New Forests 27: 2538, 2004.

2004 Kluwer Academic Publishers. Printed in the Netherlands.

(Brosimum alicastrum
The propagation of Ramon
Sw.; Moraceae) in Mayan homegardens of the
Yucatan peninsula of Mexico

A.R. GILLESPIE 1, *, D.M. BOCANEGRA-FERGUSON 1 and J.J. JIMENEZ-

OSORNIO 2
1
Department of Forestry and Natural Resources, Purdue University, 195 Marstellar Street, West
Lafayette, IN 47907 -2033, USA; 2 Department of Management and Conservation of Tropical Natural
Resources, PROTROPICO, Faculty of Veterinary Medicine and Zoology, University of Yucatan, Merida,
Mexico; * Author for correspondence

Received 25 January 2002; accepted in revised form 15 January 2003

Key words: Agroforestry, Forage production, Indigenous knowledge, Plantation establishment, Seed
germination

Abstract. In the Yucatan peninsula of Mexico, Maya natives have been propagating important species for
centuries. However, little documentation exists of these methods, and traditional knowledge is fading as
younger generations seek work in urban centers. With growing interest in using some of these species for
plantation-scale production, this knowledge should be captured to aid in propagation and cultural
methods of selected species. One such species, Ramon (Brosimum alicastrum Sw.), is grown in Mayan
homegardens primarily as a source of dry season forage. We conducted a survey of Mayan families in
several municipalities to determine basic propagation habits and procedures for cultivating Ramon, and
used this information to conduct some controlled-environment studies of reproductive ecology of the
species. Our survey showed that all or most Maya grow the Ramon tree in homegardens for forage and
that the tree is both cultivated from wild seedlings as well as planted on a small scale. Propagation is by
seed and young seedlings are irrigated until they are established. We examined the effects of temperature
on seed germination and found that maximum germination occurred between 278 and 38 8C, with best
germination and growth of healthy seedlings at 33 8C. No germination occurred below 218 or above 44
8C. Simulated Maya irrigation gave a four-fold increase in growth rate of seedlings, aiding early
establishment. Watering gave seedlings of greater biomass, greater stem and root length, and greater
numbers and size of leaves. This magnitude of response may provide an economic return for plantation
culture of Ramon for forage production.

Introduction

In the Yucatan Peninsula of Mexico, the Mayan culture has used homegarden
agroforestry for centuries. Homegardens, found in many cultures around the world,
are a diverse mixture of vegetables, fruits, medicinal plants, fodder plants, shrubs,
and trees cultivated around the family home. These agroforestry systems are often
maintained in association with slash and burn agricultural practices, which provide
the bulk of caloric intake for a family. As such, homegardens are critical for dietary
balance and family health as well as production of food and goods that may be sold
to supplement income. And the Mayan agroforestry practice of homegardening is
26

one agroforestry system that can help sustain the indigenous families of the Yucatan
peninsula of Mexico and related regions (De Clerck and Negreros-Castillio 2000;
Levasseur and Olivier 2000).
Interest has developed in understanding how the Maya manage homegardens and
what environmental factors they take into account to improve the survival and
growth of their plants. We know that they have developed a detailed soil classifica-
tion system (Dunning 1992) and propagate many species important to their
livelihood and culture, but much is unknown about their traditions and abilities in
domesticating specific species and how these efforts might be improved, despite
their being studied or described since the 1500s (e.g., De Landa in Tozzer 1941).
Among the many species found in Mayan homegardens, the Ramon tree
(Brosimum alicastrum Sw.; Moraceae) has been present for centuries. Anth-
ropological studies have found large assemblages of this species in the old ruins of
the many city-states in the Yucatan Peninsula, Chiapas, and Tabasco states of
Mexico, and ruins in Guatemala and Honduras, among other sites. While this species
likely played a role in providing medicines and food for humans and animals during
times of shortage, we know today that the high density of this tree in archaeological
ruins is due to its ecological habit of colonizing cleared areas (Miksicek et al. 1981;
Lambert and Arnason 1982; Puleston 1982; Peters 1983). Though the Ramon tree
was likely to have been important for several reasons in the past, the Ramon tree
found in modern Mayan homegardens is mainly used as forage for livestock.
Several recent studies have found that there is high nutritional value in leaves, twigs,

and seeds for humans and animals alike (Pardo-Tejeda and Sanchez-Munoz 1980;

