Memposting Status

Inga edulis Mart.
Monograph
Esteban Velasquez Mariucci

Agriculture class 2021-2022
Colegio Bolivar
Esteban Velasquez: Inga edulis
Table of Contents:
Chapter 1: Introduction 3
Chapter 2: Ecology 4
2.1 Distribution 4
Figure 1: 4
Figure 2: 5
2.1.1 Affinity and Origin: 5
Figure 3: 6
2.1.2 Present Distribution 7
Figure 4: 8
2.2 Environmental Factors Affecting Distribution 8
2.2.1 Elevation and Climate 8
Figure 5: 9
2.2.2 Geology and Soils 9
2.3 Vegetation Components 9
2.3.1 Soil Environment and Effects of Soils 9
2.3.2 Interaction of roots and relationship with animals 10
Figure 6: 10
Table 1: 10
Table 2: 12
Chapter 3: Biology 13
3.1 Chromosome Complement 13
3.2 Life Cycles and Phenology 13
Table 3: 14
3.2.1 Seedling Protection 15
3.2.2 Flower Production 15
Figure 7: 15
3.2.3 Foliage and Growth 15
3.2.4 Seed Pods 16
3.3 Productivity and Biology 17
3.3.1 Pollen 17
Figure 8: 17
3.3.2 Sexuality and Reproduction 17
3.3.3 Pollination and potential pollinators 18
3.3.4 Anthesis 18
Figure 9: 18
1
3.3.5 Fruit Development and Seed Set 18

