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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 Amazônia, Av. André Araújo, 2936, 69067-375 Manaus, AM, Brazil
2
Unidad de Recursos Naturales, Centro de Investigación Científica de Yucatán A.C., Mérida, Yucatán, 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 São 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 distribuyó a las costas del Atlántico y el Caribe. Sin embargo los registros del
cocotero en siglo XVI en la costa del Pacífico de Panamá son polémicos. Los análisis
recientes de ADN de poblaciones modernas de coco muestran que son similares a las
variedades Filipinas, lo que está de acuerdo con los análisis morfo-métricos previos. Por
lo tanto, el cocotero debe haber sido llevado en barco desde el Pacífico Occidental, pero
no hay evidencias arqueológicas, etnobotánicas o lingüísticas precolombinas. La
explicación más parsimoniosa es que fue introducido después de la conquista española.

C.R. Clement et al.

Esto es apoyado por los análisis de ADN y los registros históricos de los viajes españoles
con cocos. Nuevas colectas a lo largo de las costas del Pacífico desde México hasta
Colombia mejorarán el muestreo para el análisis genético, y se sugieren nuevos trabajos
en Filipinas para confirmar los orígenes precisos. A menos de que nuevos restos
arqueológicos se encuentren que demuestren lo contrario, esta hipótesis puede orientar
nuevas investigaciones sobre los orígenes del cocotero en la Costa del Pacífico
americano.

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

Palabras claves Cocos nucifera . Genética molecular . Historia . Arqueología .
Lingüística . Etnobotánica

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).

making this the first plant evidence for a Neotropical to Oceania dispersal. found in small areas in southern California and southern Chile. Sewn plank canoe-building technology.. but Patiño’s (1970) documental analysis does not agree. characteristic of Polynesia and Mi- cronesia. ethnobotany. 2011a) to propose that the chicken had been introduced into El Arenal. 1992). which thus explains Acosta’s record. genetic and some linguistic evidence was re- cently used by Storey et al. We thus seem to have a reasonable answer to the “where it came from” question. also suggests contact with the Pacific Islands (Jones et al. Although Acosta (1940) mentions them in the Caribbean and Mexico.. Previous work has already shown that this founder event could not be explained by coconuts drifting on ocean currents (Ward & Brookfield. but the “how it got there” remains an open question. Patiño (1970: 34. The information available for coconut is much less complete. from Polynesia a century before European conquest. No information on the archaeology and linguistics of coconut in Panama has yet been published. (2013) have now confirmed Yen’s (1974) hypothesis. 2009). with new genetic information and an historical record that has been questioned and defended by different authors. 1978).000 years before present (BP) (Yen. 2011). but is post-conquest in that region. 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. 2008a. while populations on the Pacific coast of Mexico contain consid- erable Philippine variation. 2009: 94) and thence to New Zealand and Hawaii. 2010.g. A combination of archaeological. 1974. The earliest was to Yen’s “ellipse” area of eastern Polynesia by 900– 1. which corresponds with the South American center of sweet potato domestication (Roullier et al. 2011). 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.. as the origin of these sweet potatoes. Only a fraction of the coconut genetic variation in the Philippines is present in Panama. A study by Zhang et al. linguistics and genetics. The sweet potatoes in Oceania appear to have originated from two separate introductions from America (Ballard et al. (2004) questioned whether the Ecuadorian origin was not representative of both origins and dispersals. Lebot. Ecuador.Coconuts in the Americas The evidence for sweet potatoes includes archaeology. 2011). Skeletal. or an introduction via Mexico after European conquest (Harries. artifact and non-material cultural evidence also suggests Polynesian contacts with the Chilean coast (Matisoo-Smith & Ramírez.. An associated question is “when. (2007. 2009). and concluded that Mesoamerican sweet potatoes dominate Oceania. 2005: 46–7. b). This lack of variation raises two possibilities: a direct introduc- tion from the Philippines to Panama (Baudouin & Lebrun.. but also only a fraction of the Panama genetic variation (Gunn et al.. Ramírez-Aliaga. Linguistic and other evidence is cited by Scaglion (2005: 35–41) for the Gulf of Guayaquil. Green. They counter earlier questioning of the validity of the archaeological results of the chicken remains in Chile (e. 2011. Storey et al. Ramírez-Aliaga. and seems reasonably complete. suggesting a single small founder event (Baudouin & Lebrun. 2011). Gongora et al. 2005). Roullier et al. Chile. although a trans-Pacific route is clear. (2007) affirm that Francisco Pizarro found chickens in Peru..” which raises contrasting possibilities: a very early direct introduction from the Philippines to .

Panama. 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. Such a reduction is the signature of a bottleneck. 1929.324 in this Panama Tall versus 0.250 BP. As our starting point. . 1954. 2. at a date immediately before European contact (suggested by the extremely limited endemic distribution reported at the time of European conquest). most probably in the Philippines (Baudouin & Lebrun. 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. which might help with the “how” question. C. Allele 222 is also the most frequent in Southeast Asia. Harries. Hill.e. By contrast. 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. 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. 2011). we re-examine the genetic and historical evidence. 2009). 1). e. 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. such as locus CnCir 2 (Fig. as well as with evidence from ethnobotany and history. as postulated by Baudouin and Lebrun (2009). linguistic and ethnobotanical evidence. 2011). 4) were carried by unknown mariners. This trade route did not necessarily involve a large number of introductions nor a large number of nuts at any one time.480 for all Tall coconuts. the Panama Tall on the Pacific coast is characterized by markedly reduced diversity (expected heterozygosity=0... 1971. Gunn et al. or over the Manila-Acapulco route less than 450 years ago (Small. they are readily distinguished using microsatellite markers (Baudouin & Lebrun. as the coconuts were carried for consumption by the crew and passengers during a three-month journey. and 5) were carried by Spanish galleons returning from the Philippines.R. 2) floated on ocean currents from Oceania. but once an initial planting came into bearing. i. 1978). and investigate the archaeological record.230 in this Panama Tall versus 0. a strong reduction of population size at some moment in the history of a population. 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. These hypotheses can be examined with linguistic and archaeological evi- dence.. Gunn et al. Merrill. 2009). we summarize in greater detail the new genetic information concerning its origin. as is typical of a small founder event. 2).579 for all Tall coconuts combined. 1929. This founder event is clearly seen by examining individual microsatellite loci.. 2009.g. when the Spanish crown inaugurated and managed the Manila-Acapulco trade route (Fig. observed heterozygosity=0. We apply de Candolle’s and Vavilov’s multidisciplinary methodology to the question of coconut on the Pacific coast of Panama at the time of European conquest. Although the coconuts of the Pacific coast of Mexico and of Panama came from the same general region. Clement et al. Finding that these latter lines of investigation are not fruitful. 3) were carried by Polynesians. Any early germinators not used could be planted.