Perez et al. 1995; Santos Ricalde and Abreu Sierra 1995). The Ramon tree also has
other uses, such as shade for animals and humans, medicines, timber, and ornamen-
tal value (Chavelas Polito and Devall 1988). In 1975, The US National Academy of
Sciences, in a survey of important tropical plants, included the Ramon tree as a
promising species (Peters and Pardo-Tejeda 1982).
Although the Maya intensively manage homegardens today (Gillespie et al.
1993), the Ramon tree is not planted, as are most other important species. Rather,
the species falls within a small group of native plants that are tolerated when they
seed in because of their importance (Benjamin et al. 2001). In the lowland area of
the Yucatan Peninsula of Mexico, the tree grows in an environment that is semiarid
and characterized by low precipitation, thin soil cover with rocky limestone
outcrops, and little to no surface water. Also, there is rapid water percolation due to
the karst topography and porous bedrock of the area. As such, growing conditions
for plant species are difficult, and species that thrive in such environments can be
important for rural development purposes. Additionally, an interest in cultivating the
species in a plantation setting as a perennial and dry-season source of forage is
growing in the region and throughout Latin America (Ayala and Sandoval 1995;

Perez et al. 1995). Thus, it becomes relevant to research the traditional management
of the species to illuminate patterns of successful propagation and management and
to develop new information to promote the use of the tree as a plantation crop. This
project attempts to 1) gain an understanding of the traditional knowledge of Maya
people regarding the management and survival of the Ramon tree in the sisal (Agave
 27

fourcroydes Lemarie)-growing region of the Yucatan, and 2) to determine some

basic reproduction ecology of the species.

Materials and methods

Homegarden propagation

Survey data on traditional propagation techniques were gathered in the areas of the
communities of Hocaba, Sahcaba, and Mani in the sisal region of the Yucatan
peninsula of Mexico, in the state of Yucatan. The interviews were conducted with
indigenous people, mainly focused on seniors from 40 to 801 years of age. The data
obtained in these communities were based on a total of 18 interviews with Maya
natives, 7 men and 11 women. Additional, supporting information was gathered
from interviews with a group of approximately 20 indigenous farmers in the Escuela
Campesina (Farmers School) located in the town of Mani in the southern part of the
state of Yucatan. These farmers were representatives of their individual com-
munities from all over the state, sent to the school to learn sustainable production
techniques. Their data were used to substantiate and clarify practices in the
homegardens.
In the communities of Hocaba, Sahcaba, and Mani, interviews were conducted in
the homegardens. Most often women were interviewed as women spend most of the
day in the home taking care of children, cooking, and maintaining the homegarden
while men spend most of their time working in the milpa (slash and burn agricultural
plots for maize production) or in other activities. In the majority of cases, the women
interviewees were either the mother or the grandmother of the household. If the
interviewee was a man, this person was the father or the grandfather. Several visits
were made to each garden.
The interview was informal. Though all questions or topics were covered with
each individual, they were not structured so as to provide the freedom to explore
different paths of knowledge. The interviews were conducted mainly in Spanish. On
some occasions, the person interviewed spoke only Maya, therefore, a translator was
required. The original interviews were recorded on tape and manually transcribed.
The interviews were analyzed qualitatively and are summarized here.
To determine specific Ramon cultural practices, direct observation of family-
member practices was also conducted in addition to interviews (Patton 1990). Direct
observations were done the same day as interviews. The direct observation was a
simple, non-structured observation, conducted within the homegarden area, with
some of the members of the family. For all sites, permission was granted by the
owners of the house to enter and make observations. Observations included note-
taking, photography, and videography. These observations were compared or cross-
referenced with the information obtained from all interviews and from literature
review.
28

