Design Complex - Systems Agrobiodiversity

Environment, Development and Sustainability
https://doi.org/10.1007/s10668-023-03002-9
Design of complex agroecosystems: traditional and formal

knowledge to conserve agrobiodiversity in the Santa Marta
Mountains, Veracruz, México
Carlos H. Ávila‑Bello1 · Ángel Héctor Hernández‑Romero1 ·

Dinora Vázquez‑Luna1 · Daniel Alejandro Lara‑Rodríguez1 ·
Adriana Martínez‑Jerónimo2 · Bitia N. Meneses‑García2 , et al. [full author details at
the end of the article]
Received: 11 May 2022 / Accepted: 1 February 2023

© The Author(s), under exclusive licence to Springer Nature B.V. 2023
Abstract
In the context of globalization, two major human challenges are global warming and the
loss of biological and cultural diversity. Solving the latter problem can help reduce green-
house gases, diversify production, and maintain agrobiodiversity. The objective was to pro-
pose a design of complex agroecosystems based on indigenous and scientific knowledge.
In 66 agroecosystems from seven ejidos (communal lands used for agriculture), the tree
species were determined in 400 square meter plots, considering three layers: high (up to
35 m), middle (up to 25 m), and low (< 17.5 m), estimating carbon sequestration using
allometric equations. Three participatory workshops were carried out with 36 peasants to
corroborate the species observed in the field and create lists of attributes (foliage produc-
tion, cover, growth speed, and flowering and fruiting periods). With these attributes, as
well as carbon sequestration capacity, melliferous and nectar potential, a proposal for an
agroecosystem design was developed. The species that sequestered the largest amount of
carbon was: Pouteria sapota, Manilkara sapota, Brosimum alicastrum, and Calophyllum
brasiliense. The highest valued species were: Spondias mombin, Citrus limon, and Psidium
guajava, Guazuma ulmifolia, Trema micrantha, Inga spp., Bursera simaruba, Mangifera
indica, Pouteria sapota, and Tabebuia rosea; in the shrub and understory layer Musa para-
disiaca, Coffea arabica, Chamaedorea tepejilote, and Chamaedorea spp. The combination
of indigenous and western scientific knowledge allows the structuring of complex agroeco-
logical matrices to diversify coffee agroecosystems. This can be the basis for reducing pov-
erty, conserving, or restoring different trophic relationships, and reducing global warming.
Section 1 presents the literature reviewed. Section 2 presents a description of the study area
with the main biological and social characteristics, also, the main methods used to charac-
terize coffee agroecosystems, the estimation of carbon sequestration and valuation of tree
species by peasants and the attributes used for the proposal of design of agroecosystems.
Sections 3 and 4 present the discussion of results and finally in Sect. 5, the conclusions.
Keywords Biocultural diversity · Complex systems · Indigenous seeds · Sustainability
13
Vol.:(0123456789)
Content courtesy of Springer Nature, terms of use apply. Rights reserved.
C. H. Ávila‑Bello et al.
1 Introduction
The study, planning, and management of nature to produce healthy food is a complex task,
and thus, the paradigms of complex systems and agroecological matrices can be fundamen-
tal for the design of sustainable agroecosystems, taking into account the carrying capacity
of the environment, but also the knowledge and needs of native communities (Soley & Per-
fecto, 2021). In countries like Mexico, these indigenous groups have an extensive knowl-
edge of the environment and the agrobiodiversity that they have domesticated, conserved,
and improved for centuries (Llamas-Guzmán et al., 2022). The complex systems paradigm
allows, based on interdisciplinary work teams, studying and understanding the different
elements that interact in a given environment, considering their limits and reciprocal rela-
tionships (Capra & Sempau, 1998; García, 2006). A complex agroecosystem is a dynamic
set of ecological networks and cultural practices carried out by human beings based on the
influence of economic, social and political factors; these subsystems and their elements
interact through exchanges of energy and matter, which can lead to co-evolutionary pro-
cesses of domestication that preserve fundamental ecological events for the survival of the
planet, according to Krakauer (2019), many of the current challenges and errors of this
century derive from a poor understanding of what a complex system is. Complementarily,
the agroecological matrix approach allows determining the structural characteristics of bio-
logical diversity (alpha, beta, and gamma) in a set of spaces covered by patches of natural
or cultivated vegetation managed by humans and which, depending on their distance and
complexity, can facilitate gene flow between populations and the preservation of different
groups of species, trophic chains, and biogeochemical cycles that allow a dynamic equilib-
rium in the ecosystem (Vandermeer & Perfecto, 2007). To transcend to the socioeconomic
aspect, the design of agroecological matrices must be based on local knowledge (through
the knowledge dialogue) and meet the need for the production of healthy food and other
goods for the population, and must also meet regional, national, and global needs intercon-
nected in different ways, and where appropriate, must allow generating surpluses to cover
other population needs (Polanco-Echeverry & Rios-Ososrio, 2015). The biodiversity of a
region and the associated trophic and socioeconomic networks form a complex biocultural
web, based on which it is possible to achieve healthy food production, self-sustenance,
and food autonomy. It can also be the basis of a production strategy in the face of climate
change, water crises, soil deterioration, migration, economic crises, and pandemics, which
would allow reducing carbon emissions, deforestation, and the loss of soils and microor-
ganisms, which are some of the biggest challenges in tropical countries (Gibbs et al., 2007).
Agroecosystems have been gradually recognized as important environments for carbon
(C) sequestration and storage. These include traditional coffee and cocoa agroecosystems
with diversified shade (Segura et al., 2006). According to Dixon (1995), the potential for
C sequestration in this type of systems ranges between 12 and 220 tons of carbon per hec-
tare (ton C ha−1), including soil carbon, although the potential for storage is even higher
in the humid tropics. These agroecosystems represent an option, not only because of their
ecological value, but also because they are an economic alternative source that can help
reduce global warming through the payment for C sequestration, a strategy that must be
considered, designed, and implemented in the short term (Olabi et al., 2022). Similarly,
the biological diversity of agroecological matrices that compose the biocultural landscape
can be the basis for the design of complex agroecosystems (Perfecto & Vandermeer, 2015),
with the aim of having matrices that satisfy not only fundamental ecological aspects but
also social and economic ones. Even though ecological reserves play an important role in
13
Design of complex agroecosystems: traditional and formal…
the protection of a wide range of species, the function of patches interconnected by agro-
ecological matrices of different quality and complexity is crucial (Lindenmayer & Frank-
lin, 2002; Perfecto & Vandermeer, 2015). Furthermore, the complex agroecosystems pre-
served with traditional indigenous knowledge contribute to support ecosystem functions
in every of four categories: regulation (e.g., regulation of CO2 sequestration and nutrient
cycling), production (food, firewood, medicines, genetic resources, etc.), habitat (for refuge
and reproduction of wild biota, contributing for conservation and evolutionary processes)
and cultural or information functions (e.g., for recreational, aesthetical, artistical, spiritual,
historical, educational and scientific research functions) (Chiesura & de Groot, 2003). This
paper aims to point out some aspects of all these functions, emphasizing on the contribu-
tion of traditional knowledge to maintain functional agroecosystems.
One of the most important problems faced in many coffee-producing regions is chang-
ing to agroecosystems without shade or with shade consisting of one or a few tree species,
largely due to the pressure of soluble coffee industries which seek the production of low-
quality coffee, as well as the impact caused by coffee rust (Hemileia vastratix). Eliminating
shade trees or simplifying shade can be a serious mistake, since C sequestration can be
drastically reduced, the soil is exposed to erosion and loss of microorganisms, less water
is retained, and spaces for different species that can have important functions (pollinators,
predators, antagonists, among others) are eliminated, as well as for seed dispersers. Shade
trees in coffee agroecosystems have been shown to serve as refuges for insects such as Azya
luteipes and A. orbigera, which at different stages of their biological cycle attack scales
(Coccus spp.), which are parasites of coffee (Salinas-Castro & Ávila-Bello, 2016). There is
also enough evidence to conclude that part of the natural control of H. vastatrix is related
to a complex network of interactions between ants of the genus Azteca, Azya spp., and
some fungal diseases in scales caused by Lecanicillum lecanii (Vandermeer et al., 2010);
this fungus is a very important antagonist of coffee rust and its spores are found in leaf lit-
ter deposited in the soil by different shade trees, and in turn, the trees are a refuge for ants
of the genus Azteca, as well as for Azya spp. In addition to the above, trees in shade cof-
fee agroecosystems are an important source of flowers for the conservation of melliferous
insects such as bees (Nicholls & Altieri, 2013).
Considering the above, the present study aimed to highlight traditional indigenous
knowledge as a base to design complex agroecosystems and, altogether with scientific
knowledge, contribute to find options to face global warming, the loss of biodiversity and
rural poverty.
2 Materials and methods
2.1 Study area
The different sections of this study were conducted in seven ejidos (communal lands
used for agriculture) from the municipality of Soteapan, in the Sierra de Santa Marta,
Veracruz. This region is part of the buffer zone of Los Tuxtlas Biosphere Reserve in the
littoral zone of the Gulf of Mexico (18°16′7" and 18º14′20″ N, 94°53′2″ and 94º57′23″
W, 480 to 980 m above sea level (masl), the terrain is characterized by middle to high
slopes, with slopes between 15 and 35% (Fig. 1). The predominant climate types in the
area are hot very humid (Afm), between 800 and 1000 masl, with annual precipitation
between 3500 and 4000 mm, and mean annual temperature between 22 and 24 °C; and
13
Fig. 1 Location of the study area, in red the sampling points of the present investigation. Map designed by
Daniel Alejandro Lara-Rodríguez
semi-hot very humid (A)C(fm), with a mean annual temperature higher than 18 °C and
located above 900 masl. It is important to mention that the sierra (mountain range) pro-
duces a rain shadow effect on the western plains and thus there is higher precipitation
than on the leeward side, a situation that favors the availability of water in the Sierra,
which supplies at least 600 thousand people in the most important cities in the middle
and lower basin of southern Veracruz (Gutiérrez-García & Ricker, 2011). The dominant
soil types are Andosols (52%), Acrisols (14%), Vertisols (12%), and Luvisols (11%).
In the Zoque-Popoluca classification, the predominant soil types are red (tsabats nas),
black (yik nas), and sandy (poy nas) (Ramírez-López et al., 2008).
The main vegetation types where the coffee agroecosystems are established or from
which different types of useful plants are obtained are tropical montane rain forest, trop-
ical semi-evergreen forest, tropical semi-deciduous forest, deciduous or montane cloud
forest, tropical pine forest, tropical oak forest, induced savannahs, and different succes-
sional stages of all of these vegetation types (Ramírez & Ramírez, 2009). The studied
coffee plantations are established in hillsides (slopes between 15 and 35%) as traditional
rustic or mountain agroecosystems (Moguel & Toledo, 1999), in areas formerly occu-
pied by tropical semi-evergreen forest. In these sites, people remove mainly lower strata
of the forest and maintain original species like Bursera simaruba, Cordia alliodora,
Dendropanax arboreus, Guazuma ulmifolia, Heliocarpus appendiculatus, Luehea spe-
ciosa, Spondias mombin, Trema micrantha, Vochysia guatemalensis, Swietenia macro-
phylla, Byrsonima crassifolia, Pinus oocarpa, Tabebuia rosea, and different species of
Inga and Quercus, among others, are preserved (Castillo-Capitán et al., 2014).
13
The population of Soteapan is mostly indigenous (92.3%) and it is one of the 100
municipalities with the highest marginalization index at a national level (Medel-Ramírez &
Medel-López, 2019). The main productive activities are agriculture, mainly shade-grown
coffee, maize (as a monoculture and polyculture of milpa), and subsistence livestock pro-
duction. Peasants are part of regional organizations, such as the Regional Union of Indig-
enous Popoluca Coffee Producers of the Sierra de Soteapan and the cooperative Jem Uxu-
ctero and local organizations, such as the Union of Indigenous Forest Producers Kalluy
Kotsik from the ejido San Fernando, which is a group of beekeepers that is not formally
established. The Rural Foliage and Product Production Society Wɨty Cotsɨk, made up of
peasants from nine communities, whose main objective is the production of ornamental
palms (Chamaedorea spp.) in the understory of coffee agroecosystems. Some peasants
have accessed resources to carry out forest restoration projects in the Texizapan stream
basin, which include a forest nursery in the community of Ocotal Grande (Rodríguez et al.,
2012).
2.2 Diagnosis of the coffee agroecosystems
Field trips and participatory workshops were conducted with 36 peasant leaders from the
ejidos of San Fernando, Ocotal Grande, Ocotal Chico, Ocozotepec, Santa Marta, Sotea-
pan, and El Tulín; all of them are coffee growers with complementary activities such as
palm growing, beekeeping, and planting trees for reforestation, these peasants have been
managed these agroecosystems for long time, so they have developed a fine understand-
ing of their ecological functioning. With these peasants, 66 sampling sites were located to
structurally characterize the coffee agroecosystems (Fig. 1). Sampling plots of 20 m × 20 m
were established at each site and the tree species present were recorded in a list, includ-
ing the following measurements for each species: normal diameter (at 1.30 m from the
ground), total and trunk height (m), and cover (m2). Based on the physiognomy of each
site, three layers were visually determined: upper (up to 35 m), middle (up to 25 m), and
lower (< 17.5 m). For each layer, trees were grouped into diameter categories: upper, with
four categories (< 10.0 cm, 10.1–20.0 cm, 20.1–30.0 cm, and > 30.1); middle, three catego-
ries (< 10.0 cm, 10.1–20.0 cm, and 20.1–30.0 cm); and lower, four categories (< 5.0 cm,
5.1–10.0 cm, 10.1–15.0 cm, and 15.1–20.0 cm). Similarly, the soil type where each species
was present was identified following the local Zoque-Popoluca classification (Ramírez &
Ramírez, 2009). The knowledge peasants share about the agroecosystem functioning is key
to design sustainable and complex systems not only for planning the use of their territory,
but also to face the loss of biodiversity and climate change in our planet.
2.3 Estimation of carbon sequestration
According to Navarro-Martínez et al. (2020), the lack of standardized models for the
estimation of aerial biomass (in kg) using different variables is a source of uncertainty.
To minimize this, we used allometric equations for broadleaf species obtained in tropi-
cal zones with environmental conditions like those of the study area, carefully verifying
that the physiognomic and wood density characteristics of the species were like those of
the models used (Table 1). Allometric equations are regression models that relate statisti-
cally few easy-to-measure characteristics of trees (diameter, height, wood density) to esti-
mate destructive harvest measurements (volume, biomass, carbon stored), based on that
proportions between such groups of features follow rules (represented by the models) for
13
Table 1 Models of specific and general allometric equations chosen to calculate the biomass of the tree species found in the sampling sites
Species Formula References
13
Andira galeottiana Y = exp(− 3.1141 + 0.9719 * ln(DBH2 * H)) Brown et al. (1989)
Annona cherimola
Annona muricata
Apeiba tibourbou
Calophyllum brasiliense
Cojoba arborea
Cordia megalantha
Gmelina arborea
Guarea glabra
Mangifera indica
Muntingia calabura
Pachira aquatica
Pimenta dioica
Sloanea medusula
Spondias mombin
Tabebuia crhysantha
Tabebuia rosea
Tamarindus indica
Virola guatemalensis
Ficus yoponensis Ya = exp(4.9375) * ((DBH2)^1.0583) * (1.14/1000) Hughes et al. (1999)
Heliocarpus appendiculatus Yb = exp(− 3.1141 + 0.9719 * ln (DBH2 * H))
Lonchocarpus guatemalensis
Miconia argentea
Nectandra ambigens
Pseudolmedia oxyphyllaria
Astrocaryum mexicanum Y = (exp(3.6272 + 0.5768 * ln (DBH2*H)))*1.02/106 Hughes et al. (1999)
Chamaedorea spp.
Ch. tepejilote
Brosimum alicastrum Y = 0.0336 * (DBH2)*H Cairns et al. (2003)
Bursera simaruba Y = 0.064808 * (DBH2.46998) Rodríguez-Laguna (2008)
Byrsonima crassifolia Y = exp(− 1.696 + 2.226 * ln(DBH)) van Breugel et al. (2011)
Cecropia obtusifolia
Y = (exp(-3.78 + (0.95*log(DBH) + log(H)) Hughes et al. (1999)

