ArticlePDF Available

The Avocado (Persea Americana, Lauraceae) Crop in Mesoamerica: 10,000 Years of History

Authors:

Abstract

The most ancient evidence of the existence of the avocado tree (Persea americana, Lauraceae) in Mesoamerica is about 10,000 years ago in Coaxcatlan, Puebla (Mexico). Since then, the history of the avocado has been preserved and recorded in Mesoamerica, the domestication center for the species, and in northern South America, where pre-Hispanic Mesoamerican cultures dispersed it. In colonial times, the Spanish valued the avocado fruit highly and documented it in numerous texts. Since then, the avocado tree has been introduced into places where the climate is appropriate for its cultivation. Avocado is now the fourth most important tropical fruit in the world, and Mexico is the main producer, with more than one million metric tons produced in 2005. Resumen. La evidencia más antigua de la presencia del aguacate (Persea americana, Lauraceae) en Mesoamérica es de hace 10,000 años, en Coaxcatlán, Puebla (México). Desde entonces, la historia del aguacate ha sido documentada en Mesoamérica, su centro de domesticación, y en el norte de Sudamérica, en donde fue dispersado por las culturas que habitaron en Mesoamérica en la época pre-Hispánica. En el tiempo de la Colonia, los españoles apreciaron esta fruta y la documentaron en numerosos textos. Desde entonces el aguacate ha sido introducido en los lugares donde el clima es apropiado para su desarrollo. Actualmente el aguacate es la cuarta fruta tropical más importante en el mundo y México es el principal productor con más de un millón de toneladas métricas producidas en el año 2005.
The relationship between humans and trees
in Mesoamerica is very ancient, with trees hav-
ing played a very important role in the history
of many cultures that have inhabited this area
(e.g., Mayas, Toltecs, and Aztecs; Sánchez-
Colín et al., 2001; Telliz, 2000). One example
is the Chontales, who lived near the forests of
Tabasco (Mexico). They believed that the gods
lived in the forests and that they must therefore
maintain and conserve them (Vázquez-Dávila,
2001).
Among the trees used by ancient Meso-
american cultures, the avocado (Persea ameri-
cana Mill., Lauraceae) has had an important
place not only in the diet but in the mythology
and culture of different groups that have lived in
the area. It is possible that, even before some
human groups transitioned from hunter-gatherer
life styles to agriculture, that they valued the
avocado fruit as nutritious, and managed and
cultivated the crop in its natural stands for a
long time (Gama-Campillo and Gómez-Pompa,
1992). Seed remains found in ancient human
settlements in the Tehuacan Valley suggest that
the avocado could have been used as early as
8000 to 7000 BC and possibly domesticated at
least since 5000 BC by Mesoamerican groups
(Smith, 1966, 1969). However,the relationship
between humans and the avocado has been
complex, so, in order to understand its actual
status it is necessary to document its history.
In this paper, the social and cultural impor-
tance of the avocado crop since prehistoric times
is presented along with a paleographical recon-
struction of avocado history in Mesoamerica.
Moreover, to document the relationship of the
THE AVOCADO (PERSEA AMERICANA, LAURACEAE) CROP
IN MESOAMERICA: 10,000 YEARS OF HISTORY
MARÍA ELENA GALINDO-TOVAR,1,5 AMAURY M.ARZATE-FERNÁNDEZ,2
NISAO OGATA-AGUILAR,3AND IVONNE LANDERO-TORRES4
Abstract. The most ancient evidence of the existence of the avocado tree (Persea americana, Lauraceae) in
Mesoamerica is about 10,000 years ago in Coaxcatlan, Puebla (Mexico). Since then, the history of the avocado
has been preserved and recorded in Mesoamerica, the domestication center for the species, and in northern South
America, where pre-Hispanic Mesoamerican cultures dispersed it. In colonial times, the Spanish valued the avo-
cado fruit highly and documented it in numerous texts. Since then, the avocado tree has been introduced into
places where the climate is appropriate for its cultivation. Avocado is now the fourth most important tropical
fruit in the world, and Mexico is the main producer, with more than one million metric tons produced in 2005.
Resumen. La evidencia más antigua de la presencia del aguacate (Persea americana, Lauraceae) en
Mesoamérica es de hace 10,000 años, en Coaxcatlán, Puebla (México). Desde entonces, la historia del aguacate
ha sido documentada en Mesoamérica, su centro de domesticación, y en el norte de Sudamérica, en donde fue
dispersado por las culturas que habitaron en Mesoamérica en la época pre-Hispánica. En el tiempo de la Colonia,
los españoles apreciaron esta fruta y la documentaron en numerosos textos. Desde entonces el aguacate ha sido
introducido en los lugares donde el clima es apropiado para su desarrollo. Actualmente el aguacate es la cuarta
fruta tropical más importante en el mundo y México es el principal productor con más de un millón de toneladas
métricas producidas en el año 2005.
Keywords: Persea americana,Lauraceae, avocado, Mesoamerican cultures, Pre-Hispanic, colonial.
We thank the Universidad Veracruzana, Universidad Autónoma del Estado de México, and PROMEP for their support in
this investigation. We also thank the reviewers for their comments and suggestions.
1Facultad de Ciencias Biológicas y Agropecuarias-Córdoba, Universidad Veracruzana, Camino Peñuela-Amatlán,
Amatlán de los Reyes-Peñuela, Veracruz, México. E-mail: megalindo@uv.mx; Phone: 01(271) 716 6410.
2Facultad de Ciencias Agrícolas. Universidad Autónoma del Estado de México. Carretera Toluca-Ixtlahuaca Km 15,
Toluca, Estado de México.
3Centro de Investigaciones Tropicales, Universidad Veracruzana, Calle 7 de enero No. 12, Xalapa, Veracruz, México.
4Facultad de Ciencias Biológicas y Agropecuarias-Córdoba, Universidad Veracruzana, Camino Peñuela-Amatlán,
Amatlán de los Reyes-Peñuela, Veracruz, México.
5Author for correspondence.
HarvardPapers in Botany,Vol. 12, No. 2, 2007, pp. 325334.
©President and Fellows of Harvard College, 2007.
avocado with the customs and habits of
Mesoamerican cultures from its beginnings
10,000 years ago up to the present, and follow-
ing the methodology of Messer (2003), we
examined ethnohistoric and linguistic data as
well as archaeobotanical remains (Smith, 2001)
that provide evidence for the antiquity of avo-
cado tree cultivation.
326 HARVARD PAPERS IN BOTANY Vol. 12, No. 2
HISTORICAL-GEOLOGICAL ORIGIN
The origin of the avocado, as with any other
species, cannot be explained without its histori-
cal-geological background (Graham, 1995).
Ancestors of the Lauraceae family originated in
Gondwana (Africa) and migrated to Laurasia
(Europe; Raven and Axelrod, 1974). According
to Chanderbali et al. (2001), the Lauraceae orig-
inated in Laurasia, from the Gondwanan ances-
tors. Subsequently, one part of the family
migrated to Asia, and another, including the
Perseae clade, migrated to North America
(Renner, 2004). Later, when Central America
was formed (Miocene-Pliocene) and mountain
building occurred, new habitats emerged and
speciation took place due to geographical isola-
tion (Scora and Bergh, 1992). Archaeological
evidence shows that when the climatic condi-
tions changed during the Paleocene glaciations,
avocado ancestors migrated from North
America to the south and became established in
the more hospitable habitats of Mesoamerica
(Schroeder, 1968; Storey et al., 1986; Scora and
Bergh, 1992; Bergh, 1995). Evidence suggests
that the complex geological history of Mexico
has been the main evolutive factor for the avo-
cado (Ramamoorthy et al., 1993).
THE PRE-HISPANIC PERIOD
The avocado has been consumed in Meso-
america by human groups since prehistoric
times (Mac Neish, 1964). Buckler et al. (1998)
documented that from 16,000 to 8,000 BC the
weather in this region was appropriate for avo-
cado development. It is therefore possible that
avocado cultivation began at this early time and
continued to be used by different cultures that
inhabited the Americas (Galindo et al., in press).
Examples of these cultures and their use of the
avocado are explained in more detail below.
Caral, Peru
In Peru, the Supe Valley was inhabited by the
Caral civilization, the oldest known culture in
the Americas (Solis et al., 2001). Ethnobotanical
remains found here suggest a system of agricul-
ture that depended on irrigation (Solis et al.,
2001). Furthermore, recent findings indicate that
the Supe Valley comprises a cultural complex
going back as far as 3100 BC; and domesti-
cated avocado botanical remains indicate that
avocado was grown there at least since 1200 BC
(Skidmore, 2005). Moreover, because there is no
evidence of maize or any other grain (Skidmore,
2005), it is possible that avocado was an impor-
tant staple for these people before other crops
were grown there. In addition to the Caral civ-
ilization, there is also archaeological evidence
that the avocado was a part of the diet of groups
inhabiting the Moche Valley in northern Peru in
2500–1800 BC (Pozorski, 1979), and on the
Peruvian coast in 1500 BC (Heiser, 1979).
