The Botanical Review, New York, 79(x): online first.

DOI 10.1007/s12229-013-9121-z

Coconuts in the Americas

CHARLES R. CLEMENT1, DANIEL ZIZUMBO-VILLARREAL

2, CECIL H. BROWN

3, R. GERARD

WARD4, ALESSANDRO ALVES-PEREIRA

5 AND HUGH C. HARRIES

6

1 Instituto Nacional de Pesquisas da Amazônia; Av. André Araújo, 2936; 69067-375 Manaus,

AM, Brazil; cclement@inpa.gov.br; tel. (55-92) 3643-1862; fax (55-92) 3643-1867 2 Unidad de Recursos Naturales; Centro de Investigación Científica de Yucatán A.C.; Mérida,

Yucatán, México 3 Distinguished Research Professor Emeritus, Northern Illinois University; DeKalb, IL,

60115, USA; and Faculty Associate, University of West Florida; Pensacola, FL, 32514, USA

4 Emeritus Professor, Research School of Asia and the Pacific; Australian National

University; Canberra, Australia

5 Genetics and Plant Breeding Post-graduate Program; Universidade de São Paulo, Escola

Superior de Agricultura Luiz de Queiroz; Piracicaba, SP, Brazil

6 Coconut Time Line; Weymouth, Dorset, DT3 5NP Dorset, UK

I. 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.

Key words: Cocos nucifera, molecular genetics, history, archaeology, linguistics,

ethnobotany

Resumen

Cocoteros en las Américas

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. 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.

Palabras claves: Cocos nucifera, genética molecular, historia, arqueología, lingüística,

etnobotánica

II. 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, linguistics and

archaeology. During the early 20th

century, the famous Russian geneticist, 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 Yen‟s “ellipse” area of eastern Polynesia by 900-1000 years before present (BP) (Yen,

1974; Green, 2005: 46-7; Lebot, 2009: 94) and thence to New Zealand and Hawaii.

Linguistic and other evidence is cited by Scaglion (2005: 35-41) 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

Yen's (1974) hypothesis, making this the first plant evidence for a Neotropical to Oceania

dispersal.

A combination of archaeological, genetic and some linguistic evidence was recently

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., 2008ab). Although Acosta (1940) mentions them in the Caribbean and

Mexico, Patiño (1970: 34, 35) considers that no reliable historical record exists before

Columbus took chickens on his second expedition and disembarked them in 1493 as presents

for local chiefs, which thus explains Acosta‟s record, but is post-conquest in that region.

Storey et al. (2007) affirm that Francisco Pizarro found chickens in Peru, but Patiño‟s (1970)

documental analysis does not agree.

Skeletal, artifact and non-material cultural evidence also suggests Polynesian contacts

with the Chilean coast (Matisoo-Smith & Ramírez, 2010; Ramírez-Aliaga, 2011). Sewn plank

canoe-building technology, characteristic of Polynesia and Micronesia, found in small areas

in southern California and southern Chile, also suggests contact with the Pacific Islands

(Jones, 2011; Ramírez-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 considerable 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 introduction 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

Panama, e.g., 2250 BP, as postulated by Baudouin & 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 evidence, as well as with evidence from

ethnobotany and history, which might help with the “how” question.

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. 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.

III. 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.

Fig. 1. Map of the Pacific basin with features relevant to coconut in the Americas. Inserts –

locations 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.

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, 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).

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 & Lebrun (2009).

Baudouin & 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 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

selection 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).

Fig. 3. Group assignment at K=5 for the Philippines, Mexican and Panamanian coconut

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 that country. There are two mixed groups: orange represents mixtures of the Indo-

Atlantic and Pacific origins; dark green represents mixtures of the four Pacific groups

(Dwarf, South Pacific, Papua New Guinea, Panama). Redrawn from Gunn et al. (2011).

IV. 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 Fernández de Oviedo y Valdés, who was appointed as one of the official historians

of the Indies (Spanish American colonies) in 1523, after his return from Santo Domingo

(today‟s Dominican Republic). 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

1851-1855, 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 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; Patiño, 1963, 2002).

However, the first complete 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

Patiño (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 Oviedo‟s Summary and General

History to determine if coconut was present in Panama at the time of European conquest.

Patiño (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 Patiño and Zizumbo-Villarreal & Quero cautiously accept that Oviedo‟s reports are

correct. Based on Patiño‟s analysis, this is the time line for the early Spanish references to

coconut in Panama.

1492 – Discovery of New World

1494 – Treaty of Tordesillas divided the world outside Europe between Portugal

(eastwards) and Spain (westwards), which effectively prevented Spanish mariners from

sailing to Asia (until 1580).

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. The expedition explored only a small area and returned without mentioning

coconut, although numerous documents of this expedition have been lost.

1514-1515 – Alonso de la Puente (royal treasurer in Darien, on Panama‟s Caribbean

coast – saw coconut) and Rodrigo Colmenares (ship pilot and interpreter – heard of coconut)

visited the Pacific coast of Panama and were interviewed by Pedro Mártir de Anglería,

another official crown historian, in late 1516 in Spain. Anglería‟s report was only published

in 1944, but Oviedo knew Anglería 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 Calcutta – 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. 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.

