strongly supports the Philippine Mexico connection, but is less clear about showing
relationships with Panama, given the fact that only the Philippine varieties represent-
ed the whole Pacific Ocean.
The similarity of Pacific coast Panama Tall and named varieties in Southeast Asia
has long been noted (Harries, 1978; Zizumbo-Villarreal et al. 1998; Zizumbo-
Villarreal & Quero, 2005), in particular with the San Ramon type in the Philippines
(Vargas & Blanco, 2000). Considering the lack of archaeological, ethnobotanical and
linguistic evidence for coconut in Panama at conquest, it seems reasonable to ask how
the San Ramon type might have arrived in Panama. The Manila-Acapulco galleon
route (Fig. 1) that had been suggested by Safford (Small, 1929; Hill, 1929), although
post-conquest, is the obvious candidate. Although it had been accepted by Merrill
(1954) and others (Harries, 1971, 1978; Zizumbo-Villarreal et al. 2005), it has not
been closely examined until now (see also Harries, 2012).
In 1564 an expedition to establish a Spanish settlement set out from Puerto de la
Navidad (Barra de Navidad), Jalisco, Mexico to avoid sailing through Portuguese
waters, setting a course for Cebu in the Visayas, where Magellan had landed 41 years
previously. Manila subsequently became the premier city of the Philippines, because
it already had trade links with China and Japan and, as the northernmost harbor, it
became the departure point for the galleon route. However, on the first occasion in
1565, Alonso de Arellano and Andrs Urdaneta, returning independently, both sailed
from Cebu and not from Manila, which is significant because there were superior
coconuts in that region. An agricultural observer in the seventeenth century, Father
Francisco Ignacio Alzina, who resided in the Visayas from 1634 to 1667, wrote:
There are very big ones [coconuts] which would measure more than one azumbre
(~ 2 l) (Alzina 1668). At the head of the Sulu Sea, Cebu in the Visayas was not far
from Mindanao and, when the production of copra became commercially important
in the Philippines at the beginning of the 20th century, the San Ramon coconuts in
Mindanao were highly regarded. This was because there are no records from any
other part of the world of plantation averages showing such size of nut as those of San
C.R. Clement et al.
Ramon (Copeland, 1914). However, similar coconuts were reported from Colom-
bia at the same time: Gorgona Island between 3rd and 5th parallel N of Equator 24
miles off Colombia . . . is famous for producing coconuts of immense size and are of
great use to planters as seed nuts (Bardy, 1914).
The possibility that coconuts were carried to Mexico in 1565 has previously been
discounted (Bruman, 1945), because coconuts were not recorded on the list of
provisions. However, Arellanos account of cooking oil solidifying (literally freez-
ing) is strong circumstantial evidence of coconuts as deck cargo for the crew to drink
or use when preparing food (Harries, 2012). The early germination of this
typemore than 75 % in 105 days (Harries, 1981)would have meant that, in
August 1565, at the end of a four-month voyage to Puerto de la Navidad, there would
be seedlings ready for planting. The Barra de Navidad lagoon borders Colima
province, which became, and is still today, the center for coconuts in western Mexico.
It also seems reasonable to suppose that the subsequent Manila-Acapulco galleons
would carry the same sort of coconuts. This could be done, either by arranging for
them to be collected in Mindanao or the Visayas for transshipment or, more easily, by
plantings near to Manila. This would explain Copeland's note that San Ramon
coconuts were in general cultivation in the coastal district of Pangasinan province,
Luzon and a report that the largest nuts in the world are produced around Lingayen
Gulf, Luzon Island (O.W. Barrett cited by Smith & Pape, 1914: 537). These
locations, to the north of Manila, would have been the most convenient for taking
deck cargo on board. Confirmation that seedlings were planted in Mexico comes from
the activity of skilled Filipino toddy tappers in 1580, who tapped tuba for
fermenting to coconut spirits for consumption in Mexico (Zizumbo-Villarreal
1996; Zizumbo-Villarreal & Colunga-GarcaMarn, 2008). As first the tuba market
and in the 20th century the copra market expanded, later introductions of larger
numbers of seednuts intended only for planting would have been made.
Thus, the Panama Tall coconuts could have come from one or more locations, but
still represent the San Ramon and closely related varieties. It is even likely that the
seedlings from the post-1565 introductions into Mexico were fully grown and in
bearing within five to seven years and the year-round production of seednuts, rather
than the once yearly supply from Manila, would be disseminated from Colima
southwards as far as Peru, including Gorgona Island, Colombia (Fig. 1), for example.
A single, small sample from this source might explain the extremely narrow genetic
base of the Panama Tall reported by Baudouin and Lebrun (2009). Zizumbo-
Villarreal and colleagues in Mexico are collecting and analyzing the Pacific coast
Tall varieties there, as well as in other locations along the Pacific coast southwards.
Further DNA analysis of the San Ramon and similar varieties in the Philippines and
their comparison with the Pacific coast Talls of Mexico and Panama will permit these