Mokaya, Mesoamerica
In Mesoamerica, one of the first known cul-
tures is the Mokaya (1800 BC). The Mokaya
inhabited the Soconusco area during the
Formative period (1500 BC–300 AD), and
although the Mokaya name means maize, there
is evidence that this was not their primary food
source (Taube, 2004). As the Mokayas were a
sedentary group engaged in food acquisition,
with an incipient agriculture, it is possible that
they used common fruits in the area, especially
those as nutritive as the avocado, as a source of
food. In addition, as the Mokayas were the
forerunners of the Olmec and Maya cultures
(Taube, 2004), they may have passed on to
them their knowledge of the use and cultivation
of trees, including the avocado.
Maya, Mesoamerica
Like the Chontales, the Mayas also had a spe-
cial appreciation for trees. In the Popol Vuh, a
Mayan holy book written in Mayan hiero-
glyphic script around 1550, there is a part that
refers to the Creation Myth. In this part of the
book there is an interesting reference to a place
with many trees (“the delightful place”):
“There, the creators found the yellow and the
white maize as the appropriate food for men and
from them they made the flesh when man was
formed. In this beautiful place full of delights
there were many trees of pataxte (Theobroma
bicolor Bonpl.) and cacao (Theobroma cacao
L.), and innumerable zapotes (Pouteria sapota
(Jacq.) H.E. Moore & Stearn), anonas (Annona
muricata L.), jocotes (Spondias purpurea L.),
nantzes (Byrsonima crassifolia (L.) Kunth.),
and matasanos (Casimiroa edulis La Llave &
Lex.)” (Anonymous, 2002).
There are paleoethnobotanical accounts sug-
gesting the Maya used the coyol tree
(Acrocomia aculeata (Jacq.) Lodd. ex. C. Mart.)
as a source of food, oil, and/or wine (Lentz,
1991). It is also known that other fruit trees, such
as ciruela (Spondias sp.), nance (Brysonimia
crassifolia (L.) Kunth),avocado (Persea amer-
icana), and possibly zapote (Pouteria sp.),
were common in the Copan Valley (Guatemala)
from 900 BC to 900 AD, indicating a reliance
on arboriculture as part of the subsistence strat-
egy for the Mayas (Lentz, 1991). With regard
to the avocado, Colunga-García and Zizumbo-
Villareal (2004) have suggested that domesti-
cated avocados from other cultural areas were
introduced to the Maya lowlands by at least
3400 BC. The Mayas, however, had used avo-
cado since ancient times (Fedick, 1995), and
they were among several of the Mesoamerican
cultures that domesticated this fruit (Gama-
Campillo and Gómez-Pompa, 1992).
Iconographic Data
Since many documents that could have shed
light on pre-Hispanic avocado history were
destroyed by the Spanish in their effort to
Christianize the cultures that inhabited
Mesoamerica (Ossenbach, 2005), the icono-
graphic data reveal important information
about the relationship between the avocado and
some Mesoamerican cultures. One example is
the Maya civil calendar (Haab calendar) devel-
oped between 800 and 300 BC. In this calendar
the name of each month is based on seasonal
and agricultural events, and the 14th month is
represented by a glyph representing the avo-
cado. The Mayas called this glyph Uniw or
Uniiw in their classic language and K’ank’in in
the yucatec of the 16th century (Landa, [1560]
1978; Kettunen and Helmke, 2005). Another
example is in the Pacal tomb inscriptions in
Palenque, Chiapas, Mexico. This tomb was
built in 650 AD, and on the sides of the sar-
cophagus are 10 figures representing Pacal’s
ancestors, 9 of them men and 1 a woman
(Schele, 1974). Every figure emerges from the
earth, and behind each of them there is a tree
with fruits that include the cacao, avocado,
soursop (Annona muricata L.), and chicozapote
(Manilkara zapota (L.) P. Royen) (Schele,
1974). These figures represent a forest growing
around the king’s sarcophagus and also repre-
sent the main fruit trees the Mayas cultivated
around their homes, as it was thought that their
ancestors were reborn as the trees (Schele,
1974, 1998).
Other interesting iconographic data that sup-
port the importance of the avocado in
Mesoamerica is found in “El Codice Mendoza,”
amanuscript painted by an Aztec tlacuilo
(“artist”) at the time Don Antonio de Mendoza
was viceroy of New Spain (“Nueva España”;
Mendoza, 1989). In this manuscript there is a
glyph representing Ahuacatlan, whose name
means a place where avocado is abundant. This
town was identified by a tree with a tooth in the
trunk (ahuacacahuitl)and a calli, meaning
place or town (Telliz, 2000). Castillo (1978)
has placed this town in the state of Jalisco
(Mexico), because the other towns mentioned
in the same plate belong to this area. In addition
to the iconographic data, there is also an Aztec
legend that describes the way Yaotl was trans-
formed into an avocado grasshopper (ahua-
cachapulin)as punishment when he disobeyed
the gods (Robelo, 1951).
Linguistic Data
In addition to the iconographic references,
linguistic data show the ways different human
groups used plant and animal names to record
their experiences and secure the survival of
their history.However,according to Navarijo
(1995), the presence of a plant or animal in
nature alone is not enough for it to be named. It
is necessary for the society or human group to
acknowledge the importance of the species
before naming it (Navarijo,1995). Understanding
the naming and significance of avocado is the
first step in understanding the cultural percep-
tions that various groups had of avocado; and
two or more names for a single species repre-
sented a greater cultural significance (Bye,
1993). For avocado, the fact that many cultures
recognized it with a name demonstrates that it
was an important tree. For example: some
Maya groups, such as the Pokomchi, Pokoman,
Cakchiquel, Quiche, and Uspanteca, named the
avocado oj (Gama-Campillo, 1992; Schieber and
Zentmyer,1992); the Chucutamanes, Tzental,
Tzontzil, and Chnabal, on (Gama-Campillo,
1992; Schieber and Zentmyer, 1992; Landa,
[1560] 1978); the Kekehi, oh (Popenoe et al.,
2007 GALINDO-TOVAR ET AL., 10,000 YEARS OF AVOCADO HISTORY 327
1997); the Chicomulcelteca, ou (Popenoe et al.,
1997); the Choi, um (Popenoe et al., 1997); the
Chorti and Chontal, un (Popenoe et al., 1997);
the Otomi, nttzani (Popenoe et al., 1997); the
Zapotec, yaus, yashu, ishu,or isu (Popenoe et
al., 1997); the Mixe, cuchpa (Popenoe et al.,
1997); the Chibcha, cura (Popenoe et al.,
1997); the Tarascos, cupanda (Popenoe, 1963);
the Quekchi, o(Gama-Campillo, 1992); and in
totonaco, cucata (Cortés, [1522] 1970). But the
name in contemporary Spanish—aguacate
comes from the Aztec word ahuacaquahuitl,
which means “testicle.” In South America, the
Incas gave the name palta to the avocado,
because it was brought from a place named
Palta (Vega, [1605] 1995).
328 HARVARD PAPERS IN BOTANY Vol. 12, No. 2
EARLY DISPERSION AND COMMERCE
In Mesoamerica, many cultures shared reli-
gious beliefs, art, architecture, science, and
technology for thousands of years (Wolf, 1967).
In addition, since early times, a wide network
for commercial exchange existed, and the dif-
ferent cultures inhabiting the area shared many
traits (Ortíz and Rodríguez, 2000). Because of
this, it seems reasonable to think that the avo-
cado was an important exchange product. For
instance, by 1600 to 500 BC, the Olmec was a
complex society with a wide distribution in
Mesoamerica and an extensive network of com-
merce routes (Love, 2005). Based on evidence
that suggests that the Olmec had contact with
the Papayecas (Honduras) in 1200–1000 BC
(Healy, 1978), and because this time coincides
with the avocado remains found in this area
(Healy,1978), it is probable that the Olmecs
brought the avocado as a commerce product.
Wolters (1999) proposed that the avocado
and other crops were brought from Mexico to
Ecuador and to northern Peru in pre-Columbian
times by the Valdivia culture (western Ecuador)
and their successors. He proposed that in their
travels by boat from west Ecuador to Peru and
Middle America (since 2200 BC) and to south-
ern Mexico (since 1450 BC), the Valdivias dis-
persed several plants, among them the avocado.