Mic-Micronesia. see Baudouin & Lebrun. Mex-Mexico. GuN-North coast of New Guinea.1 0 0 Pan Mex Phi SEA Ind GuN Mar GuS NBr Van Mic Pol Fig. The Philippines variety is most similar to both this Panama Tall and to the Mexican Pacific coast 1 1 0. Solomon Islands. 2 Allele frequency of microsatellite locus CnCir2. Micronesia and Polynesia. Gunn et al..2 0.7 Allele frequency 0.1 0.3 0. Pol- Polynesia. SEA-Continental Southeast Asia. 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.4 0. Ind- Indonesia. NBr-New Britain. Map prepared by the Cartographic & GIS Services. Australia and its frequency is especially high in the Philippines (65 %). Its frequency decreases progressively from Southeast Asia to Melanesia.4 0.5 0.3 0. Redrawn from Baudouin and Lebrun (2009) . Australian National University.5 0. Canberra. Van-Vanuatu. allele 222. GuS-South coast of New Guinea. and similarity index across 30 microsatellites to compare allelic diversity of the Panama Tall with possible source regions in the Pacific Ocean basin.8 Similarity 0. Pan-Panama Tall.8 Similarity Index 0. 2011). Insets—locations on the Pacific coast of Panama where coconuts were reported shortly after European conquest. Phi-Philippines.9 CnCir2-222 0. 1 Map of the Pacific basin with features relevant to coconut in the Americas.2 0. 2).6 0. locations in the Philippines mentioned in the text. Mar-Markham Valley in New Guinea. New Caledonia. 2009. Such a tendency is observed at most of the 30 loci studied (data not shown.9 0.6 0.7 0.Coconuts in the Americas Fig.

Clement et al.R. Redrawn from Gunn et al. but he had misidentified the Royal palm (Roystonia spp). 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. since these would have been less preferred for consumption. who was appointed as one of the official historians of the Indies (Spanish American colonies) in 1523. A Time Line of Spanish Contact with Coconut The journal of Christopher Columbus (Cristobal Colombo) records that. Columbus thought he had reached Asia and found coconuts. Part of his “General and Natural History of the Indies” was published in 1535. Throughout he makes Fig. Thus. before returning to the Americas. after his return from Santo Domingo (today’s Dominican Repub- lic). However. (2011). while the Polynesian island sources are the least similar. 1893: 80). de los Rios y Serrano. 1996). only a small fraction of the allelic diversity found in the Philippines is found in this Panama Tall. The similarity of Mexican and Philippine varieties is probably due to several introductions after 1565. his “Natural History” is considered an important reference about the natural resources of the Americas. They found a very large nut of the kind belonging to India. Mexican and Panamanian varieties studied by Gunn et al. the initial selection pressures may have been uniform and closely related to trans-Pacific travel. Although Oviedo was not a naturalist. The limited similarity between the Philippines and this Panama Tall (0.475) is due to the extremely small founder population of the Panama coconut. This is the genetic basis for the affirmation that both the Panama and Mexican Pacific coast coconuts were introduced from the Philippines. although they are geographically closer. Some 30 years later. and possibly different selec- tion pressures since introduction (Baudouin & Lebrun. …” (Columbus. but the whole General History was only published in 1851–1855. (2011) . coconuts. 3 Multi-locus allelic composition of the Philippine. 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. when sailing near Puerto del Principe on the north coast of Cuba: “There was a beautiful meadow and many very tall palms. 3). palms also thought to be coconuts were reported on the Pacific coast of Panama by Gonzalo Fernández de Oviedo y Valdés. He published a first Summary entitled “Natural History of the Indies” in 1526. 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. especially to supply the toddy market in the 17th century (Zizumbo-Villarreal. whereas the allelic diversity found in Mexico is quite similar to that found in the Philippines (Fig. on the 17th of November 1492. edited by J. 2009). C.A.

1977). as Patiño (1963. 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. as Spanish and Portuguese explorers visited new parts of the New and Old Worlds. The case of coconut is emblematic of this flow of information. 1963. However. 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. information was flowing into the Iberian Peninsula at a great rate. It is important to remember that they did not explore the same areas because of the Treaty of Tordesillas. based on his travels in Asia between 1501 and 1508. The expedition explored only a small area and returned without mentioning coconut.Coconuts in the Americas clear that only part of the information was obtained personally. which effectively prevented Spanish mari- ners from sailing to Asia (until 1580). on Panama’s Carib- bean coast–heard of coconut) and Rodrigo Colmenares (ship pilot and interpreter– . although he recognized that many documents have been lost and some were unavailable to him even after 40 years of effort. 1977. Based on Patiño’s analysis. Oviedo and others of the period used information from Asia and the Americas in their reports without specifying the origin of each detail. Patiño. the first com- plete information available in Iberia about coconut uses in Asia was published by Ludovico di Varthema (1510). 1501—King Manuel of Portugal wrote about coconut in detail in a letter to Queen Isabella and King Ferdinand of Spain. 1513—Vasco Núñez de Balboa crossed the Isthmus of Panama and discovered the Pacific Ocean. 2002). Hence. Both Patiño and Zizumbo & Quero cautiously accept that Oviedo’s reports are correct. extolling the prime value of coconuts as a source of drinking water and cordage for sailing ships (cited by Harries. after leaving seedlings in the Cape Verde islands. although some may have seen the nuts in Lisbon or other Iberian cities. although numerous documents of this expedition have been lost. as Vasco de Gama was the first to bring coconut to Portugal in 1499. Patiño (1963. as the Spanish crown demanded information from all explorers who received crown support. 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. 1514–1515—Alonso de la Puente (royal treasurer in Darien. which effectively divided the non-European world between Spain and Portugal. 2002) analyzed both of these documents and the origins of the information that Oviedo used to prepare his Summary and General History. and part was obtained by interviews and correspondence. Before and during the period that Oviedo was active as historian. 2002) emphasizes. a common occurrence before modern citation methods were developed. 1492—Discovery of New World 1494—Treaty of Tordesillas divided the world outside Europe between Portugal (eastwards) and Spain (westwards). where they grew to reproductive age and were distributed to the Americas (Harries. Zizumbo-Villarreal & Quero (1998) analyzed Oviedo’s Summary and General History to determine if coconut was present in Panama at the time of European conquest. this is the time line for the early Spanish references to coconut in Panama.

1539—Alvaro de Guijo (resident in the City of Panama) sent nuts to Hernán Cortés de Monroy y Pizarro (conqueror of Mexico). as well as some already germinated. Near the Cape of Corrientes (Fig. Natá was reputed to have an abundance of coconuts. Some of the riper ones can be sown by placing half the fruit in the ground. to the southeast of the current City of Panama. 1). 1526—Juan de Cabezas (pilot) discovered Cocos Island (Fig. but does not mean that it was indeed coconut. C. and I send you two dozens of ripe fruits. reported by the natives to germinate and grow spontaneously along the shore. 1944). 1. but not in the Summary. although they also reported young plants were transplanted. canon of Cape Verde (Bruman. 1524—Francisco Pizarro González (mayor of the City of Panama 1519–1524) led an expedition that explored the South American coast from the City of Panama to the northern limits of Colombia. Panama insert). The fact that they thought it to be the same as the coconut from India is significant. they reported “a large quantity of coconuts” at La Candelaria Bay. another official crown historian. possibly the first coconuts on continental America. If you like them. along with advice about how to plant them. In his letter that accompanied them. Patiño suggests that Espinosa may only have realized the importance of his observation after learning about coconut in Asia. Clement et al. Anglería states that coconuts were observed in three places along the Pacific coast: Chimán. further to the west at the frontier with modern Costa Rica (Fig. The explorers Puente and Colmenares reported on the coconut (that they thought to be the same fruit as cultivated in Calicut—the 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. 1944). Anglería also reports that “Some think that sea currents bring the seeds of these trees from unknown regions. and Burica. Anglería’s report was first published in 1530 (translated in 1944). I can send more another time. Brazil (Bruman.R. but Oviedo knew Anglería and had access to his report. to the west of the City of Panama. which explains why this information is in the Summary. 1549—Coconuts from the Cape Verde islands (Portuguese) planted in Puerto Rico (Spanish). 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. In fact. where he reported “Many beautiful and large mameys (Pouteria zapota) and many palms with the large coconuts. 1) and provided information used by Oviedo in the full General History. . Chimán is thought to be the first place that Europeans saw coconut and Patiño even suggests that Balboa’s expedition may have seen it there also. attributed to Diego Lorenzo. but Zizumbo-Villarreal & Quero (1998) caution that this possible introduction to Mexico may not have been successful. Natá. Hernán Cortés left for Spain in 1541 and never returned to Mexico.” 1519—Gaspar de Espinosa y Luna (explorer. …” It is not clear exactly when Anglería obtained this information from Espinosa. Guijo wrote: “I have heard that you do not have this fruit we call cocos.” This letter is cited in Bruman (1947). in late 1516 in Spain. 1553—Coconuts from the Cape Verde islands planted at Bahia. saw coconut) visited the Pacific coast of Panama and were interviewed by Pedro Mártir de Anglería. so I sent a boat to a place along the coast to collect some. which are reported in Oviedo’s General History.