Table 1 (continued)
2 0.9218 2 1.0451 2
Cedrela odorata Y = 0.0013*DBH *H + 0.0072*DBH *H + 0.0029*DBH Cole and Ewel (2006)
*H1.0172 + 0.0010*DBH2 *H0.8038 + 0.0893*DBH2 *H0.5326
Citrus limon Y = 6.64 + 0.279 * BA + 0.000514 * BA2 Schroth et al. (2002)
Citrus sinensis
Clethra mexicana Y = 0.4632 * DBH1.8168 Acosta et al. (2011)
Coffea arabica Y = exp(− 0.66) * DBH1.37 Acosta (2003)
Cordia alliodora Y = 10^(− 0.755) * DBH2.072 Segura et al. (2006)
Dendropanax arboreus Y = 0.037241 * DBH2.99585 Rodríguez-Laguna (2008)
Gliricidia sepium Y = 3.27 * exp(0.075*DBH) Gómez-Castro et al. (2010)
Guazuma ulmifolia Y = exp(− 1.62) * DBH2.12 Douterlungne et al. (2013)
Inga jinicuil Y = exp(− 1.76) * DBH2.26 Acosta-Mireles et al. (2002)
Inga leptoloba
Inga vera
Leucaena leucocephala
Y = exp(− 1.46 + 1.69 ln (DBH)) Gómez-Castro et al. (2010)

Luehea speciosa Y = exp(− 2.12605 + 0.868 * ln(DBH2 * H))
Liquidambar styraciflua Y = 0.180272 * DBH2.27177 Rodríguez et al. (2006)
Manilkara zapota Ya = 0.0447 * DBH2 * H Cairns et al. (2003)
Pouteria sapota Yb = 0.0034 + 0.0482 * DBH2 *H
Melicoccus oliviformis Y = 0.0493 + 0.0480 * DBH2 * H
Musa paradisiaca Y = 0.0303*DBH2.1345 Espinoza-Domínguez et al. (2012)
Pinus oocarpa Y = − 10.4113 + 5.3998 * DBH + 0.8144 * DBH2 González (2008)
Pouteria reticulata Y = − 1.716 + 2.413 * ln (DBH) van Breugel et al. (2011)
Psidium guajava Y = 0.246689 * DBH2.24992 Rodríguez-Laguna (2008)
Quercus conspersa Y = exp(− 2.27)* DBH2.39 Acosta-Mireles et al. (2002)
Q. glaucescens
Q. oleoides
Q. peduncularis
13
Table 1 (continued)
13
Swietenia macrophylla Y = 10^(− 4.00692 + 2.51389 * log10(DBH)) * 1000 Arreaga (2002)
Terminalia amazonia Y = exp (− 1.675 + 2.326* ln (DBH)) van Breugel et al. (2011)
Trema micrantha
Trichilia havanensis Y = 0.130169 * DBH2.34924 Rodríguez-Laguna (2008)
Vochysia guatemalensis Y = exp(− 1.44742 + 1.3308* DBH0.5) Fonseca et al. (2009)
DBH Diameter at breast height, H Total height, Y Biomass, ln Natural logarithm, log logarithm, exp exponent, BA Basal area
a
Diameter interval ≤ 10 cm
b
Diameter interval ≥ 10 cm