In addition, in “Los Comentarios Reales de
los Incas,” Garcilazo de la Vega ([1605] 1995)
writes about the way the avocado was brought
to the Inca valleys. He describes the origin,
kings, religion, laws, and government of the
Incas and relates that when the king Tupac Inca
Yupanqui conquered Palta (now Ecuador), he
brought the avocado to Cuzco and to the Incas’
warm valleys (Vega, [1605] 1995). It is known
that Yupanqui conquered Cuzco in 1450–1475
A.C. It is therefore not surprising that by the
time the Spanish arrived in the Americas, the
avocado was consumed from Mesoamerica to
Peru (Webber, 1936; McPherson, 1955;
Popenoe, 1963; Takashi, 1968), an area settled
by agricultural people by that time.
COLONIAL TIMES
When the Spanish arrived in the Americas,
the New World flora and fauna attracted the
attention and curiosity of many chroniclers
who were interested in the description and
identification of new species, mainly with
medicinal rather than botanical purposes. These
chroniclers recorded many descriptions of the
cultivation and use of native plants, thus pro-
viding important ethnohistoric information on
the cultures inhabiting the Americas at that
time (Ossenbach, 2005).
The first written description of the avocado
was made by Martín Fernández de Enciso in
“La Suma de Geografía,published in Sevilla,
Spain, in 1519. He found the avocado in Yaharo
(now Colombia) and described it as “an orange,
and when it is ready for eating it turns yellow-
ish; that which it contains is like butter and is of
marvelous flavor, so good and pleasing to the
palate that it is a marvelous thing.In addition
to Fernández de Enciso, other chroniclers have
also written about the avocado: Fernández de
Oviedo, in “Sumario de la Natural Historia de
las Indias” ([1526] 1996), described it as being
like a pear but better; Cervantes de Salazar, in
“Crónica de la Nueva España” ([1594] 1985),
described it as being like a fig but bigger;
Landa, in “Relación de las Cosas de Yucatán”
([1560] 1978), described it as a big and fresh
tree with fruits like a courgette; Vega, in “Los
Comentarios Reales de los Incas” ([1605]
1995), referred to the avocado as a tasty and
healthy fruit for sick people, which, when eaten
with sugar,is like a preserve; Ximénez, in
“Libros de la Naturaleza y Virtudes de las
Plantas y Aanimales, de Uso Medicinal en la
Nueva España” ([1615] 2001), described the
ahuacaquahuitl tree as being like an oak
with orange leaves, small flowers, and fruits
like figs.
At the end of the 16th century, the king of
Spain and the Indies, Felipe II, asked Francisco
Hernández for a description of the plants, ani-
mals, and minerals from New Spain, with a
particular focus on the virtues and uses of them.
In his book, “Historia de las Plantas de la
Nueva España,” he described the ahoa-
caquahuitl as a tree like an oak, with hanging
black fruits like figs and with anise-scented
leaves, and he added that this tree grows every-
where in New Spain (Hernández, [1605] 1942).
Likewise, during the same period, Antonio de
Ciudad Real ([1590] 1993) wrote “Tratado
Curioso y Docto de las Grandezas de la Nueva
España.” In his books, he described the avo-
cado in Mexico as a fruit the size and color of
an early fig, with a big seed and little flesh but
tasty and healthy even though in other places
they were bigger; and he documented the pres-
ence of avocado in the town and monastery of
Cuauhcachulan (Tlaxcalla episcopate), in
Tehuacan (Puebla), in Iztapa and Comitan
(Chiapas) in Tuchpan (Jalisco), in Michoacan,
and in Yucatan. Also, in his trip to Guatemala
he described avocados in the town of Apenega
and in the margins of the Atitlan Lake (Ciudad
Real, [1590] 1993).
Moreover, there are at least three interesting
mentions of the avocado used as a tribute. Friar
Diego Durán ([1867] 1967) wrote in the mid-
16th century about the tributes and riches that
the people governed by the Aztecs gave to
them. Among them were the fruits, especially
the ones from the warm lands, like the avocado
(Durán, [1867] 1967). In “El Códice Mendoza”
(Mendoza, 1989) there is a glyph for the avo-
cado tree (ahuacacahuitl in the Nahua lan-
guage) representing the merchandise the town
of Ahuacatlan gave as a tribute to the Aztecs. In
“Suma de la Visita de los Pueblos,” an inven-
tory of goods obtained between 1531 and 1544
from 907 towns in Central Mexico, Aculma is
reported as a town that gave in tribute, among
other merchandise, 10 avocados (Paso y
Troncoso, 1905). In addition, it has been docu-
mented that the avocado was sold, among other
fruits, in the Tlatelolco tianguis (open-air mar-
ket) in Tenochtitlan (Cervantes de Salazar,
[1594] 1985) and consumed by the Aztec nobil-
ity (Sahagún, [1570] 2002).
In South America, the avocado has been
described by several chroniclers. Among them,
Pedro de Cieza de León, in “La Crónica del
Perú,” described his experiences from 1532 to
1550, and according to him there were avoca-
dos all over the valleys of the plains in the
provinces from Tamboblanco to San Miguel
city on the Peruvian coast and in the Cali
region of Colombia. He also mentioned having
seen many avocados in Puerto Viejo province
(Costa Rica) and in the equinoctial line (Cieza
de León, [1553] 1962). Friar Gaspar de
Carvajal ([1542] 1999), in his book “Relación
del Nuevo Descubrimiento del Famoso Río
Grande de las Amazonas,” referred to Omagua,
atown located in northeastern Peru in the
Amazon rain forests, as a place where avocados
and other fruits were abundant. Later Don
Jorge Juan and Don Antonio de Ulloa ([1748]
1970), in their book “Del Viaje a la América
Meridional,” described the avocado as a com-
mon fruit in the lands around Quito (Ecuador).
Avocado, besides being consumed by people,
was used to feed domestic animals. For exam-
ple, pigs would eat the ripe fruit falling from
the trees (Labat, [1694] 1964). In his book
“Nouveau Voyage Aux Isles de L’Amerique,”
Labat mentioned that the meat of these animals
had an excellent flavor when they were fed
with avocados. According to Benavente
([1536] 2003), dogs and cats would also eat all
the avocado types; after a good meal of hen
they ate avocados, as a satiated man after eat-
ing meat has an olive.
Of course we cannot ignore the medicinal
uses of avocado. When Garcilazo de la Vega
described the avocado, he wrote that the fruit is
delicious and very healthy for sick people
(Vega, [1605] 1995). Friar Francisco Ximénez
in 1615 described the avocado leaves as dry
and warm, and for this reason they were com-
monly used in lavatories (Ximénez, [1615]
2001). The avocado seed rubbed on a swollen
instep healed it and eliminated bruises, and
because of certain astringent characteristics,
avocado healed the blood chamber and pre-
vented the splitting of the hair tips; and if you
made any mark on the seed with a pin or any
other object, the stain on a cloth would never
disappear (Ximénez, [1615] 2001).
The Spanish liked the avocado so much that
they distributed it to their other colonies in the
Americas, in the eastern hemisphere, and to
Europe (Knight, 2002). In Europe, the first
introduced avocado was the West Indian type
(Williams, 1977). Avocado was introduced to
Indonesia by 1750, to the Philippines in 1890,
and to Brazil in 1809 (Knight, 2002).
2007 GALINDO-TOVAR ET AL., 10,000 YEARS OF AVOCADO HISTORY 329
330 HARVARD PAPERS IN BOTANY Vol. 12, No. 2
AVOCADO DIVERSITY
In Mesoamerica, differentiation of many taxa,
including Persea,has occurred (Ramamoorthy
et al., 1993). When the first human groups
arrived in Mesoamerica, it is possible that,
among other species, they started to consume
and select avocado ancestral varieties (Mac
Neish, 1964; Smith, 1969), thus initiating
selection and domestication of the avocado. As
aresult, there are at least three varieties of avo-
cado with differences related to the ecological
preferences of the tree and fruit characteristics
(Ashworth and Clegg, 2003): (1) Persea ameri-
cana var. drymifolia (Schlecht. et al. Cham.
Blake) (Mexican) is characterized by a relatively
good cold tolerance and small fruits covered by
athin, purplish-black skin; (2) P. americana
var. guatemalensis L. Wms. (Guatemalan) is
somewhat cold tolerant, and the fruit has a
thick, tough skin, which remains green until
maturity; (3) P. americana var. americana
(West Indian) is adapted to humid tropical con-
ditions, and the fruit has a smooth, easy-to-peel
skin and a flesh with an almost sweet taste not
found in the other two varieties (Williams,
1976; Bergh and Ellstrand, 1986). In addition,
Ben-Ya’acov et al. (2003) proposed P.ameri-
cana var. costarisencis (Costa Rican) as another
horticultural variety, but Van der Werff (2002)
considers that much of the avocado variation
can be attributed to the cultivation process and
prefers to ignore the cultivated varieties.