relationships to be refined with much better precision than is possible with the current
dataset.
Botanical and Historical Questions
Given the doubts about the historical record, we revisit Oviedos account because,
although it was previously analyzed by Patio (1963, 2002), Allen (1965) and
Zizumbo-Villarreal & Quero, (1998), the DNA data match between the Panama and the
Philippines coconuts raises fresh concerns. Oviedos account of coconut has been
questioned on a number of occasions because the illustration does not match the
description, parts of the description can match other palm genera and parts of the
information may have come from Asia rather than the Americas, as indicated above.
Patio (1963, 2002) discards the first problem by pointing out that Oviedo finished
describing the cane palm, Bactris major, on the page where he started describing
coconut. While Patio and Zizumbo-Villarreal & Quero looked at the similarities and
cautiously accepted that Oviedo had seen coconuts, Allen saw differences and suggested
that Oviedo may have regarded them as aberrant and inferior sorts. We consider their
opinions and identify two properties, found only in genuine coconuts, which Oviedo did
not mention and apparently did not know about. The following sections of Oviedos
account (as translated for use by Allen, 1965) are worth transcribing and commenting on.
There are other palm trees whose fruit are called cocos, this being a genus of large
palm trees, and whose leaf is of the same kind as that of the date palms, This is
the first sentence of the account and suggests that numerous Neotropical palm fruits
were called cocos, which is at odds with our linguistic analysis. Hence, what we may
be seeing in this sentence is the Spanish naming of native American palm species.
Over the next centuries, 131 palm species or sub-species were attributed to the genus
Cocos. These other palms are now assigned to different genera (Beccari, 1917;
Henderson et al., 1995) and Cocos nucifera is monotypic. Some of them have statures
similar to coconut, with pinnate leaves, although many have spines that are hard to
miss (Acrocomia, Astrocaryum, Bactris). The spineless ones are now in Attalea,
which has the appropriate stature, fruit that look somewhat like small coconuts (see
archaeological evidence above), and today grow on the hills above the beaches of
Panama.
These trees or palm trees put forth a fruit which is called coco, Altogether, such
as it is on the tree, it has a much greater bulk than a mans head; A century before
Columbus crossed the Atlantic, the Arab Ibn Battuta (1929) visited East Africa and
India, and reported that coconut is the size of a mans head, so this kind of
information was available to historians in Iberia at the time that Oviedo wrote, as
was Varthemas (1510) account.
But there in these our Indies the Indians do not trouble to cure these cords and cloths
which can be made from the wool or burlap of these cocos, such as in the Levant, for
here there is much cotton and henequen and cabuya to supply such necessity for cords.
These comments by Oviedo are with respect to the fibrous mesocarp of coconuts and a
clear indication of the lack of use by Native Americans, as noted above with respect to
the lack of ethnobotanical evidence. The description appears to be a justification for lack
of use, since the Native Americans had numerous other good fibers.
This fruit which is within that burlap, is the coco, as big as the fist of a closed
hand, and some as big as two fists, and more or less, it is a sort of round nut, and some
are elongated. The crust is hard, and as thick as the width of the inscription [title] on a
Castillian silver real [coin]. Inside, attached to the crust of that nut or coco, is a meaty
part in width like half the thickness of the small finger of the hand, or as thick as a
writing quill of the kind common to geese. This part of the description is somewhat
at odds with the original description of the size of the fruit, so if these were coconuts
they had small nuts, quite unlike the Panama Pacific Tall or San Ramon varieties.
C.R. Clement et al.
This is the fruit proper of the coco and what is edible, and it is as white as a
cleaned almond and better tasting than almonds, and of smooth taste to the palate. It is
eaten the same way peeled almonds might be eaten, This is a good description of
the endosperm of a coconut, but will be contrasted below with another description.
By way of pith or marrow of this fruit which is in its middle, , is a place taking
up the remaining part or entire quantity of the coco, full of a most clear and excellent
water, and as much as would fill the shell of the egg of a hen, and more or less, in
proportion to the bigness or size of the coco: Again, this is a good description of
coconut water in an immature coconut. However, Oviedo does not mention the sound
of the water splashing in the cavity of a mature coconut when shaken, as the water is
naturally absorbed when the nuts mature.
After I wrote the report I have mentioned, I was in the province and headland of
Borica, and I ate some of these cocos and carried many with me to Nicaragua, and
came to loathe them, and others did as I did and said the same thing as well. This
statement was not cited in Patio (1963, 2002) nor in Zizumbo-Villarreal & Quero,
(1998), although both read it since they mention Oviedo going from Burica to