Nonetheless. The Archaeological Evidence The central Pacific coast of Panama has a long cultural history. Dickau (2010) mentions finds at La Pitaya.Coconuts in the Americas 1565—Philippines to New Spain (Mexico) return trip was accomplished inde- pendently by Alonso de Arellano and Andrés Urdaneta. 1. subsistence still included gathering. Coconuts. 1580—Treaty of Tordesillas became invalid when Portugal was ruled by Philip II of Spain. 1980. permitting the supposition that some of the first reports may have confused this Attalea with coconut. 1569—Alvaro de Mendaña introduced coconuts from the Solomon Islands to Colima. (Dickau. Attalea butyracea (Mutis ex L. 2010). 1998). Astrocaryum and Bactris are spiny and like Elaeis have fruits that are morphologically clearly distinct from coconut. which are microscopic silica bodies found within and between cells in many plants.5–8. Cooke and Ranere. at the frontier with Costa Rica (Fig. horti- culture started to become important.f. A. and several coastal . the Gran Choclé and Gran Chiriquí 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. 2010).000 years before the present (BP) (Piperno & Pearsall. By the late Pre-Ceramic Period (7000–5000 BP). Mexico (Sevilla de Rio. Boer. Elaeis oleifera (Kunth) Cortés and Astrocaryum sp.. expanding rapidly through the Early Ceramic Period (5000–2500 BP) until most subsistence was horticultural by the Middle Ceramic Period (2500–1500 BP).5 cm wide) that look somewhat like very small dried coconuts. The Ceramic Period extended until European Conquest (500 BP). Phytoliths. 2010).. The macro-archaeobotanic remains from these regions include Acrocomia aculeata (Jacq. and the palm has a similar stature to that of coconut (Henderson et al. but Acrocomia. It is important to mention that most of this work was carried out in the central and western parts of Panama. Panama insert). especially of palm fruits and other useful plants. Large numbers of endocarp fragments of Attalea butyracea are reported at numer- ous locations in the Gran Choclé and Gran Chiriquí regions of western Panama from 7000 BP to European conquest (Smith. became a regular item but were not always mentioned in the ships’ manifests. 1974). and the Manila- Acapulco commercial route was established by 1571 and continued until 1815 (Fig. 1992. a small coastal island about 50 km east of Point Burica. 1995).) Lodd. The macro-and micro-archaeological records show that palms were very important to the subsistence of the Native Americans in the Gran Choclé (central Panama) and Gran Chiriquí (western Panama) regions. Bactris major Jacq. of palms are also abundant in the same region during the same period (Dickau. Specifically.) Wess. 1). with human activities present in the archaeological record since 11.5 cm long by 3–4. ex Mart. In terms of the Pacific coast. carried for consumption by passengers and crew. butyracea grows on the slopes above beaches and river courses along the Pacific coast of Panama in many places today. All of these species are members of the Cocoseae tribe within the Arecaceae. Dickau. The Attalea butyracea is especially important to our discussion because it has relatively large fruits (4..

as observed by Morcote-Rios and Bernal (2001: 342). Cerro Juan Diaz). The Chibchan languages of Panama include (from west to east) Teribe. 2010). Creole. This is a remarkable anomaly given Native American uses of a wide variety of palm species. including endosperm. 2006a. even if it were growing in relatively remote locations. and Amerindian languages affiliated with two language families. This dramatic lack of evidence is a strong reason to doubt the accuracy of the first Spanish reports. paleobiolinguistics facilitates reconstruction of terms for plants and animals in the vocabularies of their ancestral or proto-language. 1995). Monagrillo. The modern languages of Panama include Spanish. 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. La Mula Sarigua. as the archaeological record tends to be determined by sampling intensity. Buglere. In comparison. Ngäbare. although the native Panamanians apparently were familiar with its propagation (see Time Line 1514– 1515 above). then. Several archaeological sites are close to historically mentioned sites. Chibchan and Choco (Fig. In summary.R. even though “the woody endocarp of the coconut is an appropriate material to be preserved at archaeological sites in wet environments. Reconstructed words for species are indicative of their substantial significance to speakers of proto- languages. but did not use it?” Patiño states that there would have been no reason for coconut not to be culturally assimilated by the native population quite rapidly. or at least to assume coconuts could not have arrived in Pacific Panama more than a few years before European Conquest. but she does not report coconut. but none are exactly at those historically mentioned sites. Clement et al. … Would one have to conclude that these Nations met this palm at least four generations ago. so future work may reveal new information. Hence. b. the absence of evidence is not necessarily proof of absence. The approach.” However. none of the archaeological sites reported are close to Chimán or La Candelária Bay. Clement. sites about 50 km southwest of Nata (Vampiros. and even the domestication of peach palm (Bactris gasipaes Kunth. . 4). The Ethnobotanical Evidence Although many large palms have been important in Native American subsistence in Panama since well before European Conquest. C. no archaeological evidence exists for coconut in Panama before European conquest. 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. there is no record of use of coconut by native Panamanians at the time of conquest (Patiño. and luck. artifact preservation. By comparing words for biological species in languages of the same language family. 2002). Patiño (2002: 253) writes: “There is no doubt that the inhabitants lacked the tradition about the use of this plant. the Gran Darien region of eastern Panama is much less well studied.