In addition, the diversity of the avocado has
been known since pre-Hispanic times.Benavente,
in his “Historia de los Indios de la Nueva
España” ([1536] 2003), made a distinction
among different avocado types: “the ones com-
mon in all this land and all the year, are like
early figs. Other avocados are as big as large
pears, and are so good like the best fruit in the
New Spain. There are others as big as a small
pumpkin; ones with a big seed and little flesh
and others with more flesh.” Sahagún ([1570]
2002) also described three different types of
avocado: the ahuacatl or ahuacacahuitl has
dark green leaves, and the fruit is black in the
outside and white and green in the inside; the
tlacazolahuacatl is like the former but bigger;
and the quilahuacatl is green in the outside and
very good to eat.
Other chroniclers not only described the dif-
ferent avocado types but situated them geo-
graphically.Friar José Acosta, in 1590, in his
“Historia Natural y Moral de las Indias” differ-
entiated between the Mexican avocado and the
one from Peru. He described the avocados from
Peru as big fruits with a hard shell that peels
easily, and the ones from Mexico as mostly
small with a thin shell that peels like an apple
(Acosta, [1590] 1985). These descriptions
coincide, respectively, with the West Indian and
Mexican avocados described by Bergh and
Ellstrand (1986).
It is also interesting the way Friar Bernabé de
Cobo, in his book “Historia del Nuevo Mundo”
([1653] 1956), described three different avo-
cado types: “The Palta in Yucatan is a tree of
very attractive appearance, of the size of a large
fig tree; its leaf is similar to that of the mulberry
and its fruit is one of the finest in the Indies; in
some regions it becomes as big as a small
squash or large citron. The Palta has a thin skin,
more tender and flexible than that of a Ceuta
lemon, green externally, and when the fruit is
quite ripe, peeling readily. It has the largest seed
that I have ever seen in any fruit, either in the
Indies or Europe; it is as large as a hen’s egg,
and spindle shaped; it is of a reddish white sub-
stance, tender like the meat of a chestnut, and
covered with a grayish parchment. It has the fla-
vor of bitter almonds, and when pressed it
yields an oil like that of the almond. Between
the seed and the outer skin is the meat, slightly
thicker than one’s finger except at the neck
where it is very thick. It is of whitish green
color, tender, buttery, and very soft. Some peo-
ple eat it with sugar or salt, others just as it
comes from the tree, it being of such good fla-
vor that it requires no seasoning. But, although
it is very pleasant to the taste, it should be eaten
in moderation because it is considered to be
heavy and indigestible. The best Paltas come
from hot, dry regions; in Peru they grow in the
Valley of Lea. A second kind of Paltas is a large,
round one which is produced in the province of
Guatemala, and which does not have as smooth
skin as the first. The third is a small Palta found
in Mexico, which in size, color, and form
resembles a breva fig; some are round and oth-
ers elongated, and the skin is as thin and smooth
as that of a plum. In some regions they cut the
immature Palta in small bits and put it in brine,
to take the place of olives. The tree wood is use-
ful in building, and for fuel.” These avocado
descriptions also resemble the ones made by
Bergh and Ellstrand in 1986: the palta in
Yucatan would be the West Indian avocado; the
second kind of palta the Guatemalan type; and
the third the Mexican type.
2007 GALINDO-TOVAR ET AL., 10,000 YEARS OF AVOCADO HISTORY 331
THE AVOCADO IN MODERN TIMES
Today the avocado is cultivated all over the
world. In 1856 avocados were brought to
California by Nicaraguan settlers. In Israel, the
first avocado was introduced in 1908. From
1933 to 1998 avocado selections were intro-
duced in Central America, the Caribbean,
North and South America, Africa, Asia,
Oceania, and Europe (Knight, 2002). In recent
years, the avocado has become the fourth most
important tropical fruit in the world, and it con-
tinues to increase in importance in many places
(Bergh, 1992). China, for example, produced
45,000 tons in 1996, whereas in 1991 there was
no avocado production reported (Knight,
2002). At present, Mexico is the main producer
(Foreign Agricultural Service, USDA, 2006)
with 1,021,515 tons in 2005 (SIAP, 2007).
However,in Mesoamerica avocado trees are
still cultivated in traditional orchards, backyard
gardens, and as living fences and they are con-
sumed and sold on a regional scale. Traditional
orchards have not been intensively managed
and they therefore preserve populations that
still resemble their wild relatives thus provid-
ing opportunities for a better understanding of
avocado domestication (Miller and Schaal,
2005).
Current varieties and rootstocks for avocado
cultivation in the world are the products of var-
ious breeding programs based on exploration,
collections, conservation, and evaluation trials
throughout their regions of origin and disper-
sion (Mijares and López, 1998). Indeed, many
modern commercial plantings are new varieties
and cultivars, obtained by hybridization of var-
ious materials collected in Mexico and Central
America (Lemus et al., 2005), or they have
arisen as chance seedlings (Bergh, 1992), prob-
ably due to the still existing genetic variability
of the avocado.
At present there is a great need for the intro-
duction of cultivars to improve quantity, qual-
ity, uniformity, and seasonal distribution; and
biotechnology techniques promise new tools
for expediting these breeding programs and
making avocado breeding much more efficient
(Bergh, 1992; Clegg et al., 1999). In fact,
research in this area has already resulted in cul-
tivars with improved disease tolerance and
resistance (Nakasone, 1976).
Since ancient times, the avocado has played
an important role in the diet of the native cul-
tures of Mesoamerica. Archaeological and
paleographical data provide an interesting look
at the history of the avocado and its introduc-
tion to northern South America before the
arrival of the Spanish. Archaeological data pro-
vide evidence of the origin and early relation-
ship of the avocado with pre-historic human
groups. The existence of local names for the
avocado used by native groups also serves as
evidence of the importance and use of the plant.
Reports by Spanish chroniclers provide a fasci-
nating look at the use of the avocado in the
Americas at the time they arrived. Moreover,
the ethnohistoric data have been important in
documenting the management, uses, diversity,
and geographic distribution of avocado in colo-
nial times. Comparison of ancient, colonial,
and modern references has provided useful data
for clarifying the origin, dispersion, and
domestication of the species.
LITERATURE CITED
ACOSTA, J. DE. [1590] 1985. Historia Natural y
Moral de las Indias.2nd ed. Fondo de Cultura
Económica. México.
ANONYMOUS. 2002. Popol Vuh: Las antiguas histo-
rias del Quiche.Epoca. México.
ASHWORTH,V. E. T. M., AND M. T. CLEGG. 2003.
Microsatellite markers in avocado (Persea ameri-
cana Mill.): Genealogical relationships among
cultivated avocado genotypes. J. Herd. 94:
407–415.
BENAVENTE,F. T. M. DE.[1536] 2003. Historia de los
Indios de la Nueva España.Special ed. Dastin,
S.L. España.
BEN-YAACOV,A., A. SOLIS-MOLINA,AND G. BUFLER.
2003. The mountain avocado of Costa Rica, a new
sub-species. In Proceedings V World Avocado
Congress 27–33, Consejería de Agricultura y
Pesca. Málaga, España, 19–24 October 2003.
BERGH,B. O. 1992. The origin, nature and genetic
improvement of avocado. Cal. Avoc. Soc. Yrbk.
76: 61–75.
––––––. 1995. Avocado: Persea americana
(Lauraceae). In J. SMARTT AND N. W. SIMMONDS,
EDS., Evolution of Crop Plants.Longman
Scientific & Technical, Essex, UK.
BERGH,B. O., AND N. ELLSTRAND. 1986. Taxonomy
of the avocado. Calf. Avoc. Soc. Yrbk. 70:
135–145.
BUCKLER IV, E. S., D. M. PEARSALL,AND T. P.
HOLTSFORD.1998. Climate, plant ecology, and
Central Mexican archaic subsistence. Curr.
Anthropol. 39:152–164.
BYE,R. 1993. The role of humans in the diversifica-
tion of plants in Mexico. Pages 707–731 in T. P.
RAMAMOORTHY,R. BYE,A. LOT,AND J. FA.,EDS.,
Biological Diversity of Mexico: Origins and
Distribution.Oxford University Press, New York.
CARVAJAL, G. DE.[1542] 1999. Relación del nuevo
descubrimiento del famoso Río Grande de las
Amazonas.Fondo de Cultura Económica, México.
CASTILLO,V. M. 1978. Matrícula de los tributos.
Comentarios y paloegrafía. Pages 523-588 in
IGNACIO BERNAL AND MIGUEL LEÓN PORTILLA,
EDS.,Historia de México.Salvat Editores de
México, México.
CERVANTES DE SALAZAR,F. [1594] 1985. Crónica de
la Nueva España.Editorial Porrúa, México.
CHANDERBALI,A. S., H. VAN DER WERFF,AND S. S.
RENNER.2001. Phylogeny and historical biogeog-
raphy of Lauraceae: Evidence from the chloroplast
and nuclear genomes. Ann. Missouri Bot. Gard.
88: 104–134.
CIEZA DE LEÓN,P. [1553] 1962. Cronica del Perú.