reproduction ecology
Ramon

Maya interviews stressed the importance of propagation by seed and the need for
early irrigation. Thus, a series of controlled-environment experiments was con-
ducted to evaluate environmental influence on early growth and survival of Ramon.
The first experiment was conducted in order to evaluate the germination potential of
Ramon seed under different temperature conditions. Fresh seeds were obtained from
the Mexican township of Xmatkuil in the Yucatan Peninsula, supplied by the
University of Yucatan, Mexico. Seed was collected from a small population of 10
trees planted more than 30 yr ago. Seed viability is quickly lost, so only seed
produced in the current year was collected from mature fruits at the end of the rainy
season. No pre-germination procedure was undertaken (e.g. scarification, dor-
mancy). Seeds were surface-treated with a 70% ethanol solution to reduce possible
fungal infection. The seeds were then planted in 7 3 5 cm pots containing a
commercial potting media (Metromix-400), a peat moss-vermiculite mix. The seeds
were planted 2 cm below the soil surface and then placed in temperature-controlled
chambers.
Seven growth chambers were set at day / night temperatures of 16 / 13 8C, 21 / 19
8C, 27 / 24 8C, 33 / 30 8C, 38 / 35 8C, 44 / 41 8C, and 49 / 47 8C. A sample size of
thirty-four pots (one seed per pot) per treatment was used. Seed germination data
were collected each day for 37 d. At the end of the test period, pots were evaluated
for overall number of seeds germinated in each treatment.
A second experiment was undertaken to determine the early growth and survival
of Ramon under the dry conditions native to the Yucatan, and under a simulated
irrigation regime used by the Maya in propagating plants in their homegardens.
Maya will hand irrigate (traditional) or hose irrigate (new with the advent of potable
water piped directly to each household property) seed and newly transplanted
seedlings for a period of time specific to each plant species and the soil type in
which it is planted. For this experiment, a group of seeds, surface-treated with 70%
ethanol, was germinated in small pots (7 3 5 cm) in the greenhouse with the
addition of bottom heat at 32.2 8C to increase soil temperature to a level consistent
with the optimum germination seen in the first experiment.
As seeds started to develop radicles, the seedlings were transplanted to the final
pots. Two types of soil were chosen for the experiment, the commercial potting soil
and a loamy sand soil (topsoil) obtained from a local commercial greenhouse. The
topsoil and potting soil were steam sterilized to kill all possible weeds and fungi.
Two treatments were established for each soil type in a randomized block design.
Each block contained two treatments, one with simulated Maya irrigation conditions
(soil water potential maintained near 1 bar or 100 kPa) and the other with significant
water stress, the simulated natural condition of the Yucatan peninsula (soil water
potential maintained near 10 bars or 1 MPa).
To determine the water potential for each treatment, a calibration curve of water
potential vs. water content was constructed using a pressure plate tension system
(Soilmoisture Equipment Co., Goleta, CA, USA). The soil moisture content for each
treatment was monitored with a soil moisture meter, a portable MC-302 Soil
 29

Moisture-Temperature meter (ELE International, Soiltest Products Division, IL,

USA). A set of fiberglass cells was connected to the meter through wire leads. Those
cells were buried at the level of the root zone in each pot. To maintain experimental
soil water potentials, the well-watered seedlings were irrigated every other day and
the water-stress treatment received water approximately once per month.
Several parameters were assessed at the end of the experiment, approximately
nine months of growth, to evaluate the influence of water potential on plant growth
and survival. These included biomass growth and allometric relationships. Carbon
allocation to shoots and roots in terms of dry mass was measured as plants were
harvested after 274 d. Ten seedlings per treatment and per soil type were randomly
chosen. The seedlings were divided into their organs (leaves, branches, and roots)
and wet weight was measured. Then they were placed in an oven and dried at 65 8C
until continuous measurements showed that tissues remained at a constant weight.
These measurements allowed estimation of the biomass (g) gained in each treat-
ment.
In terms of root and shoot growth, root weight (g) was measured as were seedling
height (cm), number of leaves, and root:shoot biomass ratios. Lastly, an analysis of
variance (ANOVA) was conducted for carbon allocation and total biomass variables
examining the influence of soil, irrigation, and their interaction.

Results and discussion

Homegarden propagation

Traditional knowledge, having evolved over time, is based on local experiences,

resources, and conditions (Morrison et al. 1996). This traditional knowledge base is
passed down from generation to generation, and lack of this transfer may cause loss
of traditional techniques and cultivated species. There is concern and evidence in the
Yucatan that local migration to the large cities of Merida and Cancun for employ-
ment is causing the loss of traditional knowledge for important species. Interviews
showed, however, that considerable knowledge of Ramon still exists with the older
Maya.
Table 1 provides the summary of responses of all individuals interviewed in the
homegardens and the classification of their answers by percentage. According to the
interviewees, the Ramon tree (Brosimum alicastrum Sw.) is present in all Mayan
homegardens located in the towns of Hocaba and Sahcaba, and is found in the
gardens of the farmer-students from across the state of Yucatan. This widespread
occurrence of the tree indicates its importance to the Maya family and the pastoral
economy of the region. It is interesting to note that the tree does not occur in the
secondary forest of the landscape, but only in cultivated or domesticated areas.
Several Maya interviewed noted this and explained it by stating that the tree is not
planted in these areas, that it is only found in areas of water wells and cenotes
(sinkholes with water). As the seed is recognized to be dispersed by bats and birds
30