). 4). maize (Zea mays). This does not mean that no indigenous languages were spoken there in the past. Other Chibchan languages are spoken in Honduras. Plants that reconstruct for Proto-Chibchan. A large portion of Panama is empty of contemporary Native American languages (white areas of Fig. Comparative evidence is such that a word for coconut is not reconstructable for any chronological stages of Chibchan and Choco language families. 2010). if any. and sweet potato.Coconuts in the Americas Fig. very little. manioc. 2009) and Kuna. in both Proto-Chibchan and Proto-Choco. bottle gourd (Lagenaria siceraria). notably the western Pacific coastal area and central parts of the country. On that map much of the contemporary empty regions of Panama (Fig. and Colombia. No words for coconut reconstruct for these prehistoric proto-languages. For this study. a term for coconut cannot . maize. both having dialects spoken in Panama and Colombia. Kaufman (1994) pub- lished a time-of-contact language-distribution map for the Caribbean region that in- cludes southern Central America. all Chibchan languages. manioc (Manihot esculenta). Doraske. evidence bears on the language or languages of these archaeological populations. Nicaragua. Constenla Umaña. 1981). The smaller Choco language family includes Emberá and Woun Weu. 2011). Archaeological sites in parts of the area attest to Native American occupation. sweet potato (Ipomoea batatas). including coconut. 4) is identified as being filled by speakers of Mobe. Those reconstructed for Proto-Choco (c. 2258 BP) include Banisteriopsis caapi. guava (Psidium guajava). hog plum (Spondias spp. suggesting that the species was not known to their speakers. include cacao (Theobroma cacao). Costa Rica. However. 4 Map of the distribution of modern languages of Panama created with Ethnologue (Lewis. For example. we undertook a paleobiolinguistic investigation of domesticated and useful plants. including those closest to the time of the European conquest. spoken at the latest around 4400 BP (Holman et al. and tobacco (Nicotiana tabacum) (cf.. perhaps as late as the 19th century (Locascio. and Bokota (in the west) and Kuna (in the east). cotton (Gossypium hirsutum).

were extracted from lexical sources available to us for Chibchan and Choco languages (Table 1). by borrowing words for these things from languages of the people that introduced them. Words for coconut in the 12 languages robustly suggest that the plant was introduced by Europeans into southern Central America.R. and artifacts. thus. both of which are so phonologically dissimilar to one another as to exclude the possibility of cognation and. All languages except one or possibly two have borrowed words for the plant and its fruit from a European language. Speakers of native lan- guages of Latin America typically have named newly encountered items. 1999). animals. including plants. In fact. but comparatively rare among native languages of Latin America (Brown. 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. in this case from Spanish and Portuguese (Brown. nine are loans based on coco. For example. Table 1 Terms for coconut in Chibchan and Choco languages Coconut Term Language Family Location siahuá Boruca Chibchan Costa Rica koko Bribri “ Costa Rica kó ko Chimila “ Colombia koko Dorasque (extinct) “ Panama coco Guatuso “ Costa Rica ógoba Kuna “ Panama köko Ngäbare “ Panama kukunúp Rama “ Nicaragua kokoha Paya (Pech) “ Honduras kóko Northern Emberá Choco Panama k’ok’o Epena “ Panama kök Woun Weu “ Panama . All but two terms in the 12 languages are unambiguous loans from European languages. 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. a daughter language of Proto-Choco spoken at the latest around 875 BP. either Spanish (coco) or English (coconut). be retrieved for Proto-Emberá. all of the terms for the referent in available lexicons are of non-native origin. 1994. 1999). However. in the Bachajón dialect of Tzeltal. and one (in Rama) is a loan from English (coconut). 1999: 92–104). On the other hand. only rarely have native terms for indigenous things been replaced by Spanish or Portuguese loanwords (Brown. French. In fact. 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. a Mayan language of southern Mexico. the possibility that a word for coconut with such reflexes pertained to Proto-Chibchan. Twelve terms for coconut. presented in original orthography. Clement et al. 1994. The two exceptions are Boruca siahuá and Kuna ógoba. C. and Russian intruders. This practice has been very common in languages spoken by Amerindians influenced by English. Of the remaining 10 terms.

We have also undertaken a preliminary paleobiolinguistic survey of many language families of Latin America for evidence of pre-Columbian coconut. i. In conclusion. designating Acrocomia aculeata. a word that later became referentially extended to the introduced coconut. and no term for A. and suggests either that coconut arrived immediately before European Conquest.250 years BP as suggested by Baudouin and Lebrun (2009). or in addition to.” a usage almost certainly motivated by the resemblance of the European sheep to the native deer. rather than 2. 1935 BP). coconut. A Preliminary Summary of the Evidence The genetic evidence concerns modern coconuts. no such terms are apparent. No term for coconut is listed in Margery’s (1989) exceptionally thorough dictionary of Cabécar. the Proto-Chinantecan word designated A. aculeata. which. Rensch (1989:78) reconstructs *há:H (H=high tone). to borrowing. Evidence for this comes from Cabécar. In either case. Plausibly. like Boruca.Coconuts in the Americas the introduced sheep is tumin čix. this term originally denoted a native palm similar to the introduced coconut. será. both the genetic evidence and historical records need further study. Zizumbo-Villarreal & Quero 1998). assigning to it the gloss coconut. a survey of reflexes of this hypothetical word in various Chinantecan languages shows that some of these denote Acrocomia aculeata rather than. If so. is a Chibchan language of Costa Rica. aculeata is found in Quesada Pacheco and Rojas Chaves’s (1999) comprehensive dictionary of Boruca. this suggests the original referent of the Boruca term was A. 1963. Mexico. 1983). However. . 2002. Cabécar contains a word similar to the Boruca term. The only word for coconut of the 12 languages that clearly is not a European loan is Boruca siahuá. Indeed. ethnobotanical or linguistic evidence suggests that the caution was warranted. mainly focusing on the reconstructibility of terms for the plant in proto-languages. For this ancestral language. The phonological similarity of Boruca and Cabécar words may be due to cognation or. so says nothing about the historical presence of coconut in Panama. This lack of interacting evidence is an unexpected result when applying de Candolle’s and Vavilov’s methodology. paleobiolinguistics evidence assembled to date fails to provide support for the prehistoric occurrence of coconut for any region of Latin America..e. The possible exception is Proto-Chinantecan (c. like the Boruca term discussed above. this evidence strongly suggests that the modern occurrence of the plant in the region is accountable to European introduction in historical times. This suggests that. if not. but the clear absence of archaeological. a palm whose fruit is fed to cattle and occasionally consumed by humans in the Cabécar region. aculeata. This finding concurs with the work of Merrill (1937). the most salient mammalian herbivore known to Tzeltal speakers (Witkowski & Brown. and that its reflexes were referentially extended to the introduced coconut. a pre-Columbian presence of coconut in Panama and surrounding areas is not attested by paleobiolinguistic evidence. The historical record had previously been accepted with caution (Patiño. whose contemporary offspring languages are spoken in northern Oaxaca state. 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. or it arrived after European Conquest. literally “cotton deer. With only one possible exception. Either way.

including the Philippines. This data set was not designed primarily to identify the origin of the Pacific coast Panama Tall. that sampling in Oceania. inland. especially those from Mexico and Panama. is not well represented (Table 2). Montpellier. although Baudouin and Lebrun (2009) used it this way with interesting results. such as coconuts in Panama. we examine the samples used in the genetic studies. 1. 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. Hence. 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. 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. includes 5 from Oxtapacab. contrast between shaded areas and the Pacific Ocean). C. where coconut is a major crop today. taken to Jamaica.R. then Ivory Coast—considerable introgression with Indo-Altantic Panama Tall Bowden 10 Taken to Jamaica Panama Tall Costa Rica 19 Costa Rica—minor introgression with Indo-Atlantic Panama Tall Monagre 19 West of Nata along coast—minor introgression with Indo-Atlantic . Yucatan. 3 plants with fruit similar to Baybay Tall Pacific Tall Michoacán 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. the sample used will affect results. Melanesia) in 1569—fruit similar to Rennell Tall. of CIRAD. Clement et al. Even the Philippines. 3 plants with fruit similar to Philippine varieties Pacific Tall Guerrero 11 8 plants with fruit similar to San Ramon. however. Mexico and the Philippines. France. It is worth mentioning. the French Agricultural Research Center for International Development. Genetic Sampling and Expanded Analysis In any study of the genetic relationships among populations. (2011) studies. does not yet represent the variability that is present in the region either (Fig. Mexico Panama Tall Agua Dulce 13 West of Nata.