3rd ed. Espasa-Calpe, Madrid, España.
CIUDAD REAL, A. DE.[1590] 1993. Tratado Curioso
yDocto de las Grandezas de la Nueva España.3rd
ed. Universidad Nacional Autónoma de México,
México.
CLEGG,M. T., M. KOBAYASHI,AND J. Z. LIN. 1999.
The use of molecular markers in the management
of avocado (Persea americana Mill.). Revista
Chapingo. Serie Horticultura. 5: 227–231.
COBO,B. DE [1653] 1956. Historia del Nuevo
Mundo.Biblioteca de Autores Españoles, España.
COLUNGA-GARCÍA MARÍN, P., AND D. ZIZUMBO-
VILLAREAL.2004. Domestication of plants in
Maya lowlands. Econ. Bot. 58 (Supplement):
S101–S110.
CORTÉS, HERNÁN.[1522] 1970. Cartas de relación
de la conquista de México.5th ed. Espasa Calpe,
Madrid.
DURÁN,F. D. [1867] 1967. Historia de las Indias de
Nueva España e Islas de Tierra firme.Editorial
Porrúa, Mèxico.
FEDICK,S. L. 1995. Indigenous agriculture in the
Americas.. J. Arch. Res. 3: 257–303.
FERNÁNDEZ DE ENCISO,M. [1519] 1546. Suma de
Geografía que trata de todas las partidas y
provincias del mundo, en especial de las indias y
trata largam ete del arte del marear ju tamente.
Biblioteca virtual de Andalucía. http://www.jun-
tadeandalucia.es/cultura/bibliotecavirtualandalu-
cia/consulta/registro.cmd?id=100034 (accesed 3
august 2007).
FERNÁNDEZ DE OVIEDO,G. [1526] 1996. Sumario de
la Natural Historia de las Indias.2nd ed. Fondo
de Cultura Económica. México.
FOREIGN AGRICULTURAL SERVICE, OFFICIAL USDA
ESTIMATES.2006. Fresh avocados: Production,
supply and distribution in selected countries.
http://www.fas.usda.gov/psd/complete_tables/HT
P-table6-101.htm (accessed january 11 2007).
GALINDO,M. E., N. OGATA ,AND A. M. ARZATE.In
press. Some aspects of avocado (Persea ameri-
cana Mill.) diversity and domestication in
Mesoamerica. Genet. Resour. Crop. Ev.
GAMA-CAMPILLO,L. A. 1992. Studies on the system-
atics and ethnobotany of the subgenus Persea. Ph.
D. diss., Department of Botany and Plant Science,
University of California, Riverside.
GAMA-CAMPILLO,L. A., AND A. GÓMEZ-POMPA.
1992. An ethnoecological approach for the study
of Persea: A case study in the Maya area. Proc. of
Sec. World Avoc. Congr.: 11–17.
GRAHAM,A. 1995. Development of affinities
between Mexican/Central American and Nothern
South American lowland and lower montane
vegetation during the tertiary. Biodiversity and
Conservation of Neotropical Montane Forests.
Pages 11–22 in S. P. CHURCHILL,H. BALSLEV,AND
E. FERRERO,EDS., Biodiversity and conservation
of neotropical montane forests.New York Bot.
Gard. I.
HEALY,P. F. 1978. Excavations at Rio Claro,
Northeast Honduras: Preliminary report. J. Field
Archaeol. 5: 16–28.
HEISER,C. B. 1979. Origins of some cultivated new
world plants. Ann. Rev. Ecol. Syst. 10: 309–326.
HERNÁNDEZ,F. [1605] 1942. Historia de las plantas
de la Nueva España.Vol. 3. Imprenta
Universitaria, UNAM, México.
JUAN, J., AND A. DE ULLOA.[1748] 1970. Relación
histórica del viaje a la América Meridional, hecho
de orden de S. Mag. Para medir algunos grados
de meridiano terrestre, y venir por ellos en
conocimiento de la verdadera figura, y magnitud
de la Tierra, con otras observaciones astronómi-
cas, y físicas.Fundación Universitaria Española,
Madrid.
KETTUNEN, H., AND C. HELMKE.2005. Introduction
to Maya hieroglyphs. http://www.mesoweb.com/
resources/handbook/WH2005.pdf (acessed 18
September 2006).
KNIGHT,R. J. J. 2002. History, Distribution and
Uses.Pages 1–14 in A. W. WHILEY,B. SCHAFFER,
AND B. N. WOLSTENHOLME,EDS., Avocado:
Botany, Production and Uses.Wiley, New York.
LABAT,J. B. [1694] 1964. The avocado—in 1694.
Cal. Avoc. Soc. 1964 Yrbk.: 48—50.
LANDA, F.D. D. [1560] 1978. Relación de las Cosas
de Yucatán.11th ed. Editorial Porrua, México.
LEMUS,S. G., E. R. FERREYRA,M. P. GIL,B. P.
MALDONADO,G. C. TOLEDO,M. C. BARRERA,AND
J. M. CELEDON DE. 2005. El cultivo del palto.
Boletín INIA No. 129. Instituto de Investigaciones
Agropecuarias, La Cruz, Chile.
332 HARVARD PAPERS IN BOTANY Vol. 12, No. 2
2007 GALINDO-TOVAR ET AL., 10,000 YEARS OF AVOCADO HISTORY 333
LENTZ,D. 1991. Maya diets of the rich and poor:
Paleoethnobotanical evidence from Copan. Lat.
Am. Antiq. 2: 269–287.
LOVE,M. 2005. Los Olmecas en la costa sur de
Guatemala. Conferencias del Museo Popol Vuh
2005–4. www.popolvuh.ufm.edu.gt/Love2005.pdf
(acessed October 28, 2006).
MACNEISH,R. S. 1964. Ancient Mesoamerican civ-
ilization. Science. 143: 531–537.
MCPHERSON,W. 1955. Early account of the avoca-
dos in America. Cal. Avoc. Soc. 1955 Yrbk. 39:
87–88.
MENDOZA,C. 1989. Colección de Mendoza o Códice
Mendocino: documento mexicano del siglo XVI
que se conserva en la biblioteca de Bodleiana de
Oxford, Inglaterra. /facsímile fototípico dispuesto
por Francisco del Paso y Troncoso; edición acom-
pañada de una introducción, anotaciones y comen-
tarios por J. Galindo y Villa. Editorial Innovación,
México.
MESSER,E. 2003. Plantas alimenticias Zapotecas:
Transformación de dos culturas. 3rd ed. Pages
312–337 in J. LONG,ED., Conquista y comida.
Consecuencias del encuentro de dos mundos.
Universidad Nacional Autónoma de México,
México.
MIJARES, P., AND L. LÓPEZ. 1998. Variedades de
aguacate y su producción en México. Memoria
Fundación Salvador Sánchez Colín CICTAMEX
S.C., México.
MILLER, A., AND B. SCHAAL.2005. Domestication of
aMesoamerican cultivated fruit tree, Spondias
purpurea.PNAS 102: 12801–12806.
NAKASONE, H. Y. 1976. Breeding and disease prob-
lems in some tropical and subtropical fruits with
emphasis on papaya. In R. FRANCIOSI AND O.
DUARTE,EDS., 1st International Symposium of
Tropical and Subtropical Fruits. Lima, Peru, 1
October 1976. ISHS Acta Hort. 57.
NAVARIJO,L. 1995. Toponimia ornitológica mexi-
cana.Cuadernos del Instituto de Biología; 28.
UNAM Instituto de Biología, Mèxico.
ORTÍZ, P., AND M. C. RODRÍGUEZ. 2000. The sacred
hill of El Manatí: A preliminary discussion of the
site’s ritual paraphernalia.Pages 75–93 in J. E.
CLARK AND M. E. PYE,EDS., Olmec Art and
Archaeology in Mesoamerica,National Gallery of
Art, Washington, D.C..
OSSENBACH,C. 2005. History of orchids in Central
America. Part I: From Prehispanic times to the
independence of the new republics. Harvard Pap.
Bot. 10: 183–226.
PASO Y TRONCOSO,F. 1905. Suma de la visita de los
pueblos por orden alfabético, anónimo de la mitad
del siglo XVI.Papeles de la Nueva España.
Impresora de la Real Casa, Madrid.
POPENOE,W. 1963. Early history of avocado. Cal.
Avoc. Soc. 1963 Yrbk. 47: 19–24.
POPENOE,W., G. A. ZENTMYER,AND E. SCHIEBER.
1997. The avocado has many names. Cal. Avoc.
Soc. 1997 Yrbk. 81: 155–162.
POZORSKI,S. G. 1979. Prehistoric diet and subsis-
tence of the Moche Valley, Peru. World Archaeol.
II(2): 163–185.
RAMAMOORTHY,T. P., R. BYE,A. LOT,AND J. FA.
