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; [email protected]; 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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