the most important coconut disease in the Americas. which explains part of the Indo- . Note also that these middle-western regions are precisely those where Native American populations have disappeared (Fig. so one might expect coconut introductions from other localities during the colonial and modern periods. they had been collected years earlier and taken to various countries before having their DNA extracted (Luc Baudouin. from orchards established in the 18th century. Mexico and Panama (Table 2). 1995). The Pacific Tall of Michoacán was intro- duced by President Lázaro Cárdenas. 2011). The Nuxco plantation was established in the 1950s. Rather. The 11 Rennell Tall-type plants are derived from a plantation established in 1890–1900 at the margins of the Coauhuayana River in Tecoman. as this variety originated in the Indo-Atlantic group of varieties (Harries. but some plants have been joined into “state-level” varieties without due consideration of their morphology (Table 2). pers. nor is the Mexican Atlantic Tall. 2011). Guerrero state (Zizumbo- Villarreal & Colunga-GarcíaMarín.. as these are unlikely to have contributed to this Panama Tall’s genetic composition. Dwarf varieties from the Philippines are not included in the table. we have not found a history of these introductions. All of these varieties have clearly different microsatellite profiles when compared to the Pacific coast Panama Tall (Gunn et al.. 1977. Thus. The sample of Mexican Pacific Tall coconut varieties used by Baudouin and Lebrun (2009) and Gunn et al. covering the Gran Choclé and Gran Chiriquí regions.. For example. possibly from progeny of the early introductions to Acapulco from Philippines. com. the sample used by Baudouin and Lebrun (2009) and Gunn et al. and was initially established in plantations in 1937–1938. However. They may be progeny of the introduction made by Alvaro de Mendaña from the Solomon Islands in 1569 (Zizumbo-Villarreal & Colunga-GarcíaMarín. (2011) did not include plants from the Gran Darien nor the Gran Chocó (present day Pacific coast of Colombia). 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. No historical record of these introductions exists. but they agree with the Philippine location of probable early introductions (see below). 2011: Table S1). 1974) it was resistant to lethal yellowing. Numerous samples of Panama Tall from the Pacific coast were taken into the Caribbean during the 20th century and were then distributed elsewhere. The seed originated in the environs of the city of Colima. 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. the western portion of Michoacán and eastern Jalisco states. 4). respective- ly.Coconuts in the Americas The varieties in the GCP data set that interest us most are those from the Philippines. where historic sources place the pre-conquest presence of coconut at Chimán and Cape Corrientes. When the Maypan hybrid (a cross between the Malayan Dwarf variety and the Panama Tall) was produced in Jamaica (Harries & Romney. not all coconut populations along the Pacific coast of Panama were sampled. 2001). apparently from the Philippines. 2001). (2011) includes several with known origins and dates of introduction. and the Panama Tall became very important in the Caribbean (Harries. the sample is biased towards the middle-western portion. Colima state. It is represented in the vast majority of plantations in Colima. Additionally. Unfortunately. This is the most representative variety between Acapulco and Lázaro Cárdenas Port. the various samples of Panama Tall from the Pacific coast listed above (Table 2) were not collected directly in Panama for the GCP study. Gunn et al.

(2011) Table S1 . Mexican and Panamanian coconut varieties studied by Gunn et al. Pandan. Hence. Because sample sizes are small and all bootstrap confidence levels are weak.. C. the length of each color in each bar is the proportion of each group in that plant. The modern Pacific coast Panama Tall is also morphologically similar to San Ramon (Vargas & Blanco.000 bootstrap iterations to obtain a preliminary idea of relationships. 5 Group assignment at K=5 for the Philippines. Part of this is due to the very small sample sizes (Table 2). this preliminary analysis suggests an introduction of coconut to Mexico that then influenced other introductions and was also introduced to Panama. the Neighbor Joining algorithm. Redrawn from Gunn et al. and used Nei et al. Atlantic alleles found in some plants that is evidently due to introgression (see Gunn et al. 5). with the very small founder event detected by Baudouin and Lebrun (2009). Clement et al.R. and 1. Baybay. All the other Mexican varieties appear to be derived from this relationship and are a sister group to the Panamanian varieties. (2011). which is curious because this Mexican variety may have originated in the Solomon Islands (Table 2). 2011: Table S1). but neither took the opportunity to look more closely at the variability within the countries involved. The two genetic studies were both well executed for the questions that they asked. we extracted the microsatellite genotypes for the Philippines. Nonetheless. we do not present the dendrogram. 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. especially in the Philippines.’s (2011) Table S1 (Fig. but also Mexico and to some extent even Panama. we extracted the Structure assignments at K=5 from Gunn et al. because this will determine the reliability of the relationships found. Four Philippine varieties (San Ramon. However. Mexico and Panama from the Global Challenge Program dataset. The colors represent the five groups identified by Structure analysis. Tagnanan). 2000). Hence. a look at these relationships can offer ideas for new studies.’s (1983) genetic distance. the size of the slice is the proportion of each group in that country. there seems to be a relationship between the Philippine San Ramon variety and the Mexican Pacific Tall of Colima. one of which (Baybay) is morphologically similar to the Pacific Tall of Guerrero (Table 2). To look even more closely at how the Panama Tall may be related to the Philippine and Mexican varieties. Fig.

1978. 2009). it seems reasonable to ask how the San Ramon type might have arrived in Panama. the San Ramon coconuts in Mindanao were highly regarded. Jalisco. with all Philippine and Mexican varieties except the Mexican Atlantic Tall in one group. wrote: “There are very big ones [coconuts] which would measure more than one azumbre” (~ 2 l) (Alzina 1668). We then took the GCP dataset and analyzed the Philippine. as the northernmost harbor. Hill. so even though San Ramon is identified as the most probable ancestor of the Pacific coast Panama Tall. ethnobotanical and linguistic evidence for coconut in Panama at conquest.3. 2005). who resided in the Visayas from 1634 to 1667.Coconuts in the Americas Pandan. 1) that had been suggested by Safford (Small. 5. both sailed from Cebu and not from Manila. in particular with the San Ramon type in the Philippines (Vargas & Blanco. 2000. From the Structure analysis for the Mexico Pacific Tall of Colima. 2005). even though the high proportion of South Pacific confirms the origin of Colima in the Solomon Islands. At the head of the Sulu Sea. Cebu in the Visayas was not far from Mindanao and. Zizumbo- Villarreal & Quero. returning independently. 1978. In 1564 an expedition to establish a Spanish settlement set out from Puerto de la Navidad (Barra de Navidad). Mexican and Panama varieties with Structure 2. Considering the lack of archaeological. Hubisz et al. as is also visible in Fig. The Manila-Acapulco galleon route (Fig. 2012). it has not been closely examined until now (see also Harries.. An agricultural observer in the seventeenth century. Although it had been accepted by Merrill (1954) and others (Harries. Zizumbo-Villarreal et al. 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 . although post-conquest. and there is some admixture of Atlantic with Panama. However. 1929). 1998. which is significant because there were superior coconuts in that region. because it already had trade links with China and Japan and. when the production of copra became commercially important in the Philippines at the beginning of the 20th century. 2000). Alonso de Arellano and Andrés Urdaneta. on the first occasion in 1565. The similarity of Pacific coast Panama Tall and named varieties in Southeast Asia has long been noted (Harries. it became the departure point for the galleon route. setting a course for Cebu in the Visayas. Nonetheless. it is possible to hypothesize that the proportions of Panama and Papua New Guinea assignments explain its similarity with San Ramon. but is less clear about showing relationships with Panama. 1929. Zizumbo-Villarreal et al.. The San Ramon variety had some Atlantic assignment. numerous plants in this small sample have enough Panama assignment to suggest that the Panama Tall is derived from the Philippines via Mexico. Mexico to avoid sailing through Portuguese waters. The Mexican varieties also have considerable proportions of Panama assignment.’s (2011) parameters. is the obvious candidate. all Panama varieties in one group. following Gunn et al. Father Francisco Ignacio Alzina. This new analysis strongly supports the Philippine – Mexico connection. although less than the Philippines overall.3 (Pritchard et al. given the fact that only the Philippine varieties represent- ed the whole Pacific Ocean. Tagnanan) have numerous plants with considerable proportions of Panama assignment. other varieties could have contributed and would not be easily detected because of the extremely reduced size of the Panama Tall founder event. This generated three groups (data not shown). where Magellan had landed 41 years previously. and the Mexican Atlantic Tall in the third group. 1971. Manila subsequently became the premier city of the Philippines.