1993. Biological diversity of Mexico: Origins and
distribution.Oxford University Press, New York.
RAVEN,P. H., AND D. I. AXELROD. 1974. Angiosperm
biogeography and past continental movements.
Ann. Mo. Bot. Gard. 61: 539–673.
RENNER,S. S. 2004. Variation in diversity among
Laurales, Early Cretaceous to present. Biol. Skr.
55: 441–458.
ROBELO,C. A. 1951. Diccionario de Mitología
Nahuatl.2nd ed. Ediciones Fuente Cultural,
México.
SAHAGÚN,F. B. [1570] 2002. Historia general de las
Cosas de la Nueva España.Cien de México,
México.
SÁNCHEZ-COLÍN,S., P. MIJARES-OVIEDO,L. LÓPEZ-
LÓPEZ,AND A. F. BARRIENTOS-PRIEGO.2001.
Historia del aguacate. Memoria Fundación
Sánchez Colín CICTAMEX S.C. México.
SCHELE,L. 1974. Observations on the cross motif at
Palenque. Primera mesa redonda de Palenque.
Pages 41–61 in M. G. ROBERTSON,ED., Pre-
Columbian ArtResearch.R. L. Stevenson School,
Pebble Beach, California
––––––. 1998. The Code of Kings.Scribner, New
York.
SCHIEBER, E., AND G. A. ZENTMYER.1992. Exploring
Persea in the Cucumatanes,Guatemala. Cal.
Avoc. Soc. 1992 Yrbk. 76: 165–168.
SCHROEDER, C. A.1968. Prehistoric avocados in
California. Cal. Avoc. Soc. 1968 Yrbk. 52: 29–34.
SCORA, R. W., AND B. O. BERGH.1992. Origin of and
taxonomic relationships within the genus Persea.
Pages 505–514 in C. J. LOVATT,ED., Proceedings
of the Second World Avocado Congress. Pebble
Beach, California.
SIAP. Servicio de Información y Estadística
Agroalimentaria y Pesquera. 2007. México.
www.siap.gob.mx/aagricola_siap/icultivo/index.js
p(acessed 23 September 2006).
Skidmore, J. 2005. Earliest complex culture in the
Americas. Mesoweb. Reports and news.
http://www.mesoweb.com/reports/caral2.html
(acessed 12 February 2007).
SMITH,B. D.2001. Documenting plant domestica-
tion: The consilience of biological and archaeo-
logical approaches. Proc. Nat. Acad. Sci. 98:
1324–1326.
SMITH,C. E. J. 1966. Archeological evidence for
selection in avocado. Econ. Bot. 20: 169–175.
––––––. 1969. Additional notes on pre-conquest avo-
cados in Mexico. Econ. Bot. 23: 135–140.
SOLIS,R. S., J. HAAS,AND W. CREAMER. 2001.
Dating Caral, a preceramic site in the Supe Valley
onthe Central Coast of Peru. Science 292:
723–726.
STOREY,W. B., B. BERGH,AND G. A. ZENTMYER.
1986. The origin, Indigenous range, and dissemi-
nation of the avocado. Cal. Avoc. Soc. 1986 Yrbk.
70: 127–133.
TAKASHI,T. 1968. The aguacate im Mexico. Cal.
Avoc. Soc. Yrbk. 52: 169–172.
TAUBE,K. A. 2004. Olmec art at Dumbarton Oaks.
Dumbarton Oaks, Washington, D.C.
TELLIZ,D. 2000. El aguacate y su manejo integrado.
MundiPrensa S.A. de C.V., México.
VAN DER WERF,H. 2002. A synopsis of Persea
(Lauraceae) in Central America. Novon 12:
575–586.
VÁSQUEZ-DÁVILA,M. A. 2001. Etnoecología Chontal
de Tabasco México. EtnoecologíaVI(8): 43–60.
VEGA,I. G. DELA.[1605] 1995. Comentarios Reales
de los Incas.2nd ed. Fondo de Cultura
Económica, México.
WEBBER,H. J. 1936. An early illustration of the avo-
cado. Cal. Avoc. Assoc. 1936 Yrbk. 21: 39–41.
WILLIAMS,L. O. 1976. The botany of the avocado
and its relatives. Pages 9–15 in J. W. SAULS,R. L.
PHILLIPS,AND L. K. JACKSON,EDS., Proceedings of
the International Tropical Fruit Short Course. The
Avocado. Miami Beach, Florida, November 5–10,
1976. University of Florida, Gainesville.
––––––. 1977. The avocados, a synopsis of the genus
Persea, sub. Persea.Econ. Bot. 31: 315–320.
WOLF,E. R. 1967. Pueblos y culturas de Meso-
america.Traduccion de Felipe Sarabia. Editorial
Era, Mexico.
WOLTERS,B. 1999. Dispersion and ethnobotany of
the cacao tree and other Amerindian crop plants.
Angn. Bot. 73: 128–137.
XIMÉNEZ,F. F. [1615] 2001. Cuatro Libros de la
Naturaleza y Virtudes de las Plantas y Animales,
de Uso Medicinal en la Nueva España.Imprenta
de Juan Pablos, S.A., México.
334 HARVARD PAPERS IN BOTANY Vol. 12, No. 2
... Our characterization has further highlighted the admixed origin of seedling rootstocks currently used at commercial orchards in the northwest Andes. Persistent admixture due to rampant gene flow is expected for a species that, as avocado, has been subjected to continent-wide animal and humanmediated migration (Bergh and Ellstrand, 1986;Galindo-Tovar et al., 2007;Larranaga et al., 2020), besides being an obligate outcrossing (via protogynous dichogamy, a sequential nonoverlapping hermaphroditism in which female function precedes male function). Regarding economical traits, the Guatemalan race typically has small seeds and exhibits late fruit maturity, whereas Mexican race shows early fruit maturity and cold tolerance. ...
... Across Mesoamerica and northern South America, avocado trees are still cultivated in traditional orchards, backyard gardens, and as living fences. They are consumed at a regional scale and harbor a strong potential to improve fruit quantity and quality, besides tree adaptation, when used as rootstocks in commercial "Hass" orchards (Galindo-Tovar et al., 2007). However, for this to occur, a better comprehension of the consequences of grafting, more concretely the rootstock-scion interaction across traits and environments, needs to be achieved, just as envisioned here. ...
... Our characterization has further highlighted the admixed origin of seedling rootstocks currently used at commercial orchards in the northwest Andes. Persistent admixture due to rampant gene flow is expected for a species that, as avocado, has been subjected to continent-wide animal and humanmediated migration (Bergh and Ellstrand, 1986;Galindo-Tovar et al., 2007;Larranaga et al., 2020), besides being an obligate outcrossing (via protogynous dichogamy, a sequential nonoverlapping hermaphroditism in which female function precedes male function). Regarding economical traits, the Guatemalan race typically has small seeds and exhibits late fruit maturity, whereas Mexican race shows early fruit maturity and cold tolerance. ...
... Across Mesoamerica and northern South America, avocado trees are still cultivated in traditional orchards, backyard gardens, and as living fences. They are consumed at a regional scale and harbor a strong potential to improve fruit quantity and quality, besides tree adaptation, when used as rootstocks in commercial "Hass" orchards (Galindo-Tovar et al., 2007). However, for this to occur, a better comprehension of the consequences of grafting, more concretely the rootstock-scion interaction across traits and environments, needs to be achieved, just as envisioned here. ...
Article
Full-text available
Grafting is typically utilized to merge adapted seedling rootstocks with highly productive clonal scions. This process implies the interaction of multiple genomes to produce a unique tree phenotype. However, the interconnection of both genotypes obscures individual contributions to phenotypic variation (rootstock-mediated heritability), hampering tree breeding. Therefore, our goal was to quantify the inheritance of seedling rootstock effects on scion traits using avocado (Persea americana Mill.) cv. Hass as a model fruit tree. We characterized 240 diverse rootstocks from 8 avocado cv. Hass orchards with similar management in three regions of the province of Antioquia, northwest Andes of Colombia, using 13 microsatellite markers simple sequence repeats (SSRs). Parallel to this, we recorded 20 phenotypic traits (including morphological, biomass/reproductive, and fruit yield and quality traits) in the scions for 3 years (2015-2017). Relatedness among rootstocks was inferred through the genetic markers and inputted in a "genetic prediction" model to calculate narrow-sense heritabilities (h 2) on scion traits. We used three different randomization tests to highlight traits with consistently significant heritability estimates. This strategy allowed us to capture five traits with significant heritability values that ranged from 0.33 to 0.45 and model fits (r) that oscillated between 0.58 and 0.73 across orchards. The results showed significance in the rootstock effects for four complex harvest and quality traits (i.e., total number of fruits, number of fruits with exportation quality, and number of fruits discarded because of low weight or thrips damage), whereas the only morphological trait that had a significant heritability value was overall trunk height (an emergent property of the rootstock-scion interaction). These findings suggest the inheritance of rootstock effects, beyond root phenotype, on a surprisingly wide spectrum of scion traits in "Hass" avocado. They also reinforce the utility of polymorphic SSRs for relatedness reconstruction and genetic prediction of complex traits. This research is, up to date, the most cohesive evidence of narrow-sense inheritance of rootstock effects in a tropical fruit tree crop. Ultimately, our work highlights the importance of considering the rootstock-scion interaction to broaden the genetic basis of fruit tree breeding programs while enhancing our understanding of the consequences of grafting.