Nicaragua. Some individuals do find the kernel indigestible and may become tired
of it, as Allen pointed out, but that is not usually a group phenomenon. Note also
Oviedos choice of ate rather than drank, as it suggests that they were fruit with
little water. Oviedos use of the word aborrec (loathe) is a surprising reactionmore
people would agree with Charles Darwin (1860: 407): After walking under a
burning sun, I do not know anything more delicious than the milk of a young
cocoa-nut. Perhaps the water had been absorbed by early germination, typical of
Panama Tall (Harries, 1981), but Oviedo does not mention the soft, sweet and very
edible haustorium (or apple) that would immediately identify a real coconut. Nor
does he say if they were being taken to Nicaragua for planting. So, the question is:
What palm was this? The answer to this question has implications for the entire time
line presented above and for the initial Spanish contacts with coconut in the
Americas. It is also possible that this is a mixture of information from Asia (size,
endosperm, water, flavor, etc.) and the Americas.
The list of palms near the Pacific coast of Panama with large enough stature to be
confused with coconut by non-specialists was presented in the archaeological evi-
dence (above). Even non-specialists would note and comment on spines on the trunk
and leaf petioles, as pointed out by Allen (1965), so the spiny species can be
discarded. That leaves Attalea butyracea, whose fruits can superficially be confused
with very small coconuts. The seeds of A. butyracea do not have liquid endosperm
when ripe, but they are perfectly edible. Like coconut, some people may not like the
flavor of the seed, and any palm seed, including coconut, can go rancid if stored in
conditions that do not allow germination but do allow respiration, but Oviedos
account does not suggest this. Coconuts on the open deck of a boat would germinate
rather than rot and, if they had been present in Panama at this time, they would be
regularly carried for refreshment on any coastal craft.
Moving on from Oviedo, there is a logical question related to the 1539 letter to
Hernn Corts (see Timeline above): If coconuts were interesting enough to send
from Panama to Mexico, why werent they also sent to Madridat least to show at
court? Given the lack of this kind of display, it seems likely that 20 years after the first
report there still were no real coconuts on the Pacific coast of Panama.
A Second Summary of the Evidence
The genetic and morphological evidence clearly shows that the Panama Tall is closely
related to varieties from the Philippines and the San Ramon variety is a likely
candidate, but the microsatellite information suggests that the original sample was
very small and did not capture the full San Ramon genetic profile. The San Ramon
variety was probably introduced into Mexico early via the Manila-Acapulco route.
The historical evidence is not as clear as might be hoped for and appears to mix
information from Asia about real coconuts with information from the Americas about
other palms. With this summary in mind, we look at the hypotheses about how and
when coconut arrived along the Pacific coast of Panama.
Hypotheses about Coconuts in Pre-conquest Panama
Five hypotheses have been presented to account for the historical observations, three
of which (24) are not mutually exclusive. These are:
Hypothesis 1 Coconuts originated in the Americas. This was held by de Candolle at
first (1855), but discarded later (1883). The hypothesis of American
origin was reinstated by Cook (1910), strongly criticized by Beccari
(1917) and Merrill (1937), and thereby decisively debunked. Gunn
(2004) placed a final nail in its coffin with a molecular genetic
phylogeny of the Cocoseae.
Hypothesis 2 Coconuts floated from mid-Pacific islands on one or many occasions
from ancient times to the present day. This hypothesis is based on the
fact that coconuts can disperse by floating over some distance, wash-
ing onto a suitable shore, striking root and growing. But, given the
genetic relationship between the Pacific coast Panama Talls and
coconuts in the Philippines, two questions arise: Could they have
floated across the Pacific Ocean (perhaps via intermediate islands), or
must they have been carried? Both cases have been argued by many
scholars. For example, Bruman (1944) and Purseglove (1972) argue
for natural dispersal, while Dennis and Gunn (1971) argue for car-
riage by man. A crucial question is the period for which coconuts will
remain viable when floating in the sea. In two experiments to test the
viability of nuts floating in the sea, the longest period any nut floated
and remained viable was 110 days (Edmondson, 1941; Ward & Allen,
1980). This suggests that the maximum flotation period may be on the
order of four months. In an experiment at the Coconut Experimental
Station in Sulawesi in 1931 (Reyne, 1948), nuts were floated in
barrels of sea water, but the water temperature would have been
significantly higher than actual sea temperatures. The lower temper-
atures of floating coconuts delay germination compared with nuts at
average ground level temperatures (Ward & Allen, 1980).
Computer simulations of Pacific wind and surface current direc-
tions and speeds, islands and coasts allowed tests of the possibility of
C.R. Clement et al.
coconuts drifting across the Pacific from several possible starting