all the trees or groups of plants if they are growing under similar conditions or ecological
zones (Chave et al., 2005, Gibbs et al., 2007). The equations used are available in the scien-
tific literature and in different databases (Rojas-García et al., 2015). In the case of species
without a specific equation, generic models were used (Acosta-Mireles et al., 2002; Brown
et al., 1989). The equations were selected under the principle of parsimony, i.e., models
that use the fewest possible variables (Otto & Day, 2011).
2.4 Species valuation by peasants
Three participatory workshops were carried out with the peasants; older people are usually
the ones with more knowledge and experience, and they are consulted about the names
of forest species. Young people also give their view but respecting the opinions of older
people. It is important to note that some producers suggested carrying out field trips to
obtain the information about tree species, once the trips were made, their participation was
more active; then altogether create lists of attributes that make them valuable for the coffee
agroecosystems. In addition to the species present, peasants identified, based under their
knowledge, desired species, or others of interest. Both types of species were ranked based
on the attributes proposed by Yépez (2001) and those identified by peasants, for the design
of coffee agroecosystems according to the following weighing: foliage phenology (ever-
green or deciduous; values from 1 to 3), umbrella-shaped crown (1 to 3), growth speed (1),
and pruning tolerance (1 to 2). This proposal originally includes an attribute called “other
products”, which in this study was replaced by type of use based upon the knowledge and
importance that the different species has for the peasants (fruit, timber, firewood, medici-
nal, forage, and ornamental); a value of 1 was given to each use. To these values were
added the corresponding values of melliferous/nectariferous potential (between 1 and 2)
and C sequestration potential (1 to 4); the latter was based on the percentiles of C seques-
tration values. The assessment of the melliferous contribution of the species was made
by reviewing the databases of Mexican institutions (CONABIO, SAGARPA, INIFAP)
and global databases (Bee World, Agroforestree Database, and FAO), as well as different
sources of information from studies conducted in tropical regions. Based on the obtained
information, the species were classified as melliferous, nectariferous, potentially mellif-
erous, or nectariferous, and without information. Thus, the maximum values that can be
obtained per attribute are: 3 for evergreen or deciduous foliage, crown shape, and carbon
sequestration; for growth speed and melliferous/nectariferous potential the maximum is 1
per indicator; for the type of use the maximum is 6 if it includes all six uses, and for prun-
ing tolerance the maximum is 2. The sum of the values obtained per attribute would result
in a maximum value of 20. The total value of each species was used to make the proposals
for the agroecosystem design. The flowering and fruiting seasons recognized by peasants
were also recorded to include species that produce flowers and fruits throughout the year to
produce honey, improve the economic income, and maintain pollinator populations.
2.5 Agroecosystem design
The agroecosystem design was based on the global valuation of the species attributes
(Yépez, 2001). The proposals include at least twelve native tree species per hectare and
non-native species of high interest to the peasants, without letting the main species exceed
60% dominance. A density of 70 trees per hectare was considered, with at least 40%
being evergreen, with a minimum height of 15 m, and at least three layers. The number of
13
individuals per plot was obtained with the mean tree canopy (using the minimum and max-
imum diameter per species, with the formula: cover = π*r2) to achieve 40% cover (4000
m2). This attribute (per species) was multiplied by the number of plants suggested for each
tree to obtain both the area covered per species and the total area. For the minimum total
area in the dry season, only the sum of the cover of evergreen species was considered.
The information was integrated and an arrangement was defined according to the altitudi-
nal gradient of the study area, considering the soil type of the sampled plots (Ramírez &
Ramírez, 2009).
3 Results
3.1 General characteristics of the agroecosystems
In the studied agroecosystems, 49 different tree species were found, several of which are
mentioned by Miranda and Hernández-X (2014) as being part of tropical rain forests (Cal-
ophyllum brasiliense, Swietenia macrophylla, Terminalia amazonia, and Vochysia gua-
temalensis, among others) and tropical semi-evergreen forests, or advanced successional
stages of these forests (Cecropia spp. and Heliocarpus spp., among others). In the present
study, the species of the middle stratum represent 63.3%, the species of the upper stra-
tum 34.1%, and those of the lower stratum only 2.6%. This reflects how important it is
for the peasants to preserve shade species for the coffee plants that do not compete spa-
tially. A total of 597 individuals were found in the 66 sampling sites located in seven ejidos
(Fig. 1), in the three layers, comprised by the following species: Inga vera (chalahuite),
32%; Liquidambar styraciflua (ocozote), 27.5%; Cordia alliodora (solerillo), 4.1%; and
Trema micrantha (capulín), 3.6%. Species such as Calophyllum brasiliense (barí), Termi-
nalia amazonia (tepesuchil), Swietenia macrophylla (caoba), and Vochysia guatemalensis
(corpo) each represent less than 1%.
3.2 Carbon sequestration by stratum and species
The species that potentially sequester the highest amount of C are found in the upper and
middle layers (Figs. 2 and 3). The most abundant species do not sequester the highest
amount of C, although this varies with the diameter category of each species. For exam-
ple, the most abundant species in the high stratum is Liquidambar styraciflua, but it only
sequesters 290.2 kg of carbon (kg C), however, it is one of the most efficient species in
the 7.5 cm diameter category. In contrast, other less abundant species in the same stra-
tum can fix more C, such as Pouteria sapota (1204.9 kg C), Manilkara sapota (958.3 kg
C), Brosimum alicastrum (720.3 kg C), Calophyllum brasiliense, Cojoba arborea, Cor-
dia megalantha, Nectandra ambigens, Pseudolmedia oxyphyllaria, and Sloanea medusula
(705.6 kg C each) (Fig. 2). In the Middle stratum, the most abundant species, Inga vera,
can sequester up to 124.15 kg C, and the three species of Quercus sequester 113.29 kg
C. However, Dendropanax arboreus, a species with low abundance (two individuals), can
sequester up to 287.08 kg C. Apeiba tiburbo, Ficus yoponensis, Heliocarpus appendicula-
tus, Lonchocarpus guatemalensis, Mangifera indica, Miconia argentea, Pachira aquatica,
Pouteria reticulata, and Tabebuia chrysantha can sequester up to 264.55 kg C (Fig. 3).
Finally, in the low stratum, the species with the highest value was Meliccocus oliviformis
(128.6 kg C), followed by Annona cherimola and Spondias mombin (93.5 kg C), Psidium
13
1200.0
Potential C sequestration tree kg
1000.0
800.0
600.0
400.0
200.0
0.0
2.5 cm 7.5 cm 12.5 cm 15 cm 17.5 cm 25 cm 35 cm
DBH
Bursera simaruba Cedrela odorata Cojoba arborea Sloanea medusula
Swietenia macrophylla Terminalia amazonia Vochisia guatemalensis Brosimum alicastrum
Clethra mexicana Liquidambar styraciflua Manilkara zapota Nectandra ambigens
Fig. 2 Potential C sequestration by species in the higher stratum of the coffee agroecosystems studied
400.0
Potential C sequestration tree kg
300.0
200.0
100.0
0.0
2.5 cm 7.5 cm 12.5 cm 15 cm 17.5 cm 25 cm
DBH
Byrsonima crassifolia Cordia alliodora Cecropia obtusifolia Dendropanax arboreus
Ficus yoponensis Heliocarpus appendiculatus Inga jinicuil Inga vera
Luehea speciosa Lonchocarpus guatemalensis Mangifera indica Miconia argentea
Pachira aquatica Pimenta dioica Quercus conspersa Quercus glaucescens
Quercus peduncularis Quercus oleoides Trema micrantha Guazuma ulmifolia
Fig. 3 Potential C sequestration by species in the middle stratum of the coffee agroecosystems studied
guajaba (77.2 kg C), Citrus sinensis (51.7 kg C), Coffea arabica (13.0 kg C), and Gliri-
cidia sepium (6.1 kg C). Banana plants (Musa paradisiaca) can sequester up to 16.3 kg C,
and C sequestration by two Chamaedorea species and the chocho palm (Astrocarium mexi-
canum) is negligible (Fig. 4).
Some pioneer species common in coffee agroecosystems, such as Trema micrantha and
Cecropia obtusifolia, have reduced C sequestration potential.
3.3 Valuation of tree species
As a result of the participatory workshops, peasants give the highest values to Mangifera
indica, Guazuma ulmifolia, and species of the genus Inga (Table 2). Peasants also give a
high value to some non-native species that produce timber or fruits and are part of the cof-
fee landscapes, such as Citrus limon, Citrus sinensis, Gmelina arborea, and Tamarindus
indica.
The knowledge that peasants have about the distribution of the flowering and fruiting
months (Table 3) of the tree species allows identifying which species can be incorporated
13
140.0
Potential C sequestration kg
120.0
100.0
80.0
60.0
40.0
20.0
0.0
2.5 cm 7.5 cm 12.5 cm 15 cm 17.5 cm
DBH
Citrus sinensis Psidium guajava Spondias mombin Coffea arabica
Gliricidia sepium Leucaena leucocephala Melicoccus oliviformis Miconia argentea
Annona cherimola Citrus limon Chamaedorea spp Chamaedorea tepejilote
Astrocarium mexicanum Musa paradisiaca Annona muricata
Fig. 4 Potential C sequestration by species in the lower stratum of the coffee agroecosystems studied
into the coffee agroecosystem to meet the demand for pollen or nectar at different times of
the year and to provide a potential source of additional income for the coffee agroecosys-
tems owners.
3.4 Agroecosystem design
Based on the information obtained in the previous sections and the mean canopy per spe-
cies, designs were proposed according to the altitudinal gradient. Two examples are shown
in Tables 4 and 5, as well as in Figs. 5 and 6.
In this case, the total number of plants is 319 (60 deciduous and 259 evergreen). The
maximum cover area for the dry season is 4040.1 m2 and the total maximum cover is
5265.3 m2. The preference of the peasants for species of the genus Inga is notable, although
it is important to note that both Bursera simaruba and Trema micrantha are highly valued
species.
In this section of the altitudinal gradient, the total number of plants would be 300
(60 deciduous and 240 evergreen). The maximum cover area expected for the dry sea-
son is 4,053 m 2. The total maximum cover would be 5,161 m 2. As in the previous case,
the peasants prefer species of the genus Inga, as well as for Bursera simaruba and Trema
micrantha.
4 Discussion
4.1 Agroecosystems, their management and value for peasants
The way that peasants manage their agroecosystems has a strong influence on the struc-
turing of each space, which has been reported in other coffee agroecosystems in Mexico
and the world (Ehrenbergerová, 2016; Häger, 2012; Hergoualc’h et al., 2012; Segura
et al., 2006; Terán-Ramírez et al., 2018; Valencia et al., 2014). Many of the species man-
aged by peasants have been observed by different authors as part of successional stages in
the humid forests of Mexico. For example, Spondias mombin, Trema micrantha, Cecro-
pia obtusifolia, Cordia alliodora, Bursera simaruba and Terminalia amazonia stand out,
13
Table 2 Desirable characteristics, based mainly on peasant knowledge, of the tree species that compose the studied agroecosystems and that can be used for the design of
diversified agroecological matrices
Uses
Scientific Common Present (P) Foliage Cover Growth Fruit/ Timber Firewood Medicinal Forage Posts Mel- C seques- Final valu-
name name or desired produc- speed Edible liferous tration ation
(D) tion potential
Mangifera Mango D 3 3 3 1 1 1 – – – 2 2 16
indica
Guazuma Guácimo P 3 3 3 – 1 1 – – – 2 1 14
ulmifolia
Inga jini- Jinicuil D 3 3 3 – 1 1 – – – 2 1 14
cuil
Inga vera Chalahuite P 3 3 3 1 – 1 – – – 2 1 14
Bursera Palo P 2 2 3 – – – 1 1 1 2 2 14
simaruba mulato
Cedrela Cedro D 3 3 1 – 1 1 1 – – 2 3 15
odorata
Quercus Encino P 3 3 3 – 1 1 1 – – 1 1 14
glause- blanco
cens
Tabebuia Primavera P 3 3 3 – 1 1 – – – 1 2 14
chrysan-
tha
Cordia Solerillo P 2 3 2 – 1 – – – 1 2 1 13
alliodora
Pimenta Pimienta D 2 2 2 – 1 1 1 – – 2 2 13
dioica
Spondias Ciruelo P 3 3 3 – 1 1 – – 1 1 1 14
mombin
Tabebuia Roble P 3 3 3 – 1 1 – – – 2 3 16
rosea
13
Table 2 (continued)
Uses
13
(D) tion potential
Terminalia Tepesúchil D 3 3 3 – 1 1 – – – 2 3 16
amazonia
Vochysia Corpo P 3 2 2 – 1 1 – – – 2 3 14
guate-
malensis
Byrsonima Nanche P 2 2 1 1 1 1 1 – 1 1 1 12
crassi-
folia
Dendro- Palo de P 3 3 3 – 1 1 – – – 2 13
panax agua
arboreus
Inga lep- Vaina D 3 3 3 – – 1 – – – 2 2 14
toloba
Pouteria Zapote D 2 2 3 1 1 1 – – – 2 4 16
sapota mamey
Andira Macayo P 3 3 2 – 1 1 – – 1 – 3 14
galeot-
tiana
Brosimum Ramón D 3 3 1 – 1 1 – – 1 1 4 15
alicas-
trum
Citrus Limón D 2 3 3 1 – – 1 – – 1 1 12
limon
Citrus Naranja P 2 2 2 1 – 1 1 – – 1 1 11
sinensis