... Diversifying selection of half-sib rootstock families and within-family o spring at seedling nurseries remains a promising and a ordable avenue, especially if coupled with indirect genetic-guided predictors. Seedling rootstock breeding also enables broadening the genetic basis of young avocado plantations in regions with diverse sources of native avocado trees, some of them still cultivated in backyard gardens, traditional orchards, and as living fences (Galindo-Tovar et al., 2007). Yet, selection of diverse clonal rootstock genotypes may be a more appealing long-term strategy to contribute diminishing tree mortality by conveying genotypic superiority and phenotypic uniformity, while controlling genetic erosion (Ingvarsson and Dahlberg, 2019). ...
... Diversifying selection of half-sib rootstock families and within-family offspring at seedling nurseries remains a promising and affordable avenue, especially if coupled with indirect genetic-guided predictors. Seedling rootstock breeding also enables broadening the genetic basis of young avocado plantations in regions with diverse sources of native avocado trees, some of them still cultivated in backyard gardens, traditional orchards, and as living fences (Galindo-Tovar et al., 2007). Yet, selection of diverse clonal rootstock genotypes may be a more appealing long-term strategy to contribute diminishing tree mortality by conveying genotypic superiority and phenotypic uniformity, while controlling genetic erosion (Ingvarsson and Dahlberg, 2019). ...
Article
Full-text available
Grafting induces precocity and maintains clonal integrity in fruit tree crops. However, the complex rootstock × scion interaction often precludes understanding how the tree phenotype is shaped, limiting the potential to select optimum rootstocks. Therefore, it is necessary to assess (1) how seedling progenies inherit trait variation from elite 'plus trees', and (2) whether such family superiority may be transferred after grafting to the clonal scion. To bridge this gap, we quantified additive genetic parameters (i.e., narrow sense heritability-h 2 , and genetic-estimated breeding values-GEBVs) across landraces, "criollo", "plus trees" of the super-food fruit tree crop avocado (Persea americana Mill.), and their open-pollinated (OP) half-sib seedling families. Specifically, we used a genomic best linear unbiased prediction (G-BLUP) model to merge phenotypic characterization of 17 morpho-agronomic traits with genetic screening of 13 highly polymorphic SSR markers in a diverse panel of 104 avocado "criollo" "plus trees." Estimated additive genetic parameters were validated at a 5-year-old common garden trial (i.e., provenance test), in which 22 OP half-sib seedlings from 82 elite "plus trees" served as rootstocks for the cv. Hass clone. Heritability (h 2) scores in the "criollo" "plus trees" ranged from 0.28 to 0.51. The highest h 2 values were observed for ribbed petiole and adaxial veins with 0.47 (CI 95%0.2-0.8) and 0.51 (CI 0.2-0.8), respectively. The h 2 scores for the agronomic traits ranged from 0.34 (CI 0.2-0.6) to 0.39 (CI 0.2-0.6) for seed weight, fruit weight, and total volume, respectively. When inspecting yield variation across 5-year-old grafted avocado cv. Hass trees with elite OP half-sib seedling rootstocks, the traits total number of fruits and fruits' weight, respectively, exhibited h 2 scores of 0.36 (± 0.23) and 0.11 (± 0.09). Our results indicate that elite "criollo" "plus trees" may serve as promissory donors of seedling rootstocks for avocado cv. Hass orchards due to the inheritance of their outstanding trait values. This reinforces the feasibility to leverage natural variation from "plus trees" via OP half-sib seedling rootstock families. By jointly estimating half-sib family effects and rootstock-mediated heritability, this study promises boosting seedling rootstock breeding programs, while better discerning the consequences of grafting in fruit tree crops.
... Avocado was domesticated in Mesoamerica where archaeological records in the Tehuacán region (Puebla State, Mexico) showed that man consumed avocado about 10,000 years ago (Smith, 1966). This fruit tree was cultivated and domesticated by the Mokayas, the first Mesoamerican culture, who passed the knowledge to the next cultures including the Mayas and Olmecs (Galindo-Tovar et al., 2007). The larger seeds in the latest archaeological strata at the Tehuacán (Puebla State, Mexico) and Oaxaca Valley (Oaxaca State, Mexico) sites propose that avocado selection may have started about 4,800 to 6,000 years ago (Smith, 1966(Smith, , 1969. ...
Thesis
Full-text available
Avocado (Persea americana Mill) has a long cultivation history in Tanzania and adds to the country’s revenue through trading within (local and commercial avocado cultivars) and outside (commercial cultivars) the country. Research has been, however, scanty to characterize the country’s avocado production, local trading and diversity. In order to contribute to this research area, the present study was conducted in eight districts of Tanzania’s southern highlands. For the purpose of describing aspects of avocado production and trading throughout the value chain, 275 avocado growers, 231 traders and 16 key informers were interviewed. As for the diversity study, 226 seed-originated avocado trees were sampled in the eight districts and 14 morphological and 10 simple sequence repeat (SSR) loci genetic characters were evaluated. The analyses of the data generated through the interviews showed that the average avocado yield per district varied from 52 to 156 kg/plant. Approximately, 28 and 79% of the growers and the traders were content with the earnings from the avocado trading. The majority of the growers and traders mentioned some challenges affecting the value chain including limited extension services, drought, pathogens and pests, poor marketing environment, and the short shelf life of the fruits. Resolving these issues is the key towards the improvement of the earnings and living standards of avocado farmers and traders, which is also applicable to many other crops sharing similar challenges. The sampled trees showed great variation in the investigated morphological features. These included 11, 21 and 17 different shapes of the leaf, mature fruit and seed, respectively. As for the genetic characteristics, an average of 16.7 ± 1.3 alleles per locus, 0.65 ± 0.04 and 0.84 ± 0.02 observed and expected heterozygosity, respectively, and a Shannon’s information index of 2.17 ± 0.10 were obtained. These results imply a high genetic diversity of the trees investigated. Most of the observed morphological and genetic characters were not unique for a particular area as revealed by the district/region-free clustering of the trees in the principal coordinates and components analyses or by dendrograms. Introduction of highly similar (overlapping) germplasm to more than one districts and cross-transfer of seeds (propagules) among the districts/regions could be the reason for this distribution.
... Soil used as inoculum was collected from three environments with different land uses: pristine, backyard, and intensive. The soil samples were collected from the following locations: a) pristine, mature, > 150-year-old, midelevation evergreen forest in Calakmul, Campeche, Mexico (18° 9′ 48″ N; 89°23′ 53″ W) previously described by García-Licona et al. (2014) and considered the best conserved environment of the original distribution of Persea americana in the northern neotropics (Galindo-Tovar et al. 2007); b) backyard, a typical ~ 50-year-old traditional Mayan family backyard described Montagnini (2006) located in Kobén, Campeche, Mexico (19° 55′ 02″ N; 90° 24′ 57″ W); and c) intensive, ~ 30-year-old orchard monoculture of P. americana var. Hass managed with a technological package, including mechanization, the use of synthetic fertilizers, pesticides, and herbicides, located in San Juan Nuevo Parangaricutiro, Michoacán, Mexico (19° 25′ 09″ N; 102° 08′ 14″ W). ...
Article
Full-text available
To explore the capability of soil mycobiota to degrade avocado peel waste and identify relevant successions and trophic guild shifts, fungal communities from three environments with different land uses were evaluated in a solid-state process. Soil samples used as inoculum were collected from a pristine mature tropical forest, a traditionally managed Mayan land, and an intensively managed monospecific avocado plantation. Soil-substrate mixes were evaluated for 52 weeks to evaluate organic matter decay and the carbon-to-nitrogen ratio. Amplicon-based high-throughput sequencing from internally transcribed spacer (ITS) analysis revealed significant differences in fungal communities widely dominated by Fusarium sp. and Clonostachys sp.; however, less represented taxa showed relevant shifts concomitantly with organic matter content drops. Trophic guild assignment revealed different behaviors in fungal communities between treatments over the 52 weeks, suggesting distinct preconditioning of fungal communities in these environments. Overall, the results lead to the identification of promising degradation moments and inoculum sources for further consortia enrichment or bioprospecting efforts.