(Table 1), an interaction between wildlife and land use may be responsible for its
current distribution in addition to the plantings of the Maya.
The species is highly desired as a source of food for livestock animals in the Maya
homegarden due to its excellent properties and nutritional value. Even animal
husbandry professionals and scientists in the region recognize its value and desire to
plant the species in high-yielding plantations (e.g., Ayala and Sandoval 1995). The
Maya give the forage to animals in corrals during the dry season or sell the rights to
the forage. Trees are divided into thirds and the forage rights sold accordingly. Sale
is to local villagers or sometimes to people who travel from larger cities that may
maintain animals on suburban properties.
Though not currently a source of food for the Maya, older interviewees re-
membered that previous generations used Ramon as a substitute or supplement for
maize. Fruits were eaten or used to prepare tortillas or fritters as well as being mixed
with beans. This agrees with early reports of Ramon use (Lundell 1938) when seeds
were still being boiled or made into flour for bread or tortillas. One Maya
remembered its use when locusts destroyed a maize crop some 60 yr earlier. The
traditional knowledge of its use as a food during famine was common. Less
common was the knowledge of past alternative uses. Reports were given of boiled
seeds being fed to cows and pigs to increase milk production as well as being mixed
with other plants in an infusion for increasing milk production of women. Other
reported medicinal uses included its use for tooth pain, respiratory diseases, and as
an anti-diarrheal for livestock. One Maya recognized that the tree produced a latex,
but did not know its use. This same interviewee remembered eating the seeds when
he was young.
The tree is usually propagated naturally or artificially by seed. Natural dispersion
of seed is mainly by bats or simply by the dropping of seeds from the parent tree.
Young farmers are now beginning to collect seed from the ground and to plant small
nurseries. The species is rarely managed (irrigated or fertilized) once established,
and several older Maya reported that their ancestors (at least the past several
generations) did not manage the species either. Though the Maya do not recognize it
as such, related research (Benjamin et al. 2001) indicates that the Maya keep a very
clean garden surface by burning household and garden waste, and the ash is left in
random spots, acting as a source of plant nutrients. What is undertaken as a form of
insect and pest control supplements the fertility of the homegarden. Animal waste is
also used sporadically as a fertilizer.
When artificially propagated by seed or seedling, interviewees agreed that the
best time to plant the seeds or transplant seedlings is during the rainy season;
otherwise, the seedlings should be watered until they are well established. All
interviewees agreed that the species is strong and that it can grow in any kind of soil,
including stony soils. This is the reason that active management is abandoned once
the trees have become established. The roots of the Ramon tree are recognized as
growing very deep, up to 15 m to reach the water table (Table 1). Extensive lateral
root growth and the ability to grow through rocky soil (lithic ustorthents or lithosols)
is also recognized by the Maya. In terms of soils that give highest productivity, it
was stated that the best soils for Ramon are Kancab (rhodic paleustalfs or eutric
 31