Luzon and a report that “the largest nuts in the world are produced around Lingayen Gulf.R. more easily. we revisit Oviedo’s account because. However. Zizumbo-Villarreal & Colunga-GarcíaMarín. Colombia (Fig. and is still today. The Barra de Navidad lagoon borders Colima province. . However. rather than the once yearly supply from Manila. later introductions of larger numbers of seednuts intended only for planting would have been made. would have been the most convenient for taking deck cargo on board. The possibility that coconuts were carried to Mexico in 1565 has previously been discounted (Bruman. would be disseminated from Colima southwards as far as Peru. to the north of Manila. Clement et al. which became. Zizumbo- Villarreal and colleagues in Mexico are collecting and analyzing the Pacific coast Tall varieties there. for example. Arellano’s 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. It also seems reasonable to suppose that the subsequent Manila-Acapulco galleons would carry the same sort of coconuts. the Panama Tall coconuts could have come from one or more locations. in August 1565. 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 . 1914). although it was previously analyzed by Patiño (1963. who tapped “tuba” for fermenting to “coconut spirits” for consumption in Mexico (Zizumbo-Villarreal 1996. 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. 2012). at the end of a four-month voyage to Puerto de la Navidad. Confirmation that seedlings were planted in Mexico comes from the activity of skilled Filipino toddy tappers in 1580. As first the tuba market and in the 20th century the copra market expanded. Allen (1965) and . but still represent the San Ramon and closely related varieties. C. by plantings near to Manila. including Gorgona Island. 2002). 1914: 537). This could be done. Thus. because coconuts were not recorded on the list of provisions. Botanical and Historical Questions Given the doubts about the historical record. The early germination of this type—more than 75 % in 105 days (Harries. Barrett cited by Smith & Pape. 1945). small sample from this source might explain the extremely narrow genetic base of the Panama Tall reported by Baudouin and Lebrun (2009). 1914). . A single. the center for coconuts in western Mexico.W. is famous for producing coconuts of immense size and are of great use to planters as seed nuts…” (Bardy. These locations. 1981)—would have meant that. Ramon…” (Copeland. Luzon Island” (O. either by arranging for them to be collected in Mindanao or the Visayas for transshipment or. 2008). there would be seedlings ready for planting. This would explain Copeland's note that San Ramon coconuts were in general cultivation in the coastal district of Pangasinan province. 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. 1).

since the Native Americans had numerous other good fibers. (1998). as big as the fist of a closed hand. and reported that coconut is the size of a man’s head. is the coco. 2002) discards the first problem by pointing out that Oviedo finished describing the cane palm. is a meaty part in width like half the thickness of the small finger of the hand. quite unlike the Panama Pacific Tall or San Ramon varieties. Bactris). this being a genus of large palm trees. The crust is hard. “This fruit which is within that burlap. as noted above with respect to the lack of ethnobotanical evidence. While Patiño and Zizumbo-Villarreal & Quero looked at the similarities and cautiously accepted that Oviedo had seen coconuts. and more or less. 1995) and Cocos nucifera is monotypic. We consider their opinions and identify two properties. Allen saw differences and suggested that Oviedo may have regarded them as aberrant and inferior sorts. …” A century before Columbus crossed the Atlantic. 1917. so this kind of information was available to historians in Iberia at the time that Oviedo wrote. 131 palm species or sub-species were attributed to the genus Cocos.” 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. Astrocaryum. as indicated above. what we may be seeing in this sentence is the Spanish naming of native American palm species. although many have spines that are hard to miss (Acrocomia. Bactris major.” This part of the description is somewhat at odds with the original description of the size of the fruit. parts of the description can match other palm genera and parts of the information may have come from Asia rather than the Americas. The following sections of Oviedo’s account (as translated for use by Allen. it is a sort of round nut. …” This is the first sentence of the account and suggests that numerous Neotropical palm fruits were called cocos. The description appears to be a justification for lack of use. Inside. and today grow on the hills above the beaches of Panama. on the page where he started describing coconut. or as thick as a writing quill of the kind common to geese. as was Varthema’s (1510) account. attached to the crust of that nut or coco. for here there is much cotton and henequen and cabuya to supply such necessity for cords. Patiño (1963.. Oviedo’s account of coconut has been questioned on a number of occasions because the illustration does not match the description. with pinnate leaves. and some are elongated. found only in genuine coconuts. The spineless ones are now in Attalea. the Arab Ibn Battuta (1929) visited East Africa and India. and whose leaf is of the same kind as that of the date palms. . “These trees or palm trees put forth a fruit which is called coco.… Altogether. Hence. which Oviedo did not mention and apparently did not know about. the DNA data match between the Panama and the Philippines coconuts raises fresh concerns. which has the appropriate stature. “There are other palm trees whose fruit are called cocos. Over the next centuries. which is at odds with our linguistic analysis.Coconuts in the Americas Zizumbo-Villarreal & Quero. 1965) are worth transcribing and commenting on. “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. and some as big as two fists. These other palms are now assigned to different genera (Beccari. Henderson et al. and as thick as the width of the inscription [title] on a Castillian silver real [coin]. fruit that look somewhat like small coconuts (see archaeological evidence above). so if these were coconuts they had small nuts. such as it is on the tree. Some of them have statures similar to coconut. it has a much greater bulk than a man’s head. such as in the Levant.