Table 2 (continued)
Uses
(D) tion potential
Gmelina Melina D 3 3 3 – 1 1 – – – – 3 14
arborea
Helio- Jobo P 3 3 3 – 1 1 – – – – 2 13
carpus
appen-
diculatus
Tama- Tamarindo D 2 2 2 1 1 1 – – – 2 3 14
rindus
indica
Luehea Tepecacao P 2 2 2 – 1 – – – 1 2 2 12
speciosa
Muntingia Capulín D 2 2 2 – – 1 1 – – 2 3 13
calabura cimarrón
Psidium Guayaba D 2 2 2 – – 1 1 – – 2 1 11
guajava
Trema Ixpepe P 3 3 3 – 1 – – – – – 1 11
micran-
tha
Virola Cedrillo D 2 2 2 – 1 1 – – – 2 3 13
guate-
malensis
Annona Guanábana D 1 2 3 1 – – – – – 2 2 11
muricata
Ficus Hule D 3 3 3 – – – – – – – 2 11
yopon-
ensis
13
Table 2 (continued)
Uses
13
(D) tion potential
Guarea Palo P 2 3 1 – 1 1 – – 1 – 3 9
glabra blanco
Lonchocar- Palo P 1 2 3 – – – – 1 – 1 2 10
pus gusano
guate-
malensis
Pinus Ocote D 2 2 2 – 1 1 1 – – – 3 12
oocarpa
Swietenia Caoba P 2 2 2 – 1 1 – – – – 3 11
macro-
phylla
Attalea Palma real D 2 3 2 1 – – – – – – – 8
butyra-
cea
Chamae- Palma D 3 2 3 – – – – – – – – 8
dorea camedor
spp.
Chamae- Tepejilote D 2 2 3 – – – – – – 1 – 8
dorea
tepejilote
Coffea Café P 2 2 2 1 – – – – 2 1 10
arabica
Manilkara Chicoza- D 2 2 2 – – 1 – – – 1 4 12
zapota pote

Table 2 (continued)
Uses
(D) tion potential
Calophyl- Barí D 2 2 2 – 1 1 – – – 3 11
lum bra-
siliense
Pachira Apompo D 2 2 3 – – 1 – – – – 2 10
acuatica
Astro- Chocho D 1 1 2 1 – – – – – 2 – 7
caryum
mexica-
num
Cecropia Chancarro D 1 1 3 – – – 1 – – 1 1 8
obtusi-
folia
Musa Plátano D 1 1 3 1 – – – – – 1 1 8
paradisi-
aca
Quercus Encino P – – – – – – – – – 2 1 3
oleoides negro
Cojoba Frijolillo P – – – – 1 – – – – 1 4 6
arborea
Meliccocus Guaya D – – – – – – – – – 2 1 3
olivi-
formis
Sloanea Palo de D – 2 – – – – – – – – 4 6
medusula achote
13
Table 2 (continued)
Uses
13
(D) tion potential
Quercus Encino P – – – – – – – – – 1 1 2
peduncu- amarillo
laris

Table 3 Calendar of the distribution of production of flowers and edible fruits by species with melliferous potential
Flower production Edible fruits production
Scientific name Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov
Annona muricata x x x x
Annona diversifolia, A. pur- x x
purea
Annona purpurea x x
Annona reticulata x x x x x
Astrocaryum mexicanum
Brosimum alicastrum x x x x
Bursera simaruba x x x
Byrsonima crassifolia x x x x
Cecropia obtusifolia x
Ceiba pentandra x
Cedrela odorata x x
Chamaedorea spp.
Chamaedorea tepejilote
Citrus limon x x x x
Citrus sinensis x x x x x x
Cojoba arborea x x x
Cordia alliodora x x x
Dendropanax arboreus
Ficus yoponensis
Gmelina arborea
Guarea glabra
Guazuma ulmifolia x x x
Heliocarpus appendiculatus
Inga jinicuil x x x x x x
Inga leptoloba x x x x x x
13
Table 3 (continued)
13
Inga vera x x x x x x
Lonchocarpus guatemalensis x
Luehea speciosa
Mangifera indica x x x x x x x
Manilkara zapota x x x x
Meliccocus oliviformis x x x x
Muntingia calabura x x x
Musa paradisiaca x x x
Pachira acuatica
Pimenta dioica x x x
Pinus oocarpa
Pouteria sapota x x x
Psidium guajava x
Quercus glausecens x x x x
Quercus oleoides x x x x
Quercus peduncularis x x x x
Sloanea medusula
Spondias mombin x x x x x
Swietenia macrophylla
Tabebuia chrysantha x x x
Tabebuia rosea x x x
Tamarindus indica x x x
Terminalia amazonia
Trema micrantha

Table 3 (continued)
Virola guatemalensis
Vochysia guatemalensis x x
Bahunia sp. x x x
Cinnamomum zeylanicum x
Citrus limon x x
Citrus sinensis x x
Cochlospernum vitifolium x
Coffea arabica x x x x x x x x x x x x x
Cordia megalantha x x
Croton schiedeanus x
Cupania dentata x
Delonix regia x
Gliricidia sepium x x
Ilex belizensis x
I. quercetorum
I. valeri
Inga vera x x x x x x x x x x x x x x x x x x x x x x x x
Liquidambar styraciflua x
Quercus affinis x
Q. skinneri
Spondias lutea x
Spondias purpurea
13
Table 4 Species selected for the design of coffee agroecosystems located between 400 and 500 masl; 23 tree species for the three shade layers
Layers Scientific name n Mean cover, m
2 C sequestra- Mean value for Number of tradi- Melliferous Global
tion shaded coffee tional uses potential valuation
13
Deciduous species
Upper Cedrela odorata 10 71 3 2 3 2 15
Middle Tabebuia rosea 10 71 3 3 2 2 16
Middle Guazuma ulmifolia 10 196 1 3 2 2 14
Middle Cordia alliodora 10 196 1 2 2 2 13
Lower Spondias mombin 10 71 1 3 3 1 14
Lower Heliocarpus appendiculatus 10 196 2 3 2 0 13
Evergreen species
Upper Terminalia amazonia 15 106 3 3 2 2 16
Upper Vochysia guatemalensis 15 106 3 2 2 2 14
Upper Swietenia macrophylla * 15 106 3 2 2 0 11
Upper Guarea glabra 15 106 3 2 3 0 9
Upper Bursera simaruba 15 295 2 2 3 2 14
Middle Virola guatemalensis 15 106 3 2 2 2 13
Middle Inga leptoloba 20 141 2 3 1 2 14
Middle Pimenta dioica 15 106 2 2 3 2 13
Middle Dendropanax arboreus 15 106 2 3 2 0 13
Middle Annona muricata* 15 106 2 2 1 2 11
Middle Lonchocarpus guatemalensis 15 106 2 2 1 1 10
Middle Inga jinicuil * 20 393 1 3 2 2 14
Middle Inga vera 20 141 1 3 2 2 14
Middle Byrsonima crassifolia 15 71 1 2 5 1 12
Middle Trema micrantha 15 295 1 3 1 0 11
Lower Citrus limon 12 85 1 3 2 1 12
Lower Citrus sinensis 12 85 1 2 3 1 11
Data for C sequestration, mean value for shaded coffee, number of traditional uses, melliferous potential, and global valuation were obtained from Table 2
n = Number of individuals by specie. Mean cover = cover provided by n individuals of each specie
*Species that can be deciduous or evergreen according to climate conditions