... Since prehistoric times, avocado (Persea americana Mill.) has played an important role in the history of many cultures, specially in Mesoamerica and latter along the Peruvian coast (Galindo-Tovar et al., 2007). This tropical fruit is native to Central and South America, and it is increasingly consumed worldwide especially for its high nutritive composition (Pineda-Lozano et al., 2021). ...
Article
Full-text available
The production of avocado in Peru has been increasing over the last five years and it represents a major export commodity. However, dieback symptoms, typical of Botryosphaeriaceae, have been observed in avocado orchards throughout the country, leading to economic losses. Thus, this study aimed to evaluate the diversity and pathogenicity of species associated with dieback of avocado trees, in the Peru's Pacific coastline. On the basis of morphological and cultural characters, as well as phylogenetic analysis of the internal transcribed spacer region (ITS) and part of the translation elongation factor 1-α gene (tef1-α), the isolates were identified as belonging to three Lasiodiplodia species , namely L. theobromae, L. laeliocattleyae and L. pseudotheobromae. Inoculation in cv. 'Hass' plants confirmed the pathogenicity of Lasiodiplodia species and revealed differences in aggressiveness among species and isolates. Lasiodiplodia laeliocattleyae is reported for the first time in avocado trees associated with dieback symptoms, which together with L. theobromae and L. pseudotheobromae may represent a threat to Peruvian avocado plantations.
Article
With the aim of recognizing and identifying both fertile and sterile specimens of Persea, leaf venation patterns of 54 taxa were studied, 36 of them being described and illustrated for the first time. Analysis of a matrix of 12 leaf venation characters and 31 states distinguishes the taxa and can be used to detect misidentifications. Additionally, lectotypes for the names Laurus cordata, L. lingue (second-step), L. subcordata, Menestrata racemosa, Persea cordata var. microphylla, P. donnell�smithii, P. laevigata, P. lingue, P. nivea and P. rugosa are designated, and the identity of Persea microneura and the status of P. jariensis are clarified.
Preprint
Full-text available
Grafting is typically utilized to merge adapted seedling rootstocks with highly productive clonal scions. This process implies the interaction of multiple genomes to produce a unique tree phenotype. Yet, the interconnection of both genotypes obscures individual contributions to phenotypic variation (i.e. rootstock-mediated heritability), hampering tree breeding. Therefore, our goal was to quantify the inheritance of seedling rootstock effects on scion traits using avocado (Persea americana Mill.) cv. Hass as model fruit tree. We characterized 240 rootstocks from 8 avocado cv. Hass orchards in three regions of the province of Antioquia, in the northwest Andes of Colombia, using 13 microsatellite markers (simple sequence repeats-SSRs). Parallel to this, we recorded 20 phenotypic traits (including morphological, eco-physiological, and fruit yield and quality traits) in the scions for three years (2015-2017). Relatedness among rootstocks was inferred through the genetic markers and inputted in a 'genetic prediction' model in order to calculate narrow-sense heritabilities (h2) on scion traits. We used three different randomization tests to highlight traits with consistently significant heritability estimates. This strategy allowed us to capture five traits with significant heritability values that ranged from 0.33 to 0.45 and model fits (R2) that oscillated between 0.58 and 0.74 across orchards. The results showed significance in the rootstock effects for four complex harvest and quality traits (i.e. total number of fruits, number of fruits with exportation quality, and number of fruits discarded because of low weight or thrips damage), while the only morphological trait that had a significant heritability value was overall trunk height (an emergent property of the rootstock-scion interaction). These findings suggest the inheritance of rootstock effects, beyond root phenotype, on a surprisingly wide spectrum of scion traits in 'Hass' avocado. They also reinforce the utility of SSR markers for relatedness reconstruction and genetic prediction of complex traits. This research is, up to date, the most cohesive evidence of narrow-sense inheritance of rootstock effects in a tropical fruit tree crop. Ultimately, our work reinforces the importance of considering the rootstock-scion interaction to broaden the genetic basis of fruit tree breeding programs, while enhancing our understanding of the consequences of grafting.
Chapter
This multiauthor book consists of 14 sections covering all aspects of avocado production and science, including the uses, taxonomy, botany, genetics, breeding, ecology, growth and development, propagation, cultivars and rootstocks, cultivation, nutrition, biotechnology, pests and diseases (and their control), and harvesting and postharvest aspects (including storage, packing and transport) of avocado. The book will be of interest to researchers, academics, students, professional growers and consultants.
Article
The vegetation of northern Latin America is a complex of assemblages arriving by different routes and at various times, depending upon the physical conditions and climates prevailing along the North Atlantic and Panamanian land bridges. The result is a modern lowland and lower montane vegetation in northern Latin America that includes four biogeographic components: 1) plants of Gondwanan; 2) Laurasian ancestry that arrived via the North Atlantic and, to a lesser extent, across Beringia, during the late Mesozoic and Paleogene; 3) ancient Gondwanan elements established in South America before separation from Africa, and moving into northern Latin America during the late Mesozoic and Paleogene; and 4) Gondwanan (South American) introductions across the isthmus in Neogene times. -from Author
Article
Mexico is one of the botanically richest countries in the New World, with two major floristic kingdoms represented by ten vegetation types and 30 000 species. Ethnobotanical richness is reflected by the utilization of >5000 vascular plants. Human enterprises resulting in erosion, livestock grazing, and agriculture alter the Mexican landscape and threaten its botanical diversity. The gathering, incipient management, and cultivation of wild, weedy and domesticated plants produce positive as well as negative effects. -from Author
Article
There are 26 chapters, organized in six parts. The first section consists of background material such as geological and historical factors, including an essay on the diversity of Mexico's flowering plant flora and its origins. The second and third parts contain chapters on selected major faunistic and floristic groups. Several contributions in these sections discuss diversity (species richness), endemism, and distribution, among other themes. The fourth part deals with phytogeographical patterns in contrasting ecosystems: the tropical rain forests and the alpine floras of Mexico. The fifth section presents two ethnobiological essays: one on the influence of humans on plant species diversification, the other on aspects of the domestication of plants in Mexico. The last section provides a brief scenario of Mexican biodiversity and a review of terrestrial habitats in Mexico. Most chapters are abstracted separately. -from Publisher
Article
Amerindian coastal shipping along the Pacific coast of America is neglected for the dispersion of pre-columbian crop plants up to now. Indians of the Valdivia Culture and their successors traveled by boat or raft along the coasts from their west ecuadorian home to Peru and Middle America since 2200 B.C. and to Southern Mexico since 1450 B.C. The traveling ceased with the arrival of Spaniards in 1526. Presumably crop plant export occurred from Western Ecuador to Peru and Middle America (sweet manioc, Annona cherimola, Carica papaya, early great-grained corn) and Mexico (tobacco). On the other hand the pre-columbian crops (Persea americana, Capsicum annuum) evolved in Mexico were brought to Western Ecuador and neighboring Northern Peru. Shipment by sea would be faster and thus more successful than the time consuming transportation by land all the way through Colombia and Central America. Looking at the distribution map of sweet manioc only shipment of living cuttings appears as a feasible explanation. Similarly one may hypothesize that living plants of Theobroma cacao were shipped from South to Middle America on this Pacific searoute. Since neither wild cacao trees nor primitive cultigens were found in the area between Ecuador and Guatemala, thus the landroute is highly unlikely. In the Mesoamerican Pre-classic Culture region the existence of cacao tree is proven since 500 B.C., about thousand years later than the first contact of west-ecuadorian Indians with Mexico. The use of T. cacao in indigenous medicine is reported. Surprisingly there is no evidence for pre-columbian domestication of cacao in its natural habitat Amazonia. This may be explained by the great popularity of similar but much stronger acting drugs like the caffeine drug Guarana and the cocaine drug Erythroxylum coca var. ipadu.
Article
: Avocados are a subtropical climacteric fruit that is generally oblong in shape and green skinned, although ‘Hass’, the dominant commercial cultivar, ripens to purple-black. They are one of the few fruits that contain significant amounts of oils (fatty acids). The high concentrations of monounsaturated fatty acids and other phytochemicals make avocado a very healthy and nutritious fruit. Avocados do not ripen unless removed from the tree, meaning fruit can be mature but ‘tree stored’ for as long as 12 months. Increased commercial maturity is associated with increased fruit size, oil and dry matter content, and decreased ripening times. Avocados have a relatively short storage life, limited primarily by the expression of internal chilling injury (CI) symptoms, but rots are a limitation for fruit grown in many countries. Optimum storage temperature is generally around 6 °C, and temperatures below 3 to 4 °C (depending on cultivar and time in the season) lead to external CI (skin blackening). Controlled atmosphere storage and 1-MCP (SmartFreshSM) are effective commercial tools for improving storage life. Since avocado cannot be consumed unripe, an increasingly important commercial tool is the use of ethylene to hasten and synchronize ripening. New processing options, such as cold-pressed oil extraction and high-pressure processing, are receiving increasing attention.