survey in the Mayan towns of Hocaba and Sahcaba, Yucatan, Mexico

Table 1. Summary of Ramon
QUESTION ANSWERS PROVIDED DISTRIBUTION SUMMARY
PERCENT
present in your
Is Ramon No mention 0 The Ramon tree is present in all
Homegarden? No 0 homegardens of the interviewees
Yes 100 and their neighbors.
What kind of propagation tech- Sexual 89 The main propagation technique
nique do you use? Vegetative 11 used is seed. Attempts to propagate
Ramon vegetatively exist,
but there are no significant results
or reports.
Do you have traditional knowledge No mention 39 Traditional knowledge among
of Ramon from your parents or Lack of knowledge 11 these indigenous Maya is decreas-
other elders? Some knowledge 50 ing, especially for new
generations. Farmers seem not to
be interested in acquiring this
traditional knowledge.
Do you know the actual use of the No mention 17 The Maya interviewed strongly
tree?
Ramon No 0 agree that the use of Ramon as a
Yes 83 forage tree is its best use,
particularly during the dry season.
It is sole as well as used by the
owner. It was also used as
food by older genera tions
when maize was scarce, and
to increase milk production in
women and animals. It has several
other medicinal uses.
Do you give any kind of care to the No mention 11 Most of the interviewees agreed
tree?
Ramon No 17 that Ramon seedlings should
Yes 72 be watered or planted in the
rainy season. Once the plant is
established, irrigation is stopped.
What kind of technique No mention 61 Most of the indigenous people in-
do you use for Sexual 6 terviewed do not use special tech-
propagating other species Vegetative 11 niques of propagation for the
found in the homegarden? Both 22 species found in their homegar-
dens. Some propagate by
stakes or cuttings for certain
species, or they plant grafted
plants, but the grafts are not done
by them.
Do you have animals in No mention 17 Most of the interviewees have ani-
your homegarden that eat No 5 mals to feed and the Ramon tree
forage?
Ramon Yes 78 is used particularly for this
purpose.
What type of soil gives No mention 50 These Maya hold the concept that
higher productivity of Kancab (red soil) 17 the Ramon tree is a strong plant
and other species
Ramon Boxluum or Pusluum that can grow in any kind of
found in your (black soil) 6 soil condition. However, the red
homegarden? Tzekel (stony soil) 5 and black soils have the highest
1 and 2 (red and black soil) 5 likelihood of giving greater
Other 17 but
production, not only for Ramon
also for all the species of the
homegarden and the milpa.
32

Table 1. (Continued)
QUESTION ANSWERS PROVIDED DISTRIBUTION SUMMARY
PERCENT
They also agreed that Ramon can
grow even in stony soils.
The interviewees regarded
the red and black soils as
having good water holding
capacity for planting.
Is Ramon important No mention 44 Many Maya agreed that Ramon is
economically for your No 17 helping to increase their family
family? Yes 39 income.
Miscellaneous: What No mention 44 The Maya seem to know some
other information can you Botany of Ramon 17 botany of the Ramon tree.
mention about the Ramon Seed dispersal 0 All agreed that roots grow very
tree? Other 0 deep, trying to find water from
1,2,3 (botany, seed dispersal and other) 22 the water table.
1,2 (botany and seed dispersal) 0 Roots also extend a great distance
1,3 (botany and other) 11 laterally, and in the first stage of
2,3 (seed dispersal and other) 6 plant growth, roots are very
sensitive to disturbance.
Seed dispersal is mainly by bats.
Ramon seems to have various
ecotypes as the Maya have
observed Ramon trees that differ in
characteristics like size and shape
of leaves, and color of seeds.
Many mentioned that Ramon
is not found in the continuous
forest, but there is no certainty
about the reason.
They agreed that Ramon is mainly
found in the gardens and
close to urban areas.

nitosols) and Pusluum (cumulic ustirendolls or rendzinas). The Maya also men-
tioned that when Ramon seedlings were planted in Kancab soils, they obtained
higher yields because this soil facilitates planting in rows in this stony landscape.
The interviewed Maya, including the farmer-students at Mani, also provided
some information on the culture and growth of Ramon, including proper spacings,
initial age for pruning, age at flowering, lack of pests, and the use of the tree flowers
for beekeeping (itself an ancient Mayan production method with the local stingless
bee species) (Table 2). Unfortunately, much of this traditional knowledge regarding
the uses and management of Ramon is being lost, especially among new genera-
tions.

Reproduction ecology

Pardo-Tejeda and Sanchez-Munoz (1980) reported from field observation in the
Mexican state of Veracruz that between 84 and 88% of Ramon seeds germinated in
the beginning of the rainy season, reaching heights of 11 to 17 cm and an average of
2 leaves during the first month. However, not all the seedlings survived. Our results
in a controlled environment show that temperature has a great effect on germination
 33

Table 2. Summary of Mayan student-farmer knowledge on the culture of Ramon
Seed Collection
Seeds are collected from under trees and the seed coat removed before planting.
Seeds can be stored for up to two months.
Planting
Seed is planted at 3 times the depth of the seed thickness.
Germination takes place in 815 d, but no more than 3 wk.
Farmers dip the seed in warm water just before planting.
Soil is watered for seed planting.
Seeds are only planted in the rainy season.
If seedlings are transplanted from nursery polyethylene bags, a 30 3 40 cm hole is made.
Initial height growth is fast until seedlings are 50 cm tall.
Management
Animals are not allowed near seedlings or seedlings are fenced.
Though the seedlings are drought resistant, seedlings are watered regularly and soil is checked for moisture.
Once is it established, it grows well and active management is stopped.
A spacing of 4 to 6 m is used to allow for root extension.
When the tree is not pruned, it flowers at an age of 4 to 5 yr.
Flowers can be used for beekeeping.