3.3.6 Ovule Development 19
3.4 Ecophysiology 19
Chapter 4: Propagation and Management 20
4.1 Natural Regeneration 20
4.2 Nursery and Propagation 21
4.2.1 Propagation from Seed 21
Figure 10: 21
Table 4: 21
4.2.2 Storage 23
4.3 Planting 23
Figure 11: 23
4.4 Management 23
4.4.1 Tending 24
4.4.2 Fruiting 25
Chapter 5: Products and Marketing 25
5.1 Imports and Exports: 25
5.2 Marketing: 26
Figure 12: 26
5.3 Uses: 26
5.3.1 Agroforestry Uses: 26
Figure 13: 27
5.3.2 Medical Uses: 27
5.3.3 Cachiri 28
Figure 14: 28
Bibliography: 28
Chapter 2: 28
Chapter 3: 32
Chapter 4: 34
Chapter 5: 37
2
Chapter 1: Introduction
Inga edulis Mart., also known as the Ice Cream Bean plant, is a peculiar yet historically
important leguminous tree, native to the Americas. It has accompanied ancient civilizations such
as the Inca and Aztec empires, providing food and wealth to their communities. In the modern
era, this exotic tree grows besides most coffee, cacao, and tea plantations within its area of
natural growth, providing shade, mulch, and nitrogen-rich soils. Its fruit is very sweet with
beautiful, soft white pulp and an exquisite vanilla flavor. Its seeds have a rapid germination
process, often growing even before the fruit touches the ground. That's why it has such a short
shelf life, meaning limited exportation. The tree can grow very tall, in various climates and
heights. It is known to be a forest regrow and a versatile tree since it prefers rough and acidic soil
conditions to establish itself.
Throughout this document, you will find four chapters diving deep into this wonderful
specimen. The first one talks about its origins, ideal environmental factors, and distribution of
Inga edulis. The second chapter talks about the biology and sexuality components of the tree.
The third chapter will contain information about the natural and human propagation of the tree,
as well as its management. The final chapter will show what uses and resources the tree can
provide to humankind, ones we sure can be grateful for!
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Chapter 2: Ecology
2.1 Distribution
Inga edulis is a South American native species with a natural range from Colombia to
Brazil.
Figure 1:
Map of Inga edulis’ native range (Orwa et al.2009).
The plant shares history with countless civilizations (ASALE & RAE, 2014). ASALE &
RAE (2014) stated that its historic distribution can be traced down to the Inca Empire. The name
Inga originates from the word Inka from the Quechua language, which references the Inca
Monarch. They played an important role in its current distribution by spreading the tree around
their empire and reaching, from the Andes mountains and Amazon river to Central America and
the Transverse Volcanic Mountain range in Mexico (ASALE & RAE, 2014) The Aztec empire
also knew of this plant's existence. They called it by the word jinicuil, which means distorted
foot in the Nahuatl tongue. This gave the scientific name to the other species of Inga called Inga
jinicuil (ASALE & RAE, 2014). In the modern era, this tree is commonly used as a shade
provider for crops, lumber, and food in the tropic.
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Figure 2:
Inga edulis leaves’ photograph from Colombia (Eden, 1977)
2.1.1 Affinity and Origin:
As stated previously, Inga edulis is a South American species. It originates from the
subtropical and tropical lands of South America and Mesoamerica. It first originated from the
Amazon, specifically the north of Brazil. Thanks to the help of cultivation, its high survival rate,
and effective reproduction spread to Colombia, Ecuador, and Peru which. Their shared
Amazonian region acted as a highway to its distribution (Lawrence, 1993). Civilizations, like the
Inca Empire, then spread it all over the tropical and subtropical regions around the Americas. It
is common to find these trees in subtropical wetlands, mountains, and plains. These trees can
outcompete most trees, the reason being they germinate quickly. This is why they can be found
in areas that are very favorable to plant life like near rivers, path trails in secondary forests
during formation, wooded swamps, and ravines (Duke, 1998).
5
Figure 3:
Inga edulis Tree Cultivation in Peru (Tularosa Flowers and Farm, 2009).
Domain: Eukaryota
Kingdom: Plantae
Phylum: Spermatophyta
Subphylum: Angiospermae
Class: Dicotyledonae
Order: Fabales
Family: Fabaceae
Subfamily: Mimosoideae
Genus: Inga
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Species: Inga edulis
2.1.2 Present Distribution
The species has been manually introduced to most tropical countries' ecosystems in South
America and Central America, ones like Panama, Venezuela, the Guianas, and Costa Rica. They
have also been grown in distant countries that maintain such climates, such as Madagascar, the
United States (Hawai’i and Florida), Australia, Uganda, New Zealand, and Indonesia (Velosa,
2021). Since the plant is only cultivated, semi-domesticated, and propagated in places of origin,
it gives it a limited distribution over the entire world. Not many uses have been found in foreign
lands.
Figure 4:
World Map of Recent Observations of Inga edulis (Velosa, 2021).
2.2 Environmental Factors Affecting Distribution
2.2.1 Elevation and Climate
Inga edulis is one of the hardiest trees that grow in the Americas. It grows in subtropical
climates that experience elevated rains and high light intensities. It can grow in regions with an
elevation up to 2200 m above sea level where there is no frost (Duke, 1998). The tree prefers
temperatures ranging from 23° to 30° but it can grow in temperatures ranging from 40° to -4°
celsius once it is established. It has a general USDA hardiness level of 9-12 (Malaga University,
2015). It needs to receive a minimum of 1200 mm of annual rainfall and can tolerate 650 mm of
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rainfall up to 4000 mm. The tree has been seen surviving plane floods that stay waterlogged for
up to three months. It can also tolerate and survive between 4 and 6 months of the dry season
(Fern, 2017).
Figure 5:
Inga edulis plantation in the mountains of Tolima, Colombia (Trapp, 2017).
2.2.2 Geology and Soils
Another aspect of its hardiness is its ability to grow on medium to high levels of acidic
soils and its capability to grow in soils low in nitrogen. The tree prefers a soil pH between 5 and
6.5 but can tolerate PHS from 4 to 8 (Fern, 2017). When it comes to chemical compounds, It
likes soils high in ammonium, since they increase the tree’s nodulation, which is important for
the tree’s nitrogen fixation process (Wang et al., 2018). Nitrates on the other hand reduce the
chances of nodulation. Phosphate levels have been recorded being absorbed in large quantities by
Inga edulis showing a liking for phosphate-rich soils (Fisher, 1995). It normally grows in soils
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with large amounts of organic matter. Its high versatility and adaptability give it the name forest
saver (SCIO, 2012).
2.3 Vegetation Components
2.3.1 Soil Environment and Effects of Soils
This legume plant is used to grow pretty much everywhere it can lay its roots in and get
mediocrely established. Being a legume means fixing its soil with nitrogen through symbiotic
processes (Masterclass, 2020). Not only does this benefit itself, but can benefit plants around it.
It transforms and provides better soils for already established plants and those to grow in soils
lacking nutrients and even on limestone (Fern, 2017). This plant is praised in its places of origin
for its positive effects on soil. Not only does it provide nitrogen, as stated before, but it also acts
as a natural land fallow accelerator. Its high biomass, organic matter disposal, and weed control
capabilities recuperate soil quality in a fast manner (Lojka et al., 2012).
2.3.2 Interaction of roots and relationship with animals
Living in the most biodiverse tropical rainforest in the world, Inga edulis is bound to
have complex and interesting relationships with other organisms. Inga edulis uses a common
symbiotic relationship found in legumes that establishes itself with a nitrogen-fixing bacteria
called Rhizobium, which is why it can survive in soils with low levels of nitrogen. The
relationship works first by Rhizobium taking nitrogen from the atmosphere and passing it on to
the plant’s roots in exchange for glucose provided by the plant (Artis, n.d.). It also leaves excess
hydrogen for other plants nearby to use. This process reduces the chances of soil corrosion by
creating healthy root systems. Animal-wise, it has a symbiotic relationship with pheidole ants,
which gather nectar from the tree’s flowers and nest on the tree, in exchange for patrolling the
tree against local herbivores (Wang et al., 2018).
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Figure 6:
Graph depicting the relationship between rhizobium and a plant’s system (Sato, 2014).
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Table 1:
List of Beneficial Organism Relations of Inga edulis.
Scientific Name Effect on I. Edulis Type of Organism Reference
Bradyrhizobium This rhizobium Bacteria Leblanc et al., 2005.