and it is as white as a cleaned almond and better tasting than almonds. but will be contrasted below with another description. Moving on from Oviedo. is a place taking up the remaining part or entire quantity of the coco. However. although both read it since they mention Oviedo going from Burica to Nicaragua. why weren’t they also sent to Madrid—at least to show at court? Given the lack of this kind of display. The seeds of A. I was in the province and headland of Borica. but they are perfectly edible. Oviedo’s use of the word aborrecí (loathe) is a surprising reaction—more people would agree with Charles Darwin (1860: 407): “After walking under a burning sun. It is also possible that this is a mixture of information from Asia (size. including coconut. 2002) nor in Zizumbo-Villarreal & Quero. as pointed out by Allen (1965). “After I wrote the report I have mentioned. whose fruits can superficially be confused with very small coconuts.” This statement was not cited in Patiño (1963. . I do not know anything more delicious than the milk of a young cocoa-nut. and others did as I did and said the same thing as well.…” This is a good description of the endosperm of a coconut. water. and came to loathe them. That leaves Attalea butyracea. and more or less. typical of Panama Tall (Harries. it seems likely that 20 years after the first report there still were no real coconuts on the Pacific coast of Panama. 1981). Coconuts on the open deck of a boat would germinate rather than rot and. there is a logical question related to the 1539 letter to Hernán Cortés (see Timeline above): If coconuts were interesting enough to send from Panama to Mexico. and any palm seed. can go rancid if stored in conditions that do not allow germination but do allow respiration. Oviedo does not mention the sound of the water splashing in the cavity of a mature coconut when shaken. C. Note also Oviedo’s choice of “ate” rather than “drank. so the spiny species can be discarded. “This is the fruit proper of the coco and what is edible. sweet and very edible haustorium (or “apple”) that would immediately identify a real coconut. Like coconut. So. this is a good description of coconut water in an immature coconut. endosperm. and I ate some of these cocos and carried many with me to Nicaragua. butyracea do not have liquid endosperm when ripe. flavor. …. It is eaten the same way peeled almonds might be eaten. 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. Some individuals do find the kernel indigestible and may become tired of it. as Allen pointed out. and of smooth taste to the palate. but Oviedo’s account does not suggest this. in proportion to the bigness or size of the coco:…” Again.” as it suggests that they were fruit with little water. “By way of pith or marrow of this fruit which is in its middle.) and the Americas. as the water is naturally absorbed when the nuts mature. but that is not usually a group phenomenon. some people may not like the flavor of the seed. Clement et al. they would be regularly carried for refreshment on any coastal craft. Even non-specialists would note and comment on spines on the trunk and leaf petioles. and as much as would fill the shell of the egg of a hen. if they had been present in Panama at this time. 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). etc. but Oviedo does not mention the soft. (1998). full of a most clear and excellent water.R.” Perhaps the water had been absorbed by early germination. Nor does he say if they were being taken to Nicaragua for planting.

wash- ing onto a suitable shore. two questions arise: Could they have floated across the Pacific Ocean (perhaps via intermediate islands). but the water temperature would have been significantly higher than actual sea temperatures. 1948). while Dennis and Gunn (1971) argue for car- riage by man. given the genetic relationship between the Pacific coast Panama Talls and coconuts in the Philippines. we look at the hypotheses about how and when coconut arrived along the Pacific coast of Panama. Computer simulations of Pacific wind and surface current direc- tions and speeds. but the microsatellite information suggests that the original sample was very small and did not capture the full San Ramon genetic profile. The hypothesis of American origin was reinstated by Cook (1910). nuts were floated in barrels of sea water. Bruman (1944) and Purseglove (1972) argue for natural dispersal. A crucial question is the period for which coconuts will remain viable when floating in the sea. 1980). Hypotheses about Coconuts in Pre-conquest Panama Five hypotheses have been presented to account for the historical observations. This was held by de Candolle at first (1855). With this summary in mind. 1980). and thereby decisively debunked. For example. But. 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. three of which (2–4) are not mutually exclusive. striking root and growing. The lower temper- atures of floating coconuts delay germination compared with nuts at average ground level temperatures (Ward & Allen. 1941. These are: Hypothesis 1 Coconuts originated in the Americas. the longest period any nut floated and remained viable was 110 days (Edmondson. or must they have been carried? Both cases have been argued by many scholars. This hypothesis is based on the fact that coconuts can disperse by floating over some distance. In two experiments to test the viability of nuts floating in the sea. The San Ramon variety was probably introduced into Mexico early via the Manila-Acapulco route. Ward & Allen. strongly criticized by Beccari (1917) and Merrill (1937).Coconuts in the Americas 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. Hypothesis 2 Coconuts floated from mid-Pacific islands on one or many occasions from ancient times to the present day. but discarded later (1883). islands and coasts allowed tests of the possibility of . This suggests that the maximum flotation period may be on the order of four months. Gunn (2004) placed a final nail in its coffin with a molecular genetic phylogeny of the Cocoseae. In an experiment at the Coconut Experimental Station in Sulawesi in 1931 (Reyne.

The voyage from Rapa Nui to the American coast would take about one month. The conclusion must be that coconuts cannot drift to the Americas within any rea- sonable period of viability. 2009: 218). Ward & Brookfield. and combined wind and current forces. 6 and 8 months. and would be expected to plant some coconuts in situations where they were not already growing. Clement et al. Experiments were conducted for drift periods of 4. Computer simulations of eastbound voyages from Polynesian islands towards America show that such voyages are possible from such starting points as Samoa.588 simulated drifts from these islands. Fitzpatrick & Callaghan. based on the lack of similarity between the microsatellite profiles of differ- ent Polynesian coconut varieties and the Panama Tall variety. 1992). coconuts did not grow in the Galapagos at the time of early European contacts. 1) (Levison et al. How- ever.R. 1992: 473–4). C. coconuts drifting across the Pacific from several possible starting places (Levison et al. From Tonga or Samoa the . or plentiful. 37 % of drifting nuts did reach the Galapagos group. Irwin. showed that nuts floating from Polynesia had no chance of drifting to the Americas from that part of Polynesia. eastern French Polynesia and Rapa Nui (Easter Island) (Fig. in the ex- treme east of French Polynesia (Fig. no nuts made the crossing.. Tonga.. 1973. It is impor- tant to note that. and from the Marquesas two months (Irwin. variations were used to cover current only. the shortest crossing being in 178 days from Christmas Island. In “wind only” experiments. No coconuts drifted to the American coast in any of the 6. only 1 % reached the Galapagos. 1). the nut’s weight increases. and Motuiti in the northern Marquesas. Fitzpatrick and Callaghan. 1992: 163–4. wind only. 1992: 214. it floats deeper in the water and the effect of wind relative to current decreases. with a mean crossing time of 207 days. in any case. Hypothesis 3 Coconuts were carried in canoes from mid-Pacific islands by Poly- nesians on one or many occasions before discovery by Europeans. and in “wind and current” experiments. the husk absorbs water. Both lengths of time far exceed the known viability period for floating coconuts and. even in those of eight month’s duration. Initial simulation experiments from Ducie and Reao. in a mean time of 225 days. even within 8 months (Ward & Brookfield. 1973. 1). In the “current only” experiment. The last two periods far exceed any known example of the time a coconut might retain viability when floating. as a coconut floats. 2009). of the 732 nuts started from Christmas Island. In the simulation experiments. 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. Polynesia (Fig. This hypothesis was ruled out by Baudouin and Lebrun (2009). voyagers throughout the Pacific Islands regularly carried green and mature coconuts on their journeys for drink and food.