among the most important (Miranda, 1961; Lecy-Tacher et al., 2002; Levy-Tacher & Agu-
irre, 2005). Martínez (1985) identifies some of these species as long-lived that seem to
"seek" clearings, however, in managed spaces, such as those studied, they are rather pre-
ferred by peasants, which is reflected in the high valuation of species such as B. sima-
ruba, S. mombin, Guarea glabra, Inga spp. and Ficus yoponensis. In this sense, peasants
manipulate the environment, eliminate trees, conserve others, contributing to establish a
co-evolutionary process with some trees, the foregoing may constitute what Zahory (2004)
calls an unconscious process of domestication. For Puettmann et al. (2009), this constitutes
another way of doing forestry or agroforestry, more holistic and comprehensive, that is to
say, peasant empirical science, contributes to generating models that are persistent, adapt-
able and variable, recognizing that they are dynamic and complex systems. Häger (2012)
studied the importance of trees in coffee agroecosystems with up to 20 different species
and considered that it is essential to determine the potential synergies that occur in highly
diversified tropical agroecosystems when sequestering C to mitigate global warming, one
of this synergies, frequently highlighted by peasants in our case and documented for many
species of secondary succession (Selaya et al., 2008), is the decomposition of different
types of leaves, for instance, Trema micrantha is a fast growing species that also has broad
crown, very useful in coffee agroecosystems since it creates a microclimate that allows the
establishment of other pioneer species. Furthermore, its high biomass productivity in terms
of photosynthetic rate, associated with a high leaf area index, allows it to intercept more
light than other pioneer species and produces more litter, protecting soils from erosion
(Vázquez-Yanez, 1998); the leaves of other species like Tabebuia sp. and Cecropia obtusi-
folia, also incorporate very quickly into the soil because they are riche in cellulose content,
stimulating microbial activity and interactions between different species of the soil and the
plants (Mesquita et al., 1998; Loranger et al., 2002). The fruits of C. obtusifolia are highly
consumed by specialized frugivores but also by opportunists, being thus highly important
for animal conservation, and its leaves and bark have medicinal uses.
4.2 Valuation of tree species and C sequestration
The structure of the studied coffee agroecosystems depends largely on the use value that
the tree species have for the peasants, in this sense, firewood is the main use (61.2%) fol-
lowed by construction (55), food (24.5), medicinal (22.4), fencepost (16.3) and forage (4).
Species like Inga vera and I. leptoloba are highly valued for construction, firewood, and
fruits or for being evergreen, and it is the same case for Byrsonima crassifolia, Spondias
mombin and Pimenta dioica, among others, the former has five different uses and the other
two, four and three different uses, and these peasants criteria increases its value when
structuring the agroecosystems (Ehrenbergerová, 2016; Häger, 2012; Hergoualc’h et al.,
2012; Segura et al., 2006; Terán-Ramírez et al., 2018; Valencia et al., 2014).
Five of the species with the highest carbon sequestration potential have at least three
different uses. For example, Brosimum alicastrum and Pouteria sapota can have restorative
effects on the soil, by providing easily degradable organic matter, and deliver habitat and
food to wild species. Bursera simaruba is one of the most frequently used species as living
fences in tropical areas of Mexico, its leaves have a high rate of decomposition (Loranger
et al., 2002), is used for medicine and forage, and to treat fever and headache (Leonti,
2002). Peasants use Swietenia macrophylla and Cedrela odorata for timber and firewood,
and the latter is also used as an anti-inflammatory or against hemorrhages (Leonti, 2002;
Blanco, 2006). Pimenta dioica is used to provide shade to coffee plants and allows the
13
Table 5 List of 17 selected species for the design of coffee agroecosystems, with three shade layers, located between 800 and 1000 masl
Layers Scientific name n Mean cover, m2 C sequestr Mean value for Number of tradi- Melliferous Global
13
shaded coffee tional uses potential valua-
tion
Deciduous species
Upper Cedrela odorata 10 71 3 2 3 2 15
Middle Tabebuia rosea 10 71 3 3 2 2 16
Middle Guazuma ulmifolia 8 157 1 3 2 2 14
Middle Cordia alliodora 8 157 1 2 2 2 13
Lower Spondias mombin 10 71 1 3 3 1 14
Lower Heliocarpus appendiculatus 8 157 2 3 2 0 13
Evergreen species
Upper Swietenia macrophylla * 15 106 3 2 2 0 11
Upper Bursera simaruba 15 295 2 2 3 2 14
Middle Inga leptoloba 20 141 2 3 1 2 14
Middle Dendropanax arboreus 15 106 2 3 2 0 13
Middle Annona muricata* 15 106 2 2 1 2 11
Middle Inga jinicuil * 20 393 1 3 2 2 14
Middle Inga vera 20 141 1 3 2 2 14
Middle Byrsonima crassifolia 15 71 1 2 5 1 12
Middle Trema micrantha 15 295 1 3 1 0 11
Lower Citrus limon 12 85 1 3 2 1 12
Lower Citrus sinensis 12 85 1 2 3 1 11
Data for C sequestration, mean value for shaded coffee, number of traditional uses, melliferous potential, and global valuation were obtained from Table 2
n = Number of individuals by specie. Mean cover = cover provided by n individuals of each specie
*Species that can be deciduous or evergreen according to climate conditions

Fig. 5 Proposed design of an agroecosystem with four layers. Acronyms mean: AnMu = Annona muri-
cata; AsMe = Astrocarium mexicanum; BuSi = Bursera simaruba; CeOd = Cedrela odorata; CiLi = Citrus
limon; CiSi = Citrus sinensis; CoAll = Cordia alliodora; CoAr = Coffea arabica; ChEl = Chanaedorea ele-
gans, GuUl = Guazuma ulmifolia; HeAp = Heliocarpus appendiculatus; InJi = Inga jinicuil; PiDi = Pimenta
dioica; SpMo = Spondias mombin; SwMa = Swietenia macrophylla; TaRo = Tabebuia rosea; TeAm = Termi-
nalia Amazonia; TrMi = Trema micrantha; VoGu = Vochisia guatemalensis. In top-right, schematic in-plant
agroecosystem design, example of trees distribution in coffee agroecosystem. Designed by Diego Ávila-
Ruiz
control of different weeds, has potential for reforestation, and is also used as a living fence
and to provide shade to livestock (Vázquez-Yanes et al., 1999). In the study area, allspice
seeds are a source of income for some Popoluca communities, its wood is used in construc-
tion, tool manufacturing, and as fuel, and its bark and leaves appear to have anti-diarrheal
and anti-dysenteric properties (Leonti, 2002). In addition to its documented uses, Vochysia
guatemalensis is used for construction, furniture, domestic tools, and toys (Blanco, 2006),
as well as a windbreak barrier, and its high litter production increases organic matter lev-
els, improving degraded soils, and also accumulates high aluminum levels in its leaves
(Cordero, 2003). Luehea speciosa is a species widely used in the study area to provide
shade to coffee plants, for domestic tools, and as fuel (Blanco, 2006). Manilkara zapota
is an important tree not only for its high carbon sequestration capacity, but also for the
recovery of degraded land, with more or less open canopy (Vázquez-Yanes et al., 1999).
The fruit of this species is highly valued for its flavor, its wood is used in construction,
and its bark has medicinal use (Leonti, 2002; Blanco, 2006). It is important to note that
the peasants showed great interest in Astrocaryum mexicanum, Chamaedorea tepejilote,
Chamaedorea spp., and Attalea butyracea due to their commercial importance as orna-
mental and edible species, even though the first three sequester an insignificant amount of
carbon and an adequate equation was not found for the last one. Some species with high
13
Fig. 6 Proposed design of an agroecosystem with four layers. Acronyms mean: AnMu = Annona muri-
cata; AsMe = Astrocarium mexicanum; BuSi = Bursera simaruba; CeOd = Cedrela odorata; CiLi = Cit-
rus limon; CiSi = Citrus sinensis; CoAll = Cordia alliodora; CoAr = Coffea arabica; ChEl = Chanaedorea
elegans, GuUl = Guazuma ulmifolia; HeAp = Heliocarpus appendiculatus; InJi = Inga jinicuil; InLe = Inga
leptoloba; InVe = Inga vera; PiDi = Pimenta dioica; SpMo = Spondias mombin; SwMa = Swietenia macro-
phylla; TaRo = Tabebuia rosea; TrMi = Trema micrantha. In top-right, schematic in-plant agroecosystem
design, example of trees distribution in coffee agroecosystem. Designed by Diego Ávila-Ruiz
carbon sequestration potential, such as Manilkara zapota, Pouteria sapota, Cojoba arborea
and Sloanea medusula, could be an important part of the design of coffee agroecosystems,
even though the last two have low use value for peasants. Other species that should be con-
sidered are those that are registered as threatened in the official Mexican standard (NOM-
059-SEMARNAT-2010) (SEMARNAT, 2010), such as Calophyllum brasiliense and Tabe-
buia chrysanta; others, not so highly valued, such as Trema micrantha, Andira geleotiana
and Pachira aquatica, have important environmental functions such as the incorporation of
litter into the system, soil protection, riverbank protection or shelter for birds and insects
that can establish different trophic relationships, which can range from predation, parasit-
ism or mutualism, which must also be studied to know them in detail.
4.3 Other functions of trees for agroecosystem design: melliferous potential