japonicum microsybiotically
creates nodulation
and hydrogen-fixates
roots.
Bradyrhizobium As a hybrid of Bacteria Leblanc et al., 2005.

liaoningense Bradyrhizobium
Japonicum, it also
nodulates and
provides hydrogen.
Pheidole These ants are highly Insect Melo et al., 2022.

megacephala aggressive, attacking
anything in their path.
Doing so they guard
the tree against local
herbivores
Table 2:
List of pests and diseases of Inga Edulis.
Scientific Name Affect on Inga Edulis Type of Pest Disease Reference
Lepidoptera larvae The larvae consume Insect Agroforestry

the leaf into complete Database 4.0,
defoliation leaving 2009.
only the stem.
Atta cephalotes The ant cuts and Insect Agroforestry

consumes the leaf Database 4.0,
leaving only the 2009.
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Scientific Name Affect on Inga Edulis Type of Pest Disease Reference
midrib and some

veins.
Hyphopolynema In an oval shape, the Fungus Pinho et al., 2010.

ingae fungus browns then
whiten the leaf until
holes appear in both
live and dead leaves.
Colletotrichum It consumes the fruit Fungus Gamboa, 2012.

by browning and
feeding on the rot.
Anastrepha distincta The organism Insect Rozendale, 2008.

consumes the freshly
established root
systems below the soil.
Septobasidium The fungus grows by Fungus Sistema Nacional

saccardoanum forming a cover over Argentino de
the mealybugs, which Vigilancia y
produce secretions that Monitoreo de
are used by the fungus Plagas, 2019.
for its growth.
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Chapter 3: Biology
3.1 Chromosome Complement
Inga edulis Mart. is a diploid fruiting tree species (2n = 2x = 26) native to the Amazon
Rainforest (Figueiredo et al., 2014).
3.2 Life Cycles and Phenology
The guaba tree, as referred to in Ecuador, reacts quickly to changes in climates. This
shows its high adaptive capabilities. The plant’s flowering goes hand in hand with ambient
conditions such as light, water, and nutrients (Garcia, 2008). As Winrock International (2015)
states, the plant requires hot, humid climates between 26°S and 10°N, and an elevation below
1600-1800 m above sea level. “It is most widespread in areas without a dry season (Andean
South America, western Brazil) or with a dry season of three to four months and minimum
annual rainfall of around 1200 mm.” (Winrock International, 2015).
Table 3:
Biologische Bundesanstalt, Bundessortenamt, and CHemical industry (BBCH) scale codification

for Inga edulis (Quijia-Quijia et al., 2020) translated from Spanish by Google Traductor.
Stage Description
5 Presence of flower buds, still surrounded by leaves.
5 21 First buds outside the leaves.
5 25 First buttons alone on the outside with closed leaves.
5 29 Several buttons on the bunches still remain closed.
6 Flowering.
6 20 First flowers uncovered.
6 21 Presence of 1 to 2 open flowers in the first cluster.
6 23 Presence of 3 open flowers in the clusters per plant.
6 25 Full bloom: bare flowers, 5 clusters per plant.
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Stage Description
6 27 Flowering declines.
6 29 The flowering phase ends.
7 Fruit development.
7 20 Initial pods reach initial length (“flat pod”).
7 23 30% of the pods reach the final length of the pod.
7 29 The vast majority of pods reach the long end.
3.2.1 Seedling Protection
A mutualism exists with certain species of ants and Inga edulis. The ants protect the fruit
along with the whole tree (Janzen & Martin, 1980). The pods are hard, especially in the corners,
to avoid premature openings (Lim, 2012). Water retention is very important for the Inga edulis’
reproduction. Seedlings lose their vitality after failing to retain 65% of their water content
(Barbedo & Bilia, 1998).
3.2.2 Flower Production
As Marín-Gómez et al. (2012) state, the flowering phase happens twice a year, in periods
of 4-6 months. The process of transformation of a tree’s bud to flower takes from 98-120 days
when they grow in size. The total expansion goes from 10.25 mm to 46.98 mm (Quijia-Quijia et
al., 2020). Flowers are described to be very fragrant despite their size. They are attached directly
to the stalk in groups of five growing in an axle (Lim, 2012). They are usually white and yellow
but can have a light pink color at early stages.
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Figure 7:
Photo of Inga edulis flowers’ axle growth (Abatia, 2012).
3.2.3 Foliage and Growth
It is a fast-growing tree, starting up as a light yellow sprout (Lojka, 2010). When