places (Levison et al., 1973; Ward & Brookfield, 1992). It is impor-
tant to note that, as a coconut floats, the husk absorbs water, the nuts
weight increases, it floats deeper in the water and the effect of wind
relative to current decreases. In the simulation experiments, variations
were used to cover current only, wind only, and combined wind and
current forces. Experiments were conducted for drift periods of 4, 6
and 8 months. The last two periods far exceed any known example of
the time a coconut might retain viability when floating.
Initial simulation experiments from Ducie and Reao, in the ex-
treme east of French Polynesia (Fig. 1), showed that nuts floating
from Polynesia had no chance of drifting to the Americas from that
part of Polynesia, even within 8 months (Ward & Brookfield, 1992:
4734). The Equatorial Counter Current offers the most likely possi-
bility for coconuts drifting from west to east and therefore starting
points used for the remaining simulated eastward drifts from Micro-
nesia were Christmas Island (in eastern Kiribati) and Palmyra in the
Line Islands, and Motuiti in the northern Marquesas, Polynesia
(Fig. 1). No coconuts drifted to the American coast in any of the
6,588 simulated drifts from these islands, even in those of eight
months duration. In the current only experiment, 37 % of drifting
nuts did reach the Galapagos group, the shortest crossing being in
178 days from Christmas Island, with a mean crossing time of
207 days. Both lengths of time far exceed the known viability period
for floating coconuts and, in any case, coconuts did not grow in the
Galapagos at the time of early European contacts. In wind only
experiments, no nuts made the crossing, and in wind and current
experiments, of the 732 nuts started from Christmas Island, only 1 %
reached the Galapagos, in a mean time of 225 days. The conclusion
must be that coconuts cannot drift to the Americas within any rea-
sonable period of viability.
Hypothesis 3 Coconuts were carried in canoes from mid-Pacific islands by Poly-
nesians on one or many occasions before discovery by Europeans.
This hypothesis was ruled out by Baudouin and Lebrun (2009), based
on the lack of similarity between the microsatellite profiles of differ-
ent Polynesian coconut varieties and the Panama Tall variety. How-
ever, voyagers throughout the Pacific Islands regularly carried green
and mature coconuts on their journeys for drink and food, and would
be expected to plant some coconuts in situations where they were not
already growing, or plentiful. Computer simulations of eastbound
voyages from Polynesian islands towards America show that such
voyages are possible from such starting points as Samoa, Tonga,
eastern French Polynesia and Rapa Nui (Easter Island) (Fig. 1)
(Levison et al., 1973; Irwin, 1992: 1634; Fitzpatrick and Callaghan,
2009). The voyage from Rapa Nui to the American coast would take
about one month, and from the Marquesas two months (Irwin, 1992:
214; Fitzpatrick & Callaghan, 2009: 218). From Tonga or Samoa the
journey might take between 66 and 128 days (Fitzpatrick and Calla-
ghan, 2009: 218). Simulated voyages from eastern Polynesia ap-
proaching the American coast tend to be carried northwards by
winds and the Peru Current in the latter part of their journey, and
some make landfall as far north as the northern Ecuador coast. Those
from Hawaii reach the coast of Nicaragua and Costa Rica
(Fitzpatrick & Callaghan, 2009). The distance and likely duration of
voyages from eastern Micronesia (Kiribati) are similar to those from
Samoa or Tonga, and such voyages would be more likely to reach the
Panama coast.
Hypothesis 4 Coconuts were carried in canoes directly from the Philippines,
bypassing any island where Polynesian coconuts grew at that time.
This hypothesis is based on Baudouin and Lebruns (2009) analysis
that showed the close genetic relationship between coconuts in the
Philippines and Panama, and has never previously been analyzed. As
coconuts cannot have drifted to the Americas within any reasonable
period of viability, one must conclude that they were carried there by
voyagers from the western Pacific Islands. If coconuts from the
Philippines reached Panama before the 17th century and did not come
from Polynesia, we need to consider whether voyagers from Microne-
sia, or elsewhere in the western Pacific, may have had the capacity to
carry coconuts to Panama. There is no record to suggest that Philippine
mariners had the necessary technologies and knowledge.
Micronesian communities, however, have striking traditions of long-
distance voyaging in the western Pacific Islands. Long-distance two-
way voyaging by Micronesian people and their sophisticated methods
of navigation are well documented (Lewis, 1972; Gladwin, 1970;
Finney, 1979; Hezel, 1983; Thomas, 1987; Rainbird, 2004). The sea
lanes between islands extending over more than 1,600 km were named
(DArcy, 2006:1545) and maps constructed of sticks and shells were
used. Sophisticated concepts of estimating distances sailed and direc-
tions followed were taught. Two-way voyages in large outrigger canoes
were conducted annually for economic and socio-cultural reasons over
the 2,400 km length of the Caroline archipelago (Fig. 1). Anson reported
in the 1740s that Marianas canoes (with their asymmetrical hulls) were
designed to sail as close as possible to the wind and that they could
reach a speed of 20 knots (quoted by Horridge, 1995: 148).
It is clear that Micronesians had the seamanship, navigational skills
and canoes to make long voyages of exploration. They did so by settling
groups of islands extending over 5,600 km from west to east and over
1,600 km northsouth, and maintained regular links between these
islands for many centuries. Irwin has pointed out that the general