rate and speed. Figure 1 indicates that the optimum range of germination of Ramon
seed from our study area was between 278C and 388C. Germination success
decreased as temperatures fell above or below this range. Additionally, the time to
germination increased (Figure 2). Also during our research, it was noticed that low
storage temperatures (refrigeration) killed the seeds of Ramon and these seeds
would likely be classed as recalcitrant in their germination.
Though the optimum temperature range for germination falls between 27 8C and

seed to temperature.
Figure 1. Germination response of Ramon
34

germination response time for temperature treatments having succesful germination.

Figure 2. Ramon

38 8C, the best temperature in this experiment was 33 / 30 8C when considering the
speed of germination and subsequent growth. Seedlings at this temperature appeared
more vigorous and had a greener color. Even though seed cultured at 38 / 35 8C and
27 / 24 8C provided 100% germination, these seedlings had a weak appearance;
smaller in height and less vigorous in comparison to seedlings grown at 33 / 30 8C.
The 38 / 35 8C treatment gave plants with small leaves, likely in response to the high
temperature and water deficit. Seedlings grown at 27 / 24 8C were smaller with
slower growth. The other four temperature treatments had either poor or no
germination and the seed that germinated gave seedlings of lower quality.
Time to 100% germination of Ramon seed, under greenhouse conditions, was 12,
16, and 19 d, for day temperatures of 38, 33, and 27 8C, respectively (Figure 2). This
period falls within the reported range of 10 to 20 d for 82% germination success
under greenhouse conditions in Costa Rica with local conditions of 3912 mm annual
rainfall and an average annual temperature of 25.7 8C (Gonzalez 1993)
and Quiros
and a range of 10 to 24 d (maximum germination of 48%) under natural conditions
in the state of Quintana Roo in the Yucatan peninsula (Lezama Conrado and Morfin
Francisco 1992). Morikawa et al. (1994) found Ramon seed in Jamaica required
several weeks to germinate, regardless of storage time (from fresh seed to 6 m
storage). These seed were germinated in a shaded nursery environment with a
maximum of only 90% germination success after one month. The authors report that
the local average daily temperature is 24 to 28 8C, below the optimum range found
in our study. Additionally, seed was germinated under shade. Thus, air and soil
temperatures were likely further reduced.
According to the interviewed Maya, the period of Ramon seed production in the
Yucatan is between August and October, the end of the rainy season and the period
of hurricanes or storms from the Gulf of Mexico. Meteorological information was
 35

seedlings grown under varying levels

Table 3. Root, shoot, total biomass, and (standard errors) of Ramon
of water stress in two soil types
Soil Type Soil Root Weight Shoot Weight Root:Shoot Total
Water (g) (g) ratio Biomass
Potential (g)
Top Soi1 0.1 MPa 3.19 (0.25) 13.69 (0.94) 0.23:1 16.88 (1.11)
Top Soil 1.0 MPa 1.12 (0.08) 3.06 (0.32) 0.37:1 4.18 (0.34)
Potting Soil 0.1 MPa 2.47 (0.21) 9.40 (0.65) 0.26:1 11.88 (0.81)
Potting Soil 1.0 MPa 0.70 (0.10) 1.56 (0.26) 0.45:1 2.26 (0.34)