maturing, its leaves are dark green, with three to four pairs of leaflets per cluster and with
winged central leaf stalks. They generate moderate to dense shade. Plants in the genus Inga have
nectaries on their leaves, producing nectar during leaf development to attract beneficial insects
like parasitic wasps and ants (GardenOracle, 2021). After reaching maturity, fruiting, sprouting,
and foliage fall are constant. The intensity of some phenophases can be traced to maintain a
coincidence with the precipitation (Marín-Gómez et al., 2012).
3.2.4 Seed Pods
The fruits are ribbed cylindrical leguminous pods that can grow straight, curved, or
spirally twisted. They hang like pendants and grow up to one meter long. They produce 10-20
ovoid seeds that have a dark purple to black olive color. The plant’s seeds have been seen
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sprouting before the pods mature (Barbedo & Bilia, 1998). Pods are often seen being dispersed
by rivers thanks to air trapped in the pods. They still sprout under these conditions and their
genetic diversity may be greater (Russell et al., 1999).
3.3 Productivity and Biology
3.3.1 Pollen
This tree needs another individual’s pollen to correctly fruit (Lim, 2012). The flower has
a calyx tube with five fine, striate lobes, a corolla with five silky petals. They contain numerous
white stamens where pollen is found. The stamens are long and very dispersed to facilitate
pollination from other organisms and even from strong winds. Pollen and nectar are also
produced in extrafloral nectaries located in the leaves (GardenOracle, 2021).
Figure 8:
Example of Inga edulis’ pollen morphology (Barth, 2008).
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3.3.2 Sexuality and Reproduction
When a flower is fully opened and mature, pollinators will take pollen generated from the
stamen, the male part of the flower which measures between two to three millimeters, and
deposit it in the pollen tubes. Individual microscopic pieces of pollen go down the pistil, the
female part of the flower until they reach the ovary where fertilization ends. As you can see this
plant contains male and female organs (Barros et al., 2013).
3.3.3 Pollination and potential pollinators
Flowers of this type are typically pollinated at night by small mammals, birds, and
insects. Bats are usually the biggest pollinators since their furry faces get powdered with pollen
as they search for nectar. Recent research into Inga has shown that at least in some species
hummingbirds are the first pollinator after the flowers open, followed by bats as well as
hawkmoths (Growables, 2020).
3.3.4 Anthesis
This species coordinates its flowering process to make most individual flowers reach
maturity at the same time. This behavior generates a massive blossom to maximize the number
of pollinator visits (Marín-Gómez et al., 2012). This process is taken advantage of by the male
stamens, which release pollen in massive amounts, for multiple pollination processes to fertilize
the female organs.
Fructification is constant throughout the year, with the exception of the months of
December, January, and February when production is lowered. When the tree is fruit-bearing,
leaf loss is increased temporarily (Marín-Gómez et al., 2012). Mature fruit can reach up to a
meter long and three to five cm in diameter (Winrock International, 2015). Their shape can vary,
some individuals can be straight, while others are twisted or even looped. Each pod can contain
10-20 oval-shaped seeds embedded with soft, white pulp (Barbedo & Bilia, 1998).
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Figure 9:
Inga edulis’ fruit on tree (Abatia, 2012).
3.3.5 Fruit Development and Seed Set
Even though the plant produces over 100,000 flowers each season, the fruit set is only
about 1%. As the Rare Fruit Club (2009) states, this plant’s flowers are very fragrant and with
the complement of extrafloral nectaries bees and other insects are easily attracted, which are
thought to affect pollination. This leaves insects and birds to pollinate during the day, and small
nocturnal mammals, like bats, pollinate at night. This process maximizes the quality of fruit
producción, despite their low fertility rate.
3.3.6 Ovule Development
Authors Eduardo Barros, Antonio Webber, and Isabel Machado (2013), in their research
article they published called “Limitação de polinizadores”, describe the ovule development
process of Inga edulis. They state that after a pollinator introduces pollen into the plant, it takes
about eight hours for a fraction of it to reach the plant’s tube. Although the tree has a very
favorable bloom for fruiting, only 1% of the flowers get fertilized. Their studies show that the
blossom’s pollen tubes had no obstructions for pollination, incompatibility occurred after the
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penetration of the tubes into the micropyles of the eggs. This process has a 0.3% to 0.4% chance
of correctly developing an ovule.
3.4 Ecophysiology
Inga edulis reacts accordingly to the environment. This species shows better growth in
non-flooding fields. The rapid growth of this species is also reported on very acidic soils, low in
nutrients and high in aluminum toxicity in pastures located in the Amazon (Lojka, 2010).
Flowering commonly starts halfway through the dry seasons, meaning that fruiting is most
commonly processed during the beginning of the rainy seasons (Rare Fruit Club, 2009).
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Chapter 4: Propagation and Management
4.1 Natural Regeneration
This tree propagates from seed very easily. Once pods are open seeds don't have a long
time to survive because they lack moisture. They commonly sprout at the time the fruit matures
to be ready for propagation once the pods open.
Inga Edulis is considered to be a pioneer tree because of its ability to populate deserted
forest grounds. Its unique capabilities to survive in harsh conditions surpasses other competitors.
In volume 217 of the book Forest Ecology and Management (Park et al., 2005), Inga Edulis is
described to be dominant over large gaps of time in deforested lands by seed spreading. These
trees have been seen growing in coarse dirt and even on highly vegetated grounds. Guamas,
smaller than 1.5 meters tall and 10 centimeters in diameter at breast height, were more abundant
within a year than any other plant. Even in recent logging grounds, they excel in successful
reproduction and natural regeneration.
4.2 Nursery and Propagation
Farmers don't use nurseries often. Instead, they usually plant the seeds directly on fields.
At times when they do use nurseries, the general process is: separate seed by seed, cover them in
plastic sheets to maintain moisture, place them in semi-shaded areas and water them every two
days. Bare roots are equivalently capable of growing as soil-covered saplings (FACT Net, 1993).
4.2.1 Propagation from Seed
Seeds start to germinate inside the pods. After being expelled from the pods, seeds have
two weeks to find themselves with a humidity source before dying. Seeds better propagate with
their pulp taken out (Growables, Inc, 2020). This tree can only reproduce from seeds. Farmers
from South America deferienciate the seeds by color, the darker ones are called female while the
lighter ones are called males. They prefer female seeds, because of this they plant dark seeds
since they carry genes to produce more dark seeds. Inga edulis can only reproduce via seeds.
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Figure 10:
Seed shape and color variations, alongside a germinated individual (TROPICALBOTANY,