Anglería states that coconuts were observed in three places along the Pacific coast:

Chimán, to the southeast of the current City of Panama; Natá, 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). 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. Natá 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.

Anglería also reports that “Some think that sea currents bring the seeds of these trees from

unknown regions.”

1519 – Gaspar 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 Anglería obtained this information from

Espinosa. Patiño suggests that Espinosa may only have realized the importance of his

observation after learning about coconut in Asia.

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. Near the Cape of Corrientes (Fig. 1), they reported “a large quantity of

coconuts” at La Candelaria Bay, which are reported in Oviedo‟s General History, but not in

the Summary.

1526 – Juan de Cabezas (pilot) discovered Cocos Island (Fig. 1) and provided

information used by Oviedo in the full General History.

1539 – Alvaro de Guijo (resident in the City of Panama) sent nuts to Hernán Cortés 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, Hernán Cortés left

for Spain in 1541 and never returned to Mexico.

1549 – Coconuts from the Cape Verde islands (Portuguese) planted in Puerto Rico

(Spanish), attributed to Diego Lorenzo, canon of Cape Verde (Bruman, 1944).

1553 – Coconuts from the Cape Verde islands planted at Bahia, Brazil (Bruman,

1944), possibly the first coconuts on continental America.

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

independently by Alonso de Arellano and Andrés 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.

1569 – Alvaro de Mendaña introduced coconuts from the Solomon Islands to Colima,

Mexico (Sevilla de Rio, 1974).

1580 – Treaty of Tordesillas became invalid when Portugal was ruled by Philip II of

Spain.

V. 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 (7000-5000 BP), horticulture started to

become important, expanding rapidly through the Early Ceramic Period (5000-2500 BP) until

most subsistence was horticultural by the Middle Ceramic Period (2500-1500 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 Choclé (central Panama) and Gran

Chiriquí (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) Cortés 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 (4.5-8.5 cm long by 3-4.5 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 butyracaea are reported at numerous

locations in the Gran Choclé and Gran Chiriquí regions of western Panama from 7000 BP to

European conquest (Smith, 1980; Cooke and Renare, 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 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, 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 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 Chimán or La Candelária 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.

VI. 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 (Patiño, 2002), although the native Panamanians

apparently were familiar with its propagation (see Time Line 1514-1515 above). Patiño

(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?” Patiño 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.

VII. The Linguistic Evidence

Paleobiolinguistics uses the comparative method of historical linguistics to

reconstruct 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, Ngäbare, Buglere, and Kuna. Other Chibchan

languages are spoken in Honduras, Nicaragua, Costa Rica, and Colombia. The smaller Choco

language family includes Emberá 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) published a

time-of-contact language-distribution map for the Caribbean region that includes 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.

Fig. 4. Map of the distribution of modern languages of Panama from Ethnologue (Lewis,

2009).

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 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 be retrieved for

Proto-Emberá, 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 I). All but two terms

in the 12 languages are unambiguous loans from European languages. The two exceptions are

Boruca siahuá and Kuna ógoba, 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).

Table I. 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

Words for coconut in the 12 languages robustly suggest that the plant was introduced

by Europeans into southern Central America. Speakers of native languages 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: 92-104).

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

Bachajón dialect of Tzeltal, a Mayan language of southern Mexico, the introduced sheep is

tumin čix, 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 and Brown, 1983).

The only word for coconut of the 12 languages that clearly is not a European loan is

Boruca siahuá. Plausibly, this term originally denoted a native palm similar to the introduced

coconut. Evidence for this comes from Cabécar, which, like Boruca, is a Chibchan language

of Costa Rica. Cabécar contains a word similar to the Boruca term, i.e., será, designating

Acrocomia aculeata, a palm whose fruit is fed to cattle and occasionally consumed by

humans in the Cabécar region. The phonological similarity of Boruca and Cabécar 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 Margery's (1989) exceptionally

thorough dictionary of Cabécar, and no term for A. aculeata is found in Quesada Pacheco and

Rojas Chaves's (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 *há: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.

VIII. 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 (Patiño, 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 Candolle‟s and

Vavilov‟s methodology, and suggests either that coconut arrived immediately before

European Conquest, rather than 2,250 years BP as suggested by Baudouin & Lebrun (2009),

or it arrived after European Conquest. Either way, both the genetic evidence and historical

records need further study.