trajectory of Pacific colonization was first upwind and that this im-
plies pragmatic strategies of exploration as it is safest to search in the
direction from which one can most easily return in the event of not
finding new land (1992: 81). Micronesians following such strategies
could have made long easterly voyages, for example from Kiribati or
C.R. Clement et al.
Christmas Island (Fig. 1). Finney (1985) also points out that eastward
journeys would be easier in El Nio years. Once east of about 125 West
longitude, they could readily take advantage of south-westerly winds to
reach Panama (see Irwin, 1992: 916). Using such equatorial routes
they would not encounter any Polynesian islands en route so that any
coconuts they were carrying would have come from stock in their home
islands. Although the Micronesian sample used by Baudouin and Le-
brun (2009) and Gunn et al. (2011) is only distantly related to the Pacific
coast Panama Tall, Micronesia borders on Melanesia, which has more
closely related coconuts in New Britain and northern Papua New
Guinea (Fig. 1), as well as being closer to the Philippines.
However, until more intensive genetic sampling of coconuts is done
in Micronesia, and clear archaeological evidence is found of Microne-
sian contacts with America, we cannot claim that Micronesians were the
trans-Pacific carriers. As Storey et al. (2011b) affirm, there is currently
no such evidence. However, if Polynesia is ruled out as a source by the
wide genetic gap between the sampled Polynesian and Panamanian
coconuts, the history and capacities for navigation of Micronesians
suggest they may be candidates for any pre-Spanish carriage.
Hypothesis 5 Coconuts were not present until carried by Spanish galleons returning
from the Philippines. This hypothesis is supported by the lack of
archaeological, ethnobotanical and linguistic evidence for the viabil-
ity of the other four hypotheses, and by the doubts raised about the
historical record. Baudouin and Lebrun (2009) appear to have as-
sumed that the great number of coconuts carried between the Philip-
pines and Mexico over a 250 year period was inconsistent with the
limited similarity between the Philippines and the Panama Tall, which
they regarded as typical of an extremely small founder population,
while different selection pressures accounted for observed differ-
ences. However, regardless of the number of coconuts carried for
consumption from Cebu to Navidad and then from Navidad to Pan-
ama, the reduction in population size each time was due to most of the
nuts being consumed before arrival and the selection pressure every
time was for the early germinators that survived to be planted. The
genetic evidence suggests a relationship that is amenable to future study.
In order to determine if coconut was on the Pacific coast of Panama at
the time of European conquest, it is essential to collect a truly represen-
tative sample of the coconuts along that coast, considering that the
historical record suggests that not all the coconuts reported may have
been similar to the San Ramon variety, even though the modern Pacific
coast Panama Tall is indeed so. This type of collection has already started,
led by Daniel Zizumbo-Villarreal, and will be analyzed with the same
microsatellite markers used by Baudouin and Lebrun (2009) and Gunn
et al. (2011). A larger sample of the relevant Philippine, Melanesian
and Micronesian varieties is also needed to allow a more precise
genetic analysis with the Pacific coast Panama Tall and Mexican
Tall varieties.
Conclusions
The new genetic evidence is quite clear that modern coconut varieties from the
Pacific coast of Panama are closely related to known modern Philippine varieties,
as previously shown by morphometric analysis. There is, however, no archaeological,
ethnobotanical or linguistic evidence that supports a pre-Columbian origin of these
Pacific coast Panama Talls. A reanalysis of the historical record strongly suggests that
early explorers made honest mistakes in identification. Hence, the most parsimonious
explanation is that the Panama coconuts were introduced after Spanish conquest. The
Manila-Acapulco galleon trade route that was active between 1565 and 1815 is very
probably the means by which the Spanish introduced Philippine varieties of coconut
to the Pacific coasts of the Americas. This is supported by the DNA analysis, and
history records Spanish voyages with coconuts. The very small founder event that
gave rise to the Pacific coast Panama Tall variety probably came from Mexico soon
after the first Mexican plantations were established. New collections along the
Mexican to Colombian Pacific coasts are improving the sampling for genetic analy-
sis, and new work in the Philippines is suggested to confirm precise origins. Unless
new archaeological remains are found to prove otherwise, this hypothesis can direct
new research on the origins of American Pacific coast coconuts.
Acknowledgements Our special thanks to Luc Baudouin, CIRAD, Montpellier, France, for information
about the samples chosen for coconut genetic analysis and for stimulating our reconsideration of pre-
Columbian coconuts; to Madhavan Nayar, formerly Director, Central Plantation Crops Research Institute,
Kasaragod, India, for providing bibliographic material; to Michael G. Price, Michigan Center, Michigan,
Kenneth M. Olsen, Washington University, St. Louis, Missouri, Lalith Perera, Coconut Research Institute,
Sri Lanka, for reading the manuscript in draft and making numerous useful suggestions, about the
Philippines, the genetic analysis and the presentation; and to Pamela Brown, Pensacola, Florida, for careful
review of the language.
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Coconuts in The Americas