2013).
Table 4:
Implications and Pre-preparations for Inga edulis’ germination.
Implications/ Description Citation

Pre-preparations
Frost Tolerance Mature plants can tolerate up Fern, 2017.

to -2 degrees celsius. New
growth gets damaged at 0
degrees celsius.
Sun Exposure The plant prefers to position Fern, 2017.

itself in sunny areas but it has
been recorded to grow in
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Implications/ Description Citation

Pre-preparations
shade successfully.
Origin Its origins can be traced down Orwa et al.2009.

from the Andes Mountain
Range to the Amazonian
region.
Growth Habits Tropical legume trees that can Growables, Inc, 2020.
grow up to 40 m (130 ft.) in
native climates.
Water Needs It can survive 4-6 months of Growables, Inc, 2020.

drought and waterlogged for
2-3 months
Propagation The tree only propagated Growables, Inc, 2020.

through seed.
4.2.2 Storage
Barbedo and Bilia (1998) state that the shelf life of a seed is two weeks. Within that time
period, it needs to find moisture to keep functional. Conservationists have found multiple ways
of keeping the seed viable for extended periods of time using low-temperature drying, special
packages for storage, and chemical and fungicide treatments.
4.3 Planting
It is recommended that an area of one meter in diameter should be kept clear of any
living things around the tree for the first six months of growth to enable a correct rooting and
foliage environment. Inga edulis regrows fairly well after pruning. It should not be cut below
0.75 meters. It has been recorded that the tree has a better response to pruning if the height and
range of cuts are varied and a few branches are left untouched. Precise cuts should be made at
least three centimeters above a node from which the branch shoots will grow again. (Orwa et
al.2009.)
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Figure 11:
Manual grow bag planting of Inga edulis (TROPICALBOTANY, 2013).
4.4 Management
Managing this tree is very easy. Orwa et al. (2009) states that its adaptations make it
highly resistant to leaf-cutting ants, which tend to be the highest threat to trees in its native area.
The real threat is Lepidoptera larvae, which have been recorded to completely defoliate the tree.
While fruit flies are not considered to be a threat, it has been observed that they often can
damage the seed testa, especially in the seed’s late maturity stage. Some slight damage from
fungal attack, specifically Rhizoctonia, towards the seedlings has been noticed.
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4.4.1 Tending
This tree is well known for its survival capabilities. Especially for growing pretty much
anywhere in tropical and subtropical areas of the world. This means that growth variations are as
diverse as the places it grows. The plant does tend to grow better in certain conditions. Its growth
is better in fields that don't experience flooding. The rapid growth of this species is also reported
on highly acidic soil, which is low in nutrients and high in aluminum that is normally found in
abandoned pastures in the Amazonian region (Lojka et al., 2012).
4.4.2 Fruiting
The fruit is described as “... ribbed, cylindrical pods, straight or often spirally twisted,”
that can grow up to a meter long. “They contain fleshy green seeds in a sweet, white, cottony
pulp. They are produced during the wet season, and monkeys and birds eat the sweet pulp and
scatter the soft seeds.” (Orwa et al., 2009).
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Chapter 5: Products and Marketing
5.1 Imports and Exports:
Ice cream beans are not a common product to import or export due to their short shelf
life, limited availability, and conservation regulations. Most of the production and consumption
stays within the country it was grown.
Colombia signed a treaty with the European Union in Brussels, on June 26 of 2012, that
permitted the importation of the fruit, along with other produce, into Europe. This treaty is of
great importance since demand exceeds supply, and 56% of all fruits consumed are imported
(Osorio, 2021). Not much is known about the importation or exportation process.
5.2 Marketing:
Inga edulis is sold to distributors in loads tied together or in bags. Its hard pod shell gives
it great protection. It is most often sold in local markets, not far from where they were grown.
They can also be found internationally under exotic fruit markets. The fruit is highly praised for
its taste similar to ice cream, and although uncommonly done, their seeds when boiled are edible.
Its seed can be bought over the internet, which is commonly imported to similar climates to
where it is found, so consumers can grow their own tree (Miami Fruit, 2021).
Figure 12:
Display of Inga edulis in a market (Bolivar, 2017).
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5.3 Uses:
5.3.1 Agroforestry Uses:
This tree species has incredible growth rates and abilities to improve soil quality which is