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.600 km from west to east and over 1. Two-way voyages in large outrigger canoes were conducted annually for economic and socio-cultural reasons over the 2.Coconuts in the Americas journey might take between 66 and 128 days (Fitzpatrick and Calla- ghan. bypassing any island where Polynesian coconuts grew at that time. 2004). may have had the capacity to carry coconuts to Panama. This hypothesis is based on Baudouin and Lebrun’s (2009) analysis that showed the close genetic relationship between coconuts in the Philippines and Panama. There is no record to suggest that Philippine mariners had the necessary technologies and knowledge. 2006:154–5) and maps constructed of sticks and shells were used. 1970. Rainbird. have striking traditions of long- distance voyaging in the western Pacific Islands. Hezel.600 km north–south. and has never previously been analyzed. Long-distance two- way voyaging by Micronesian people and their sophisticated methods of navigation are well documented (Lewis. Micronesian communities. and some make landfall as far north as the northern Ecuador coast. The sea lanes between islands extending over more than 1. or elsewhere in the western Pacific. 2009). Hypothesis 4 Coconuts were carried in canoes directly from the Philippines. and maintained regular links between these islands for many centuries. 1983. If coconuts from the Philippines reached Panama before the 17th century and did not come from Polynesia. As coconuts cannot have drifted to the Americas within any reasonable period of viability. The distance and likely duration of voyages from eastern Micronesia (Kiribati) are similar to those from Samoa or Tonga. 2009: 218). however. Thomas. we need to consider whether voyagers from Microne- sia. Finney. It is clear that Micronesians had the seamanship. for example from Kiribati or . 1987. 1). 1979. They did so by settling groups of islands extending over 5. and such voyages would be more likely to reach the Panama coast. one must conclude that they were carried there by voyagers from the western Pacific Islands. 1972.600 km were named (D’Arcy. 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). navigational skills and canoes to make long voyages of exploration.400 km length of the Caroline archipelago (Fig. Gladwin. Sophisticated concepts of estimating distances sailed and direc- tions followed were taught. 1995: 148). Those from Hawai’i reach the coast of Nicaragua and Costa Rica (Fitzpatrick & Callaghan. Micronesians following such strategies could have made long easterly voyages. 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.

ethnobotanical and linguistic evidence for the viabil- ity of the other four hypotheses. Micronesia borders on Melanesia. it is essential to collect a truly represen- tative sample of the coconuts along that coast. until more intensive genetic sampling of coconuts is done in Micronesia. . they could readily take advantage of south-westerly winds to reach Panama (see Irwin. 1). (2011b) affirm. Hypothesis 5 Coconuts were not present until carried by Spanish galleons returning from the Philippines. Clement et al. As Storey et al. In order to determine if coconut was on the Pacific coast of Panama at the time of European conquest. which they regarded as typical of an extremely small founder population. considering that the historical record suggests that not all the coconuts reported may have been similar to the San Ramon variety. and clear archaeological evidence is found of Microne- sian contacts with America. Christmas Island (Fig. while different selection pressures accounted for observed differ- ences. 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. there is currently no such evidence. regardless of the number of coconuts carried for consumption from Cebu to Navidad and then from Navidad to Pan- ama. Once east of about 125° West longitude. and by the doubts raised about the historical record. (2011) is only distantly related to the Pacific coast Panama Tall. led by Daniel Zizumbo-Villarreal. Finney (1985) also points out that eastward journeys would be easier in El Niño years. A larger sample of the relevant Philippine. However. Although the Micronesian sample used by Baudouin and Le- brun (2009) and Gunn et al.R. However. as well as being closer to the Philippines. (2011). The genetic evidence suggests a relationship that is amenable to future study. 1992: 9–16). the history and capacities for navigation of Micronesians suggest they may be candidates for any pre-Spanish carriage. However. and will be analyzed with the same microsatellite markers used by Baudouin and Lebrun (2009) and Gunn et al. which has more closely related coconuts in New Britain and northern Papua New Guinea (Fig. This hypothesis is supported by the lack of archaeological. we cannot claim that Micronesians were the trans-Pacific carriers. 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. even though the modern Pacific coast Panama Tall is indeed so. if Polynesia is ruled out as a source by the wide genetic gap between the sampled Polynesian and Panamanian coconuts. This type of collection has already started. 1). 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. Melanesian and Micronesian varieties is also needed to allow a more precise genetic analysis with the Pacific coast Panama Tall and Mexican Tall varieties. C.

and history records Spanish voyages with coconuts. Acknowledgements Our special thanks to Luc Baudouin. 1940. for information about the samples chosen for coconut genetic analysis and for stimulating our reconsideration of pre- Columbian coconuts. Brown. México. and to Pamela Brown. M. Buenas Aires. The sweet potato in Oceania: a reappraisal. however. about the Philippines. Beccari. Kenneth M.). Hence. Ethnology Monographs 19. Oxford University Press.asp Baudouin. Editorial Bajel.. formerly Director. 1917. P. Imprensa Manuel León Sánchez. P. Current Anthropology 35: 95–117. B. Philippines Journal of Science. 2005. London. to Madhavan Nayar. Historia natural y moral de las Indias. Genetic Resources and Crop Evolution 56: 257–262. ethnobotanical or linguistic evidence that supports a pre-Columbian origin of these Pacific coast Panama Talls. 1944. France. Fondo Cultural Económica. Coconut (Cocos nucifera L. Harwood (eds. R. O. Bourke & T. 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. Philippine Agriculturalist 20: 435–446. C. The origin and dispersal of Cocos nucifera. Broadway House. Central Plantation Crops Research Institute. to Michael G. and new work in the Philippines is suggested to confirm precise origins. for careful review of the language. for providing bibliographic material. Anglería. the most parsimonious explanation is that the Panama coconuts were introduced after Spanish conquest. Bardy. C. 56. Uichanco. Price. Online: http://www. Unless new archaeological remains are found to prove otherwise. I. as previously shown by morphometric analysis. M. Oceania Monograph. London. 1994. 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. C. Oviedo. Olsen. 1965. 1931). no archaeological. Lexical acculturation in Native American languages. A reanalysis of the historical record strongly suggests that early explorers made honest mistakes in identification. India. F. the genetic analysis and the presentation. ——— 1999. Travels in Asia and Africa 1325–1354 (Translated and edited by H. Coconut Research Institute. 1668. I. Gibb). Principes 9: 62–66. L. & P. [Revised by Edmundo O’Gorman from the 1st Edition published in Sevilla. Botany 12: 27–43. for reading the manuscript in draft and making numerous useful suggestions. New collections along the Mexican to Colombian Pacific coasts are improving the sampling for genetic analy- sis. Pittsburgh. A. Missouri. P. Ballard. CIRAD.) DNA studies support the hypothesis of an ancient Austronesian migration from Southeast Asia to America. H. Pensacola. J. St. Michigan. 1914. Literature Cited Acosta. 1929. Montpellier. W. Lebrun. . Florida. BT 31/22185/135025 UK Public Record Office. New York. Battuta. There is. Lalith Perera. this hypothesis can direct new research on the origins of American Pacific coast coconuts. Michigan Center. Prospectus for Gorgona Island Coconut Estate Ltd. Sri Lanka. Washington University. R.edu/halsall/source/1354-ibnbattuta. on “Cocos”. On the palms which are called Cocos and their great usefulness (Translated by L. Kasaragod. Lexical acculturation in Native American languages. Décadas del Nuevo Mundo. Sydney. 2009. This is supported by the DNA analysis. H.Coconuts in the Americas 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.fordham. Brown. Spain] Allen. Alzina. Louis.

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