and fruit production
Sixty percent of the species have melliferous potential, species such as Pouteria sapota,
Manilkara zapota, Brosimum alicastrum, Tabebuia rosea, Terminalia amazonia and
Vochysia guatemalensis deserve attention, because in addition to having a high rate of
13
C capture, they have several uses for peasants, of 21% of the species no information was
found about their nectar or pollen contribution, and 10% were recorded as possibly mel-
liferous (Montoya-Pfeiffer et al., 2014). The species of the genus Inga, along with Cedrela
odorata, and Annona muricata, are recognized for the peasants and in different regions
as important melliferous sources for honey bees (Florez et al., 2002; Montoy, 2010). In
areas with high water availability, the melliferous and polliniferous potential of species of
the genus Quercus, particularly Q. oleoides, may be relevant since trees of this genus pro-
duce sweet sap through their pores, which is consumed by bees of the genus Melipona
(Ramírez-Arriaga et al., 2011).
The distribution of flowers and fruit production throughout the year complements pro-
ductive activities related to coffee cultivation. In general, the highest number of flowering
species occurs in the dry season (Cuevas et al., 2022) and the extension of the flowering
period depends on the region, being longer in the Pacific slope and the Yucatan Peninsula
in Mexico, due to the distribution of rainfall. On the other hand, the months with the lowest
number of flowering species are the months of rainfall (Villegas-Durán et al., 2000) or of
lower temperatures (Roman & Palma, 2007). It is important to mention that the peasants
could not specify the flowering season of species such as Anona muricata, Calophyllum
brasiliense, Inga vera, Manilkara sapota, and Inga jinicuil. Thus, future phenological stud-
ies can complete this information.
The production of fruits can represent a source of income or an improved diet for most
peasants. For most peasants, coffee production is complemented with shifting cultivation
of maize, harvested regularly in September and tapachol harvested in February. In the case
of flowering season, the production of edible fruit and seeds of some species varies with
orographic and micro climatic factors. The variation in the production periods can rep-
resent an advantage for peasants, since potential markets would be supplied at different
times of the year, preventing competition, and generating economic income. However, it is
common for only a small part of these products to be used for self-consumption and most
of them go to waste. This is not only the case of the ejidos evaluated here but has also been
reported in other studies conducted in coffee-growing areas (Philpott et al., 2007; Pinoar-
gote et al., 2017; Rice, 2011).
Even though the economic contribution of edible species was not quantified in the pre-
sent study, their importance can be inferred based on the price of the products in the market
and the value of coffee production. For example, fruit trees, such as orange, lemon, mango,
banana, and inga, in coffee agroecosystems in Peru and Guatemala have been estimated to
represent 11% of the value of coffee production (Rice, 2011). If the value of production
for self-consumption is also included, it represents between 5 and 63% of the income of
Peasants households in Chiapas (Philpott et al., 2007). The production system used also
influences the economic contribution, since income ranges from 20 to 60% of the produc-
tion value in shade polycultures with banana and citrus plants in Nicaragua, while income
is less than 20% in more diversified coffee agroecosystems (Pinoargote et al., 2017). In
organic production systems, despite diversification, income from edible species is less than
30% because of the higher production value of coffee, although they may represent addi-
tional income (Pérez-Grovas, 2000). However, it would be necessary to evaluate the impor-
tance of functional biodiversity, that is, more complex agroecological matrices could allow
the recovery of trophic relationships, whose role in pest and disease control is fundamen-
tal, and tree species that are under some protection category and whose populations have
declined sharply and could also allow the possible restoration of biogeochemical cycles.
It is important to note that the selected species were agreed upon with the owners of the
coffee agroecosystems with the purpose of having designs adapted to the studied altitudinal
13
gradient. Special emphasis was placed on including species with melliferous potential or
fruit or edible production, but species requested by the peasants were also included, such as
trees of the genus Inga. In the latter case, Bird Friendly suggests using species of this genus
as the main shade in coffee agroecosystems (SMBD, 2004), and it is also reported as one
of the most frequent genera in other coffee-producing regions (Haggar et al., 2001; Peeters
et al., 2003; Somarriba et al., 2004). Another aspect that could be relevant in further stud-
ies is that the perception of the species and uses identified is different between men and
women and it is related to historic division of tasks and daily assignments. For example,
women recognized more edible species (fruit and spices) and species for handicrafts. Men
mentioned as many species for medicinal uses as women, although women know in more
detail the use of each plant, as well as the form of use.
As mentioned before, in study the region coffee agroecosystems are traditional rustic
(Moguel and Toledo, 1999), peasants maintain original species favoring the growth of cer-
tain useful tree species. These species have different uses, shade trees or timber, medicine,
or food (Table 2). Our proposal seeks to maximize the different benefits and their spatial
arrangement looking for the distribution of benefits over the time. This proposal contem-
plates two types of change: on the one hand, the improvement of already established coffee
agroecosystems and on the other, the implementation of new coffee agroecosystems under
the proposed designs. We consider that this design can be also used in another agrofor-
estry systems (e. g., cacao-based agroecosystems) under similar environmental and cultural
conditions.
5 Conclusions
Traditional indigenous knowledge about different species that can be used to design agro-
ecosystems is fundamental, together with formal scientific knowledge, to be able to structure
complex agroecological matrices that represent, as in the case of the present study, an alter-
native to reverse global warming, diversify production, reduce poverty in the studied commu-
nities, and conserve or restore different trophic relationships in the landscape, among other
services. An example of this is that several of the species with lower C sequestration capacity
are highly valued for other attributes: in the lower stratum, Spondias mombin, Citrus limon,
and Psidium guajava; in the middle stratum, Guazuma ulmifolia, Trema micrantha, Tabebuia
chrisanta, different species of Inga, and Luehea speciosa; in the upper stratum, Terminalia
amazonia, Mangifera indica, Tabebuia rosea, Pouteria sapota, and Bursera simaruba; and in
the shrub and understory layer, Musa paradisiaca, Coffea arabica, Astrocaryum mexicanum,
Chamaedorea tepejilote, and Chamaedorea spp. Evergreen species should be favored in the
design of agroecosystems, since they will not only provide diversified shade but their cover
will also protect the soil from erosion. Fruit production can represent an important basis of
resources for peasants, but this remains to be evaluated. It is necessary to conduct phenologi-
cal studies along the altitudinal gradient in the case of Anona muricata, Calophyllum brasil-
iense, Inga vera, Inga jinicuil, and Manilkara sapota, which are species that can be used to
diversify the shade in coffee agroecosystems, obtain fruits, and sequester C.
The complex and dynamic management of traditional coffee agroecosystems makes it
necessary to carry out studies that allow understanding the dynamics of the populations
that compose them. For example, a fundamental aspect to be studied in the short term is
the role of functional biodiversity, such as the presence of antagonistic fungi, predatory
organisms, and pollinators, among others, which may allow gene flow between species
13
found in different vegetation patches. Twenty percent of the carbon emissions is caused by
tropical deforestation (Niles et al., 2002), so, studies like the present acquire more impor-
tance because management decisions of peasants, could be influenced by external policies,
for example the adoption of monocultures of coffee for the industry, affecting the structure
and functionality of coffee agroecosystems, and the ecosystem services they provide, for
instance, Balvanera et al. (2005), found that when tree diversity falls 40%, bee diversity
decreases in the same magnitude. Structuring and conserving complex coffee agroeco-
system matrices based upon traditional knowledge and formal science is very important
because of the synergies that exist between production of different goods and benefits to
the environment, for instance, conservation of soil microorganisms and fertility, avoiding
soil erosion; of course C sequestration and conservation of insects, traditional coffee agro-
ecosystems contains more birds and ant species than coffee monocultures in Brazil (Per-
fecto & Vandermeer, 2015). Also, increasing plant diversity has a positive effect on natural
enemy and pollinators abundance (Christmann, 2019; Harterreiten-Souza et al., 2014).
Based on participatory research and methods, traditional knowledge can be the basis
for structuring public policies that, from the local level and in similar environments, con-
tribute to the conservation of biological and cultural diversity, obtaining healthy food and
achieving food autonomy for each region. It is also important to recognize, both nationally
and internationally (for example in the Kyoto Protocol or the Bonn Agreement), that tradi-
tional shade coffee agroecosystems are complex forest spaces in which a high diversity of
trophic relationships are preserved and, therefore, they contribute significantly to reducing
climate change. Of course, a fundamental aspect that must be considered is respect for the
customary and legal rights of indigenous peoples and peasants over their territories and the
genetic resources contained therein.
Data availability The data that we have presented in the article "Design of complex agroecosystems: tradi-
tional and scientific knowledge to conserve agrobiodiversity in the Santa Marta Mountains, Veracruz, Méx-
ico" can be provided to anyone interested in reviewing or analyzing it, if requested.
Declarations
Conflict of interest The authors have no conflicts of interest to declare that are relevant to the content of this
article.
References
Acosta, M. (2003). Diseño y aplicación de un método para medir los almacenes de carbono en sistemas con
vegetación forestal y agrícolas de ladera de México Tesis doctoral. ColPos: Colegio de Postgradados.
Acosta, M. M., Carrillo, A. F., & Gómez, V. R. G. (2011). Estimación de biomasa y carbono en dos especies
de bosque mesófilo de montaña. Revista Mexicana De Ciencias Agrícolas, 2(4), 529–543.
Acosta-Mireles, M., Vargas-Hernández, J., Velázquez-Martínez, A., & Etchevers-Barra, J. D. (2002). Esti-
mación de la biomasa aérea mediante el uso de relaciones alométricas en seis especies arbóreas en
Oaxaca. México. Agrociencia, 36(6), 725–736.
Arreaga, G. W. E. (2002). Almacenamiento del carbono en bosques con manejo forestal sostenible en la
Reserva de la Biosfera Maya, Petén, Guatemala. Carbon storage in forests with sustainable manage-
ment in the Reserva de Biosfera Maya, Petén, Guatemala Mag. Sc. , CATIE, Turrialba (Costa Rica).
Balvanera, P., Kremer, C., & Martínez-Ramos, M. (2005). Appliying community structure analysis to
ecosystem function: Examples from pollination and carbon storage. Ecological Applications, 15(1),
360–375.
Blanco R., J. L. (2006). Erosión de la agrodiversidad en la milpa de los Zoque Popoluca de Soteapan:
Xutuchincon y Aktevet.
13
Brown, S., Gillespie, A. J. R., & Lugo, A. E. (1989). Biomass estimation methods for tropical forests
with applications to forest inventory data. Forest Science, 35(4), 881–902. https://doi.org/10.1093/
forestscience/35.4.881
Cairns, M. A., Olmsted, I., Granados, J., & Argaez, J. (2003). Composition and aboveground tree bio-
mass of a dry semi-evergreen forest on Mexico’s Yucatan Peninsula. Forest Ecology and Manage-
ment, 186(1), 125–132. https://doi.org/10.1016/S0378-1127(03)00229-9
Capra, F., & Sempau, D. (1998). La trama de la vida. Anagrama Barcelona.
Castillo-Capitán, G., Ávila-Bello, C. H., Lopez-Mata, L., & de León González, F. (2014). Structure and
tree diversity in traditional popoluca coffee agroecosystems in the Los Tuxtlas Biosphere Reserve.
Mexico. Interciencia, 39(9), 608–619.
Chave, J., Andalo, C., Brown, S., Cairns, M. A., Chambers, J. Q., Eamus, D., Fölster, H., Fromard, F., Higu-
chi, N., Kira, T., Lescure, J. O., Nelson, B. W., Ogawa, H., Puig, H., Riéra, B., & Yamakura, T. (2005).
Tree allometry and improved estimation of carbon stocks and balance in tropical forests. Oecologia,
145, 87–99. http://doi.org/10.1007/s00442-005-0100-x
Chiesura, A., & de Groot, R. (2003). Critical natural capital: A socio-cultural perspective. Ecological
Economics, 44, 219–231. https://doi.org/10.1016/S0921-8009(02)00275-6
Christmann, S. (2019). Do we realize the full impact of pollinator loss on other ecosystem services and
the challenges for any restoration in any terrestrial areas? Restoration Ecology, 27(4), 720–725.
Cole, T. G., & Ewel, J. J. (2006). Allometric equations for four valuable tropical tree species. Forest
Ecology and Management, 229(1), 351–360. https://doi.org/10.1016/j.foreco.2006.04.017
Cordero, J. (2003). Árboles de Centroamérica: un manual para extensionistas. Bib. Orton IICA/CATIE.
Cuevas, L. Z., López, N. M. N., Guzmán, R. C., Arias, J. G. M., Rodríguez, E. V. S., Hernández, L. G.,
& Magallanes, J. A. S. (2022). Fenología reproductiva de árboles en una zona montañosa del occi-
dente de México. Revista Mexicana De Ciencias Forestales, 13(69), 155–176. https://doi.org/10.
29298/r mcf.v13i69.1176
Dixon, R. K. (1995). Agroforestry systems: Sources of sinks of greenhouse gases? Agroforestry Systems,
31(2), 99–116. https://doi.org/10.1007/BF00711719
Douterlungne, D., Herrera-Gorocica, A. M., Ferguson, B. G., Siddique, I., & Soto-Pinto, L. (2013).
Ecuaciones alométricas para estimar biomasa y carbono de cuatro especies leñosas neotropicales
con potencial para la restauración. Agrociencia, 47(4), 385–397.
Ehrenbergerová, L. (2016). Carbon stock in agroforestry coffee plantations with different shade trees in
Villa Rica. Peru. Agroforestry Systems, 90(3), 433–445. https://doi.org/10.1007/s10457-015-9865-z
Espinoza-Domínguez, W., Krishnamurthy, L., Vázquez-Alarcón, A., & Torres-Rivera, A. (2012).
Almacén de carbono en sistemas agroforestales con café. Revista Chapingo Serie Ciencias
Forestales y Del Ambiente, 18(1), 57–70.