why some farmers use it as an alley crop. It eliminates the necessity of slash and burns
techniques in cultivation.
The most famous use for Guama is shading coffee, cocoa, and tea plantations. Its
generous height, deep roots, and nitrogen fixations give it great value in the cultivation
ecosystem. Most importantly, the tree can hold to its leaves during a dry season, giving shade
when most needed. Fallen leaves make for great mulch and fresh cuttings for animal feed.
Farmers often also use leaves and wood for fire-making (Lojka et al., 2010).
Figure 13:
Coffee plantations under the shade of giant Guama trees (Cafe La Meca, 2021).
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5.3.2 Medical Uses:
The tree's leaves have a long history with the South American native tribe’s medicine.
The process normally follows a pattern which is a boil and direction of the leaves, along with
roots or bark, which are left to cool and then digested. These concoctions are said to treat
diarrhea, arthritis, and rheumatism. Lightly boiled leaves are ingested to relieve coughing or
sores, possibly healing herpes infections. Studies show that these leaves contain polyphenols and
antioxidants (Silva et al., 2007).
5.3.3 Cachiri
In the northeast and south of Colombia, an alcoholic beverage is made from this plant’s
arils. It is called Cachiri. A festival under the same name takes place where women chew and
spit the arils which are left to ferment (Duke., 1983). The beverage has a shelf life of about four
days, but with modern refrigeration, it can last over two weeks.
Figure 14:
Cachiri being ingested (Flavors of Brazil, 2012).
27
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Eden, H. J. (1977). HerbWeb—Image Page.
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Fern, K. (2017, October 4). Inga edulis Ice Cream Bean, Monkey Tamarind PFAF Plant
Database. https://pfaf.org/User/Plant.aspx?LatinName=Inga+edulis
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Fisher, R. F. (1995). Amelioration of Degraded Rain Forest Soils by Plantations of Native
Trees. Soil Science Society of America Journal, 59(2), 544–549.
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Lawrence, A. (1993, September). Winrock International - Inga edulis: A tree for acid soils
in the humid tropics.
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in-the-humid-tropics/
Leblanc, H., McGraw, R., Nygren, P., & Le Roux, C. (2005). Neotropical Legume Tree Inga
edulis Forms N 2 -fixing Symbiosis with Fast-growing Bradyrhizobium Strains. Plant
and Soil - PLANT SOIL, 275, 123–133. https://doi.org/10.1007/s11104-005-0808-8
Lojka, B., Preininger, D., Van Damme, P., Rollo, A., & Banout, J. (2012). Use of the
amazonian tree species INGA edulis for soil regeneration and weed control. Journal of
Tropical Forest Science, 24, 89–101.
Malaga University. (2015, June 29). Inga edulis—JB-34-06. Jardín Botánico.
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Masterclass. (2020, November 8). How Nitrogen-Fixing Plants Enrich the Soil—2021.
MasterClass.
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Melo, T. S., Koch, E. B. A., Andrade, A. R. S., Travassos, M. L. O., Peres, M. C. L., &
Delabie, J. H. C. (2022). Ants (Hymenoptera: Formicidae) in different green areas in
the metropolitan region of Salvador, Bahia state, Brazil. Brazilian Journal of Biology,
82, e236269. https://doi.org/10.1590/1519-6984.236269
29
Miller, T. (2017). No. 78 Thank you very much! And Inga edulis.
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Pinho, D., Firmino, A., & Pereira, O. (2010). Hyphopolynema ingae sp. Nov., associated
with leaf-spot disease on Inga edulis in Brazil. Mycotaxon -Ithaca Ny-, 114, 55–59.
https://doi.org/10.5248/114.55
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Sato, T. (2014, January 29). Effects of Rhizobium Inoculation on Nitrogen Fixation and
Growth of Leguminous Green Manure Crop Hairy Vetch (Vicia villosa Roth) |
IntechOpen. https://www.intechopen.com/chapters/46074
Sistema Nacional Argentino de Vigilancia y Monitoreo de Plagas. (2019). Septobasidium
saccardoanum | Sistema Nacional de Vigilancia y Monitoreo de plagas.
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Trapp, A. E. B. (2017, noviembre 6). Maintenance of Inga Edulis Reforestation: Video Log
#2. Steemit.
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og-2
Tularosa Flowers and Farm. (2009, marzo). IMG_2426—Inga edulis [Guaba] | Pucallpa,
Peru. March 2009.… | Flickr.
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Velosa, L. (2021). Guama (Inga Edulis). Tropicos.Org. Missouri Botanical Garden, 4.
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Wang, Q., Liu, J., & Zhu, H. (2018). Genetic and Molecular Mechanisms Underlying
Symbiotic Specificity in Legume-Rhizobium Interactions. Frontiers in Plant Science, 9,
313. https://doi.org/10.3389/fpls.2018.00313