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Bot. Rev.

Coconuts in the Americas

Charles R. Clement1,8 & Daniel Zizumbo-Villarreal2 &

# The New York Botanical Garden 2013

Keywords Cocos nucifera . Molecular genetics . History . Archaeology . Linguistics .

Palabras claves Cocos nucifera . Gentica molecular . Historia . Arqueologa .

The evidence for sweet potatoes includes archaeology, ethnobotany, linguistics

The Genetic Evidence

A Time Line of Spanish Contact with Coconut

1492Discovery of New World

1565Philippines to New Spain (Mexico) return trip was accomplished inde-

The Archaeological Evidence

sites about 50 km southwest of Nata (Vampiros, La Mula Sarigua, Monagrillo, Cerro

The Ethnobotanical Evidence

The Linguistic Evidence

Paleobiolinguistics uses the comparative method of historical linguistics to recon-

be retrieved for Proto-Ember, a daughter language of Proto-Choco spoken at the

Table 1 Terms for coconut in Chibchan and Choco languages

Coconut Term Language Family Location

siahu Boruca Chibchan Costa Rica

A Preliminary Summary of the Evidence

Genetic Sampling and Expanded Analysis

In any study of the genetic relationships among populations, such as coconuts in

Pandan, Tagnanan) have numerous plants with considerable proportions of Panama

Botanical and Historical Questions

A Second Summary of the Evidence

Hypotheses about Coconuts in Pre-conquest Panama

coconuts drifting across the Pacific from several possible starting

Harries, H. C. 1971. Coconut varieties in America. Oleagineux 26: 235242.

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