obtained from a weather station in the town of Sotuta, a town close to the study site
Escalante 1998). This information indicated that during the period
(Montanez
between August and October, the average temperature is between 27 and 32 8C.
Thus, intensive efforts to propagate Ramon in the Yucatan on a large scale during
the rainy season can be undertaken with little climatic control. However, the species
is native throughout the tropical regions of North America and northern South
America as well as the Caribbean, growing at altitudes of 0 to 1000 m (e.g., Standley
1922, Peters and Pardo-Tejeda 1982). And flowering and fruiting times vary from
January to November depending on region (Chavelas Polito and Devall 1988).
Analysis of seed proteins with electrophoresis techniques indicates ecotypic vari-
ation in Ramon from the Pacific to the Gulf of Mexico. This genetic variation is
expressed in seeds that vary in seed and root phenotype, likely as an adaptation to
water limitations as well as temperature differences (Lopez-Mata 1987). Thus,
ecotypes would likely provide seed adapted to local conditions and temperatures and
these would aid propagation of plants locally. Local trials might be needed to
identify optimum temperatures in greatly different regions. Any efforts to germinate
seed and grow seedlings in a greenhouse or typical sheltered nursery setting would
need to account for altered air temperatures.
Plants from savannahs, desert areas, and other dry regions can develop root
systems reaching maximum depths of up to 30 m (cited by Parker 1968). From rock
cuts and wells, the Maya recognize the importance of roots and the depth to which
their trees reach to obtain water from the water table. Though most now have
potable water brought in by pipe to the household, many still have hand-dug wells
that reach the water table, approximately 15 to 18 m. The Maya irrigate new
plantings and seedlings by hand or hose, and they do so until the new plants are
deemed established (Table 1, 2).
Our simulation of Maya irrigation effects shows that seedling growth is indeed
accelerated (Table 3). After 274 d, total biomass is on the order of four times
greater. Irrigation makes a significant difference (p 5 0.0001) in this early growth
phase. Such a strategy not only provides greater shoot growth, it provides greater
root growth, aiding the plants ability to reach the water table and become
independent of irrigation. Plant growth under the normal, xeric condition is limited,
with low water availability slowing plant root and shoot growth. These biomass
differences were reflected in plant size measurements (Table 4).
Sporadic photosynthesis measurements (data not reported) also showed that
36

seedlings grown under varying

Table 4. Morphological differences (and standard errors) among Ramon
levels of water stress in two soil types
Soil Type Soil Mean Shoot Mean Leaf Mean Leaf
Water Height Number Size
Potential (cm) (cm)
Top Soi1 0.1 MPa 90.4 (3.7) 33 (0.3) 11.3 (0.6)
Top Soil 1.0 MPa 54.0 (3.2) 24 (3.5) 10.9 (0.2)
Potting Soil 0.1 MPa 91.8 (10.4) 38 (0.0) 15.4 (0.8)
Potting Soil 1.0 MPa 54.8 (2.9) 16 (0.9) 13.3 (0.7)

photosynthetic rates in irrigated plants were three to four times greater than plants
under water stress. Root to shoot ratios increase significantly (p 5 0.0001) with
water stress as more roots are produced per unit of shoot (Table 3). Under different
irrigation rates under forest canopy, Blain and Kellman (1991) also found less
growth of Ramon seedlings with less water in Veracruz, Mexico, and some evidence
of greater allocation to root growth. Such knowledge is critical for large-scale
planting of Ramon as a plantation crop. Not only must seedlings be of a size to
survive transplanting, but root length at outplanting becomes a critical issue where
soil depth is limited and limestone outcrop dominates the surface landscape.

Conclusions

Ramon is an important species in tropical America for subsistence and small market
production. It serves as nutritional forage for animal production, firewood, a shade
tree, and a source of medicines. The tree species also has the potential for
commercial forage production. Traditional knowledge of its propagation, ecology,
and culture still exist in the indigenous Mayan community of the Yucatan peninsula.
With some concern that this knowledge is fading, what remains should be extracted
if the species is to be grown on a commercial scale. The Yucatan is a tough
environment for plant production. Indeed, it is what constrained ancient Maya
populations and perhaps led to their decline in the region. But this knowledge is
useful in propagating and growing the species in this environment. Adaptation of
traditional production systems like raised platforms, Kaanche (Vargas Riveros
1983), can help to provide large numbers of seedlings in a more controlled
environment. Sheltering of these raised beds might also aid in controlling water use,
but temperatures and airflow would have to be monitored to maintain levels below
killing temperatures for seed and seedlings. In our study, temperatures above 44 8C
resulted in no germination. Too much shade may also reduce temperatures to levels
below optimum for germination, and our results showed that temperatures below 21
8C gave no germination. Irrigation appears critical in the early growth of the plant,
whether produced in a nursery setting or in situ. However, irrigation must continue
even after transplanting or outplanting to ensure that root systems become estab-
lished and perhaps reach the water table before irrigation is ceased.
While we expected increased growth of Ramon given irrigation, Maya manage-
 37

ment is seen to increase early growth four-fold, pointing the way for possible
commercial production. Would this magnitude of response pay for irrigation
installation and plantation management on a larger scale? This will vary by location
with labor and infrastructure costs. But in a region where plantings often involve the
movement of truckloads of soil, a tree capable of growing on rock may pay for itself.

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