Florez, J. A., Muschler, R., Harvey, C. A., Finegan, B., & Roubik, D. W. (2002). Biodiversidad funcional
en cafetales: El rol de la diversidad vegetal en la conservación de abejas. Agroforestería En Las
Américas, 9(35–36), 29–36.
Fonseca, W., Alice, F., & Rey, J. M. (2009). Modelos para estimar la biomasa de especies nativas en
plantaciones y bosques secundarios en la zona Caribe de Costa Rica. Bosque, 30(1), 36–47.
García, R. (2006). Sistemas Complejos. Conceptos, Métodos y Fundamentación Epistemológica de la
Investigación Interdisciplinaria. Gedisa.
Gibbs, H. K., Brown, S., Niles, J. O., & Foley, J. A. (2007). Monitoring and estimating tropical forest
carbon stocks: Making REDD a reality. Environmental Research Letters, 2(4), 045023. https://doi.
org/10.1088/1748-9326/2/4/045023
Gómez-Castro, H., Pinto-Ruiz, R., Guevara-Hernández, F., & Gonzalez-Reyna, A. (2010). Estimaciones
de biomasa aérea y carbono almacenado en Gliricidia sepium (Lam.) y Leucaena leucocephala
(Jacq.) y su aplicación en sistemas silvopastoriles. ITEA, 106(4), 256–270.
González, Z. M. (2008). Estimación de la biomasa aérea y la captura de carbono en regeneración natural
de Pinus maximinoi H E Moore, Pinus oocarpa var. ochoterenai Mtz, y Quercus sp. en el norte del
estado de Chiapas, México Centro Agronómico Tropical de investigación y Enseñanza. Costa Rica.
Gutiérrez-García, G., & Ricker, M. (2011). Climate and climate change in the region of Los Tuxtlas
(Veracruz, Mexico): A statistical analysis. Atmósfera, 24(4), 347–373.
Häger, A. (2012). The effects of management and plant diversity on carbon storage in coffee agro-
forestry systems in Costa Rica. Agroforestry Systems, 86(2), 159–174. https://doi.org/10.1007/
s10457-012-9545-1
Haggar, J., Schibli, C., & Staver, C. (2001). ¿Cómo manejar árboles de sombra en cafetales. Agroforest-
ería En Las Américas, 8(29), 37–41.
Harterreiten-Souza, E. S., Brum Togni, P. H., Soares Pires, S., & Ryoiti Sujii, E. (2014). The role of
integrating agroforestry and vegetable planting in structuring communities of herbivorous insects
and their natural enemies in the Neotropical region. Agroforestry Systems, 88, 205–219.
13
Hergoualc’h, K., Blanchart, E., Skiba, U., Hénault, C., & Harmand, J.-M. (2012). Changes in carbon
stock and greenhouse gas balance in a coffee (Coffea arabica) monoculture versus an agroforestry
system with Inga densiflora, in Costa Rica. Agriculture, Ecosystems & Environment, 148, 102–110.
https://doi.org/10.1016/j.agee.2011.11.018
Hughes, R. F., Kauffman, J. B., & Jaramillo, V. J. (1999). Biomass, carbon, and nutrient dynamics of
secondary forests in a humid tropical region of Mexico. Ecology, 80(6), 1892–1907. https://doi.org/
10.1890/0012-9658(1999)080[1892:BCANDO]2.0.CO;2
Krakuer, C. D. (2019). Complexity: worlds hidden in plain sight. In D. C. Krakauer (Ed.), Worlds hid-
den in plain sigth. The evolving idea of complexity in the Santa Fe Institute (pp. 229–233). Santa Fe
Institute Press.
Leonti, M. (2002). Moko/La Rosa Negra. México ETH Zurich: Ethnobotany of the Popoluca Veracruz.
Levy-Tacher, S., & Aguirre Rivera, J. R. (2005). Succesional pathways derived from different vegetation
use patterns by Lacandon Mayan Indians. Journal of Sustainable Agriculture, 26(1), 49–79.
Levy-Tacher, S., Aguirre Rivera, J. R., Martínez Romero, M. M., & Durán Fernández, A. (2002). Carac-
terización del uso tradicional de la flora espontanea en la comunidad lacandona de Lacanhá, Chia-
pas. México. Interciencia, 27(9), 1–9.
Lindenmayer, D. B., & Franklin, J. F. (2002). Conserving forest biodiversity: a comprehensive multi-
scaled approach. Island press.
Llamas-Guzmán, L. P., Lazos Chavero, E., Perales Rivera, H. R., & Casas, A. (2022). Seed exchange
networks of Native Maize, Beans, and Squash in San Juan Ixtenco and San Luis Huamantla, Tlax-
cala, Mexico. Sustainability, 14(7), 3779. https://doi.org/10.3390/su14073779
Loranger, G., Ponge, J.-F., Imbert, D., & Lavelle, P. (2002). Leaf decomposition in two semi-evergreen
tropical forests: Influence of litter quality. Biology and Fertility of Soils, 35, 247–252.
Martínez Ramos, M. (1985). Claros, ciclos vitales de los árboles tropicales y regeneración natural de
las selvas altas perennifolias. In A. Gómez-Pompa & S. R. del Amo (Eds.), Investigación sobre la
regeneración de selvas altas en Veracruz, México (pp. 191–239). Alhambra: INIREB.
Medel-Ramírez, C., & Medel-López, H. (2019). Nota metodológica para la estimación del índice de
pobreza multidimensional en el Estado de Veracruz 2010 (Methodological Note for the Estimation
of Multidimensional Poverty Index in the State of Veracruz 2010). Available at SSRN 3493873.
Mesquita, R. C. G., Workman, S. W., & Neely, C. L. (1998). Slow litter decomposition in a Cecropia-
dominated secondary forest of Central Amazonia. Soil Biology and Biochemistry, 30(2), 167–175.
Miranda, F. (1961). Tres estudios botánicos en la selva Lacandona, Chiapas, México. Boletín de la
Sociedad Botánica De México, 26, 133–176.
Miranda, F., & Hernández-X, E. (2014). Los tipos de vegetación de México y su clasificación. Fondo de
Cultura Económica: Sociedad Botánica de México. CONABIO.
Moguel, P. & V.M. Toledo. (1999). Biodiversity conservation in traditional coffee systems of Mexico.
Conservation Biology, 12, 1–11. http://doi.org/10.1046/j.1523-1739.1999.97153.x
Montoy, L. (2010). Estudio apibotánico para un mejor aprovechamiento de los recursos naturales en la
región de las montañas, Veracruz. Campeche: Tesis de grado para Ingeniero Agrónomo. Instituto
Tecnológico de Chiná.
Montoya-Pfeiffer, P. M., León-Bonilla, D., & Nates-Parra, G. (2014). Catálogo de polen en mieles
de Apis mellifera provenientes de zonas cafeteras en la Sierra Nevada de Santa Marta, Magda-
lena, Colombia. Revista De La Academia Colombiana De Ciencias Exactas, Físicas y Naturales,
38(149), 364–384.
Navarro-Martínez, J., Godínez-Jaimes, F., López-López, M. Á., Rosas-Acevedo, J. L., Juárez-López,
A. L., & Reyes-Umaña, M. (2020). Ajuste de ecuaciones alométricas para estimar biomasa aérea
en Pinus oocarpa y Quercus resinosa en Guerrero México. Madera y Bosques. https://doi.org/10.
21829/myb.2020.2611964
Nicholls, C. I., & Altieri, M. A. (2013). Plant biodiversity enhances bees and other insect pollinators in
agroecosystems. A Review. Agronomy for Sustainable Development, 33(2), 257–274. https://doi.
org/10.1007/s13593-012-0092-y
Niles, J. O., Brown, S., Pretty, J., Ball, A. S., & Fay, J. (2002). Potential carbon mitigation and income in
developing countries from changes in use and management of agricultural and forest lands. Philo-
sophical Transactions of the Royal Society. London a., 360, 1621–1639.
Olabi, A. G., Obaideen, K., Elsaid, K., Wilberforce, T., Sayed, E. T., Maghrabie, H. M., & Abdelka-
reem, M. A. (2022). Assessment of the pre-combustion carbon capture contribution into sustainable
development goals SDGs using novel indicators. Renewable and Sustainable Energy Reviews, 153,
111710. https://doi.org/10.1016/j.rser.2021.111710
Otto, S. P., & Day, T. (2011). A biologist’s guide to mathematical modeling in ecology and evolution.
Princeton University Press.
13
Peeters, L. Y. K., Soto-Pinto, L., Perales, H., Montoya, G., & Ishiki, M. (2003). Coffee production, tim-
ber, and firewood in traditional and Inga-shaded plantations in Southern Mexico. Agriculture, Eco-
systems & Environment, 95(2), 481–493. https://doi.org/10.1016/S0167-8809(02)00204-9
Pérez-Grovas, G. (2000). Evaluación de la sustentabilidad del sistema de manejo de café orgánico en la Unión
de Ejidos Majomut, región de los Altos de Chiapas. O. Masera and S. López-Ridaura, eds pp 45–81.
Perfecto, I., & Vandermeer, J. (2015). Coffee agroecology: a new approach to understanding agricul-
tural biodiversity, ecosystem services and sustainable development. Routledge. https://doi.org/10.
4324/9780203526712
Philpott, S. M., Bichier, P., Rice, R., & Greenberg, R. (2007). Field-Testing Ecological and Economic
Benefits of Coffee Certification Programs [https://doi.org/10.1111/j.1523-1739.2007.00728.x].
Conservation Biology, 21(4), 975–985. https://doi.org/10.1111/j.1523-1739.2007.00728.x
Pinoargote, M., Cerda, R., Mercado, L., Aguilar, A., Barrios, M., & Somarriba, E. (2017). Carbon
stocks, net cash flow and family benefits from four small coffee plantation types in Nicaragua. For-
ests, Trees and Livelihoods, 26(3), 183–198. https://doi.org/10.1080/14728028.2016.1268544
Polanco-Echeverry, D. N., Álvarez-Salas, L. M. & Rios-Osorio, L. A. (2015). Proposed methodology for
research into the socioecological resilience of agroecosystems. Tropical and Subtropical Agroeco-
systems, 18(2), 207–219.
Puettmann, K. J., Coates, K. D., & Messier, Ch. (2009). A critique of silviculture. Managing for com-
plexity. Island Press.
Ramírez, W. M., & Ramírez, J. M. (2009). Municipios con mayor biodiversidad en Veracruz. Foresta
Veracruzana, 11(2), 43–50.
Ramírez-Arriaga, E., Navarro-Calvo, L. A., & Díaz-Carbajal, E. (2011). Botanical characterisation of
Mexican honeys from a subtropical region (Oaxaca) based on pollen analysis. Grana, 50(1), 40–54.
https://doi.org/10.1080/00173134.2010.537767
Ramírez-López, F., Hernández-Romero, Á. H., Ávila-Bello, C. H., & Retureta-Aponte, A. (2008).
Clasificación de suelos en el sistema Zoque-Popoluca en Soteapan, Veracruz, México. Sociedades
Rurales, Producción y Medio Ambiente, 8(16), 51–76.
Rice, R. A. (2011). Fruits from shade trees in coffee: How important are they? Agroforestry Systems,
83(1), 41–49. https://doi.org/10.1007/s10457-011-9385-4
Rodríguez, L. R., Jiménez, P. J., Aguirre, C. Ó. A., & Treviño, G. E. J. (2006). Estimación del carbono
almacenado en un bosque de niebla en Tamaulipas, México. Ciencia UANL, 9(2), 179–187.
Rodríguez, O. N., Cruz, R. M., Ramírez, T. J., Hernández, R. A. H., & Graillet, E. M. (2012). Diag-
nóstico de las organizaciones de producción agropecuaria en el municipio de Soteapan, Veracruz.
Ciencia Administrativa, 2, 1–11.
Rodríguez-Laguna, R. (2008). Carbono contenido en un bosque tropical subcaducifolio en la reserva de la
biosfera El Cielo, Tamaulipas. México. Revista Latinoamericana De Recursos Naturales, 4(2), 215–222.
Rojas-García, F., De Jong, B. H., Martínez-Zurimendí, P., & Paz-Pellat, F. (2015). Database of 478 allometric
equations to estimate biomass for Mexican trees and forests. Annals of Forest Science, 72(6), 835–864.
Roman, L., & Palma, J. (2007). Árboles y arbustos tropicales nativos productores de néctar y polen en el
estado de Colima. México. Avances En Investigación Agropecuaria, 11(3), 3–24.
Salinas-Castro, A., & Ávila-Bello, C. (2016). Traditional coffee agroecosystems in the Los Tuxtlas
Biosphere Reserve, Veracruz (Mexico): A refugee for ecologically important Coleoptera. Global
Advanced Research Journal of Agricultural Science, 5(6), 224–234.
Schroth, G., D’Angelo, S. A., Teixeira, W. G., Haag, D., & Lieberei, R. (2002). Conversion of secondary
forest into agroforestry and monoculture plantations in Amazonia: consequences for biomass, litter
and soil carbon stocks after 7 years. Forest Ecology and Management, 163(1), 131–150. https://doi.
org/10.1016/S0378-1127(01)00537-0
Segura, M., Kanninen, M., & Suárez, D. (2006). Allometric models for estimating aboveground biomass
of shade trees and coffee bushes grown together. Agroforestry Systems, 68(2), 143–150. https://doi.
org/10.1007/s10457-006-9005-x
Selaya, G. N., Oomen, R. J., Netten, J. J. C., Werger, M. J. A., & Anten, N. P. R. (2008). Biomass alloca-
tion and leaf life span in relation to light interception by tropical forest plants during the first years
of secondary succession. Journal of Ecology, 96(6), 1211–1221. https://doi.org/10.1111/j.1365-
2745.2008.01441.x
SEMARNAT. (2010). Norma Oficial Mexicana NOM-059-SEMARNAT-2010. Protección ambiental-
Especies nativas de México de flora y fauna silvestres-Categorías de riesgo y especificaciones para
su inclusión, exclusión o cambio-Lista de especies en riesgo. Diario Oficial, 30 de diciembre de 2010.
SMBD. (2004). Normas para la Producción, el Procesamiento y la Comercialización de Café “Bird
Friendly®”. Certificado Orgánico Bajo Sombra (p. 19). Washington, DC. USA: Smithsonian
Migratory Bird Center, National Zoo.
13
Soley, F. G., & Perfecto, I. (2021). A way forward for biodiversity conservation: High-quality landscapes.
Trends in Ecology & Evolution, 36(9), 770–773. https://doi.org/10.1016/j.tree.2021.04.012
Somarriba, E., Harvey, C. A., Samper, M., Anthony, F., González, J., Staver, C., & Rice, R. (2004). Biodi-
versity conservation in Neotropical coffee (Coffea arabica) plantations. Agroforestry and biodiversity
conservation in tropical landscapes, pp. 198–226.
Terán-Ramírez, M. A., Rodríguez-Ortiz, G., Enríquez-del Valle, J. R., & Velasco-Velasco, V. A. (2018).
Biomasa aérea y ecuaciones alométricas en un cafetal en la Sierra Norte de Oaxaca. Ecosistemas y
Recursos Agropecuarios, 5(14), 217–226. https://doi.org/10.19136/era.a5n14.1444
Valencia, V., García-Barrios, L., West, P., Sterling, E. J., & Naeem, S. (2014). The role of coffee agroforestry
in the conservation of tree diversity and community composition of native forests in a Biosphere Reserve.
Agriculture, Ecosystems & Environment, 189, 154–163. https://doi.org/10.1016/j.agee.2014.03.024
van Breugel, M., Ransijn, J., Craven, D., Bongers, F., & Hall, J. S. (2011). Estimating carbon stock in sec-
ondary forests: Decisions and uncertainties associated with allometric biomass models. Forest Ecology
and Management, 262(8), 1648–1657. https://doi.org/10.1016/j.foreco.2011.07.018
Vandermeer, J., & Perfecto, I. (2007). The Agricultural Matrix and a Future Paradigm for Conservation
[https://doi.org/10.1111/j.1523-1739.2006.00582.x]. Conservation biology, 21(1), 274–277. https://
doi.org/10.1111/j.1523-1739.2006.00582.x
Vandermeer, J., Perfecto, I., & Philpott, S. (2010). Ecological complexity and pest control in organic coffee
production: Uncovering an autonomous ecosystem service. BioScience, 60(7), 527–537. https://doi.
org/10.1525/bio.2010.60.7.8
Vázquez-Yanes, C., Batis-Muñoz, A., Alcocer-Silva, M., Gual-Díaz, M., & Sánchez-Dirzo, C. (1999). Árbo-
les y arbustos potencialmente valiosos para la restauración ecológica y la reforestación. Reporte Téc-
nico Del Proyecto J, 84, 201–204.
Vázquez-Yanez, C. (1998). Trema micrantha (L.) Blume (Ulmaceae): A promising neotropical tree for site
amelioration of deforest land. Agroforestry Systems, 40, 97–104.
Villegas-Durán, G., Bolaños-Medina, A., Miranda-Sánchez, J. A., & Zenón-Abarca, A. J. (2000). Flora nec-
tarífera y polinífera en el estado de Chiapas. COTECOCA- SAGAR.
Yépez, P. C. (2001). Selección de árboles para sombra en cafetales diversificados de Chiapas. Turrialba
(Costa Rica): Mag. Sc. CATIE.
Zahory, D. (2004). Unconscious selection and the evolution of domesticated plants. Economic Botany,
58(1), 5–10.
Publisher’s Note Springer Nature remains neutral with regard to jurisdictional claims in published maps and
institutional affiliations.
Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a
publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript
version of this article is solely governed by the terms of such publishing agreement and applicable law.
Authors and Affiliations
Carlos H. Ávila‑Bello1 · Ángel Héctor Hernández‑Romero1 ·