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Chapter 3:
Abatia. (2012). Photos of the fruits of Colombia, Inga edulis. Meet Latin Women for
Marriage, an Exotic Latin Bride.
https://www.latin-wife.com/blog/colombia/inga-edulis/
Barbedo, C. J., & Bilia, D. a. C. (1998). Evolution of research on recalcitrant seeds. Scientia
Agricola, 55, 121–125. https://doi.org/10.1590/S0103-90161998000500022
Barros, E. C. de O., Webber, A. C., & Machado, I. C. (2013). Limitação de polinizadores e
mecanismo de autoincompatibilidade de ação tardia como causas da baixa formação de
frutos em duas espécies simpátricas de Inga (Fabaceae—Mimosoideae) na Amazônia
Central. Rodriguésia, 64(1), 37–47.
https://doi.org/10.1590/S2175-78602013000100005
Barth, O. M. (2008). Fig. 3. Examples of pollen morphology. 1 -Inga edulis (400x). 2-...
ResearchGate.
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x-2-Albertina-brasiliensis-1000x_fig3_235365322
Figueiredo, M. F., Bruno, R. L. A., Barros e Silva, A. E., Nascimento, S., Oliveira, I. G., &
Felix, L. P. (2014). Intraspecific and interspecific polyploidy of Brazilian species of the
genus Inga (Leguminosae: Mimosoideae). Genetics and Molecular Research: GMR,
13(2), 3395–3403. https://doi.org/10.4238/2014.April.29.18
GardenOracle. (2021, February). Growing Ice Cream Bean / Pacay: Inga feuilleei.
https://gardenoracle.com/images/inga-feuilleei.html
Growables. (2020, November 3). Ice Cream Bean, Inga edulis.
https://www.growables.org/information/TropicalFruit/IceCreamBean.htm
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Legume | Definition & Examples | Britannica. (1980). Retrieved January 25, 2022, from
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Lim, T. K. (2012). Inga edulis. In T. K. Lim (Ed.), Edible Medicinal And Non-Medicinal
Plants: Volume 2, Fruits (pp. 715–719). Springer Netherlands.
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Lojka.pdf. (2010). Retrieved February 2, 2022, from
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Marín-Gómez, O. H., Castaño-González, A. F., & Gómez-Marín, G. D. (2012).
FENOLOGÍA DEL GUAMO Inga edulis (FABALES: MIMOSOIDEAE) EN DOS
AGROECOSISTEMAS DEL QUINDÍO, COLOMBIA. Revista de Investigaciones
Universidad del Quindío, 23(2), 127–133. https://doi.org/10.33975/riuq.vol23n2.412
Quijia-Quijia, M., Castillo-Torres, S., Vasquez-Castillo, W., & Racines-Oliva, M. (2020).
Fenología floral de la guaba (Inga edulis) en un valle interandino del Ecuador. Enfoque
UTE, 11(3), 25–34.
Rare Fruit Club. (2009). Inga, also called Ice Cream Bean [Organization]. Rare Fruit Club.
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The Defenses of Legumes against Herbivores.pdf. (n.d.). Retrieved February 1, 2022, from
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0of%20Legumes%20against%20Herbivores.pdf?sequence=1&isAllowed=y
TyC_2008_22_03.pdf. (2008). Retrieved January 21, 2022, from
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Winrock International. (2015). Winrock International - Inga edulis: A tree for acid soils in
the humid tropics.
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in-the-humid-tropics/
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Chapter 4:
Orwa C, A Mutua, Kindt R , Jamnadass R, S Anthony. 2009 Agroforestree Database:a tree
reference and selection guide version 4.0
http://www.worldagroforestry.org/sites/treedbs/treedatabases.asp
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Scientia Agricola, 55, 121–125.
FACT Net. (1993, September). Winrock International - Inga edulis: A tree for acid soils in
the humid tropics.
https://winrock.org/factnet-a-lasting-impact/fact-sheets/inga-edulis-a-tree-for-acid-soils
-in-the-humid-tropics/
Fern, K. (2017, October 4). Inga edulis Ice Cream Bean, Monkey Tamarind PFAF Plant
Database. https://pfaf.org/user/Plant.aspx?LatinName=Inga+edulis
Growables, Inc. (2020, November 3). Ice Cream Bean, Inga edulis.
https://www.growables.org/information/TropicalFruit/IceCreamBean.htm
Lojka, B., Preininger, D., Van Damme, P., Rollo, A., & Banout, J. (2012). Use of the
amazonian tree species INGA edulis for soil regeneration and weed control. Journal of
Tropical Forest Science, 24, 89–101.
Park, A., Joaquin Justiniano, M., & Fredericksen, T. S. (2005). Natural regeneration and
environmental relationships of tree species in logging gaps in a Bolivian tropical forest.
Forest Ecology and Management, 217(2), 147–157.
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Times.
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1-story.html
TROPICALBOTANY. (2013, December 29). How to prepare Inga seeds for sowing. Tropical
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Bolivar, F. (2017). Venta de Plantas de Guama, Guamo. Facebook.
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adearbolesfrutalesensanfelix%2Fposts%2F805028329556818%2F
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Café la Meca.
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38