IX. 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 & 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 II).

Table II. Coconut varieties included in the Generation Challenge Program data set that were

analyzed with microsatellite markers for the Baudouin & 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 1569 – fruit 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 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 (Table S1); includes

5 from Oxtapacab, Yucatan, Mexico

Panama Tall Agua Dulce 13 West of Nata, inland, taken to Jamaica, then Miami –

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

The varieties in the GCP data set that interest us most are those from the Philippines,

Mexico and Panama (Table II). Dwarf varieties from the Philippines are not included in the

table, as these are unlikely to have contributed to this Panama Tall‟s 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 & 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 II). 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 1890-1900 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 Michoacán 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 Mendaña from

the Solomon Islands in 1569 (Zizumbo-Villarreal & Colunga-GarcíaMarín, 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 Michoacán was introduced by President Lázaro

Cárdenas, apparently from the Philippines, and was initially established in plantations in

1937-1938. This is the most representative variety between Acapulco and Lázaro Cárdenas

Port, Guerrero state (Zizumbo-Villarreal & Colunga-GarcíaMarín, 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 & Lebrun (2009) and

Gunn et al. (2011) did not include plants from the Gran Darien nor the Gran Chocó (present

day Pacific coast of Colombia), where historic sources place the pre-conquest presence of

coconut at Chimán and Cape Corrientes, respectively. Thus, the sample is biased towards the

middle-western portion, covering the Gran Choclé and Gran Chiriquí 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 II) 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-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 II), 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 1000 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 II).

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 II). 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 & 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, 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 represented 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 & Quero, 1998; Zizumbo-Villarreal et

al., 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).

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.

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 Andrés

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 litres) (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 Ramon…”

(Copeland, 1914). However, similar coconuts were reported from Colombia 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, Arellano's account of cooking oil solidifying (literally “freezing”) 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 type – more 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, p. 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-GarcíaMarín, 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 & 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.

X. Botanical and Historical Questions

Given the doubts about the historical record, we revisit Oviedo‟s account because,

although it was previously analyzed by Patiño (1963, 2002), Allen (1965) and Zizumbo-

Villarreal & Quero (1998), the DNA data match between the Panama and the Philippines

coconuts raises fresh concerns. Oviedo's 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. Patiño (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 Patiño 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

Oviedo‟s 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 man's head; …” A century before Columbus

crossed the Atlantic, the Italian Ibn Battuta (1929) visited East Africa and India, and reported

that coconut is the size of a man‟s head, so this kind of information was available to

historians in Iberia at the time that Oviedo wrote, as was Varthema‟s (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.

“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 Patiño (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 Oviedo's choice of “ate” rather than “drank,” as it suggests that

they were fruit with little water. Oviedo's use of the word aborrecí (loathe) is a surprising

reaction – more people would agree with Charles Darwin (1860, p.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 evidence

(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 Oviedo's 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

Hernán Cortés (see Timeline above): If coconuts were interesting enough to send from

Panama to Mexico, why weren‟t they also sent to Madrid – at 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.

XI. 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.

XII. Hypotheses about Coconuts in Pre-conquest Panama

Five hypotheses have been presented to account for the historical observations, three

of which (2-4) 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, washing 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 carriage 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, 1951; 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 temperatures of floating coconuts delay germination

compared with nuts at average ground level temperatures (Ward & Allen, 1980).

Computer simulations of Pacific wind and surface current directions and speeds,

islands and coasts allowed tests of the possibility of coconuts drifting across the Pacific from

several possible starting places (Levison et al., 1973; Ward & Brookfield, 1992). It is

important to note that, as a coconut floats, the husk absorbs water, the nut‟s 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 extreme 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: 473-

4). The Equatorial Counter Current offers the most likely possibility for coconuts drifting

from west to east and therefore starting points used for the remaining simulated eastward

drifts from Micronesia 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

month‟s 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 reasonable period of viability.

Hypothesis 3 – Coconuts were carried in canoes from mid-Pacific islands by

Polynesians on one or many occasions before discovery by Europeans. This hypothesis was

ruled out by Baudouin & Lebrun (2009), based on the lack of similarity between the

microsatellite profiles of different Polynesian coconut varieties and the Panama Tall variety.

However, 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: 163-4; 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 and Callaghan, 2009: 218).

From Tonga or Samoa the journey might take between 66 and 128 days (Fitzpatrick and

Callaghan, 2009: 218). Simulated voyages from eastern Polynesia approaching 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

Hawai‟i reach the coast of Nicaragua and Costa Rica (Fitzpatrick and 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 & Lebrun‟s (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 Micronesia, 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 (D‟Arcy, 2006:154-5) and

maps constructed of sticks and shells were used. Sophisticated concepts of estimating

distances sailed and directions 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 north-south, 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 “implies 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 Christmas Island (Fig. 1).

Finney (1985) also points out that eastward journeys would be easier in El Niño years. Once

east of about 125° West longitude, they could readily take advantage of south-westerly winds

to reach Panama (see Irwin, 1992: 9-16). 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 & Lebrun (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 Micronesian 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 viability of the other four hypotheses, and by

the doubts raised about the historical record. Baudouin & Lebrun (2009) appear to have

assumed that the great number of coconuts carried between the Philippines 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 differences. However, regardless of the

number of coconuts carried for consumption from Cebu to Navidad and then from Navidad to

Panama, 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 representative 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 & 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.

XIII. 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 analysis,

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.

XIV. 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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