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Inga edulis Mart.

Esteban Velasquez Mariucci

3.3.5 Fruit Development and Seed Set 18

Map of Inga edulis’ native range (Orwa et al.2009).

Inga edulis leaves’ photograph from Colombia (Eden, 1977)

2.1.1 Affinity and Origin:

Species: Inga edulis

2.1.2 Present Distribution

2.2 Environmental Factors Affecting Distribution

2.2.1 Elevation and Climate

Inga edulis plantation in the mountains of Tolima, Colombia (Trapp, 2017).

2.2.2 Geology and Soils

2.3 Vegetation Components

2.3.1 Soil Environment and Effects of Soils

2.3.2 Interaction of roots and relationship with animals

List of Beneficial Organism Relations of Inga edulis.

Scientific Name Effect on I. Edulis Type of Organism Reference

Bradyrhizobium This rhizobium Bacteria Leblanc et al., 2005.

Bradyrhizobium As a hybrid of Bacteria Leblanc et al., 2005.

Pheidole These ants are highly Insect Melo et al., 2022.

List of pests and diseases of Inga Edulis.

Scientific Name Affect on Inga Edulis Type of Pest Disease Reference

Lepidoptera larvae The larvae consume Insect Agroforestry

Atta cephalotes The ant cuts and Insect Agroforestry

Scientific Name Affect on Inga Edulis Type of Pest Disease Reference

midrib and some

Hyphopolynema In an oval shape, the Fungus Pinho et al., 2010.

Colletotrichum It consumes the fruit Fungus Gamboa, 2012.

Anastrepha distincta The organism Insect Rozendale, 2008.

Septobasidium The fungus grows by Fungus Sistema Nacional

3.1 Chromosome Complement

3.2 Life Cycles and Phenology

Biologische Bundesanstalt, Bundessortenamt, and CHemical industry (BBCH) scale codification

5 Presence of flower buds, still surrounded by leaves.

5 21 First buds outside the leaves.

5 25 First buttons alone on the outside with closed leaves.

5 29 Several buttons on the bunches still remain closed.

6 20 First flowers uncovered.

6 21 Presence of 1 to 2 open flowers in the first cluster.

6 23 Presence of 3 open flowers in the clusters per plant.

6 25 Full bloom: bare flowers, 5 clusters per plant.

6 29 The flowering phase ends.

7 20 Initial pods reach initial length (“flat pod”).

7 23 30% of the pods reach the final length of the pod.

7 25 50% of the pods reach the final length of the pod.

7 27 70% of the pods reach the final length of the pod.

7 29 The vast majority of pods reach the long end.

3.2.1 Seedling Protection

3.2.2 Flower Production

Photo of Inga edulis flowers’ axle growth (Abatia, 2012).

3.2.3 Foliage and Growth

It is a fast-growing tree, starting up as a light yellow sprout (Lojka, 2010). When

3.2.4 Seed Pods

3.3 Productivity and Biology

Example of Inga edulis’ pollen morphology (Barth, 2008).

3.3.2 Sexuality and Reproduction

3.3.3 Pollination and potential pollinators

Inga edulis’ fruit on tree (Abatia, 2012).

3.3.5 Fruit Development and Seed Set

3.3.6 Ovule Development

Chapter 4: Propagation and Management

4.1 Natural Regeneration