Dinora Vázquez‑Luna1 · Daniel Alejandro Lara‑Rodríguez1 ·
Adriana Martínez‑Jerónimo2 · Bitia N. Meneses‑García2 ·
Xochitl M. Sánchez‑Sandoval2
* Carlos H. Ávila‑Bello
cavilab2001@yahoo.com
1
Centro de Estudios Interdisciplinarios en Agrobiodiversidad, Universidad Veracruzana, Carretera
Costera del Golfo Km. 220. Col. Agrícola y Ganadera Michapan, 96100 Acayucan, Veracruz,
México
2
Facultad de Ingeniería en Sistemas de Producción Agropecuaria (FISPA), Universidad
Veracruzana, Carretera Costera del Golfo Km. 220. Col. Agrícola y Ganadera Michapan,
96100 Acayucan, Veracruz, México
13
Terms and Conditions
Springer Nature journal content, brought to you courtesy of Springer Nature Customer Service Center
GmbH (“Springer Nature”).
Springer Nature supports a reasonable amount of sharing of research papers by authors, subscribers
and authorised users (“Users”), for small-scale personal, non-commercial use provided that all
copyright, trade and service marks and other proprietary notices are maintained. By accessing,
sharing, receiving or otherwise using the Springer Nature journal content you agree to these terms of
use (“Terms”). For these purposes, Springer Nature considers academic use (by researchers and
students) to be non-commercial.
These Terms are supplementary and will apply in addition to any applicable website terms and
conditions, a relevant site licence or a personal subscription. These Terms will prevail over any
conflict or ambiguity with regards to the relevant terms, a site licence or a personal subscription (to
the extent of the conflict or ambiguity only). For Creative Commons-licensed articles, the terms of
the Creative Commons license used will apply.
We collect and use personal data to provide access to the Springer Nature journal content. We may
also use these personal data internally within ResearchGate and Springer Nature and as agreed share
it, in an anonymised way, for purposes of tracking, analysis and reporting. We will not otherwise
disclose your personal data outside the ResearchGate or the Springer Nature group of companies
unless we have your permission as detailed in the Privacy Policy.
While Users may use the Springer Nature journal content for small scale, personal non-commercial
use, it is important to note that Users may not:
1. use such content for the purpose of providing other users with access on a regular or large scale
basis or as a means to circumvent access control;
2. use such content where to do so would be considered a criminal or statutory offence in any
jurisdiction, or gives rise to civil liability, or is otherwise unlawful;
3. falsely or misleadingly imply or suggest endorsement, approval , sponsorship, or association
unless explicitly agreed to by Springer Nature in writing;
4. use bots or other automated methods to access the content or redirect messages
5. override any security feature or exclusionary protocol; or
6. share the content in order to create substitute for Springer Nature products or services or a
systematic database of Springer Nature journal content.
In line with the restriction against commercial use, Springer Nature does not permit the creation of a
product or service that creates revenue, royalties, rent or income from our content or its inclusion as
part of a paid for service or for other commercial gain. Springer Nature journal content cannot be
used for inter-library loans and librarians may not upload Springer Nature journal content on a large
scale into their, or any other, institutional repository.
These terms of use are reviewed regularly and may be amended at any time. Springer Nature is not
obligated to publish any information or content on this website and may remove it or features or
functionality at our sole discretion, at any time with or without notice. Springer Nature may revoke
this licence to you at any time and remove access to any copies of the Springer Nature journal content
which have been saved.
To the fullest extent permitted by law, Springer Nature makes no warranties, representations or
guarantees to Users, either express or implied with respect to the Springer nature journal content and
all parties disclaim and waive any implied warranties or warranties imposed by law, including
merchantability or fitness for any particular purpose.
Please note that these rights do not automatically extend to content, data or other material published
by Springer Nature that may be licensed from third parties.
If you would like to use or distribute our Springer Nature journal content to a wider audience or on a
regular basis or in any other manner not expressly permitted by these Terms, please contact Springer
Nature at
onlineservice@springernature.com

Design Complex - Systems Agrobiodiversity

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Design Complex - Systems Agrobiodiversity

Uploaded by

Copyright:

Available Formats

Environment, Development and Sustainability

Design of complex agroecosystems: traditional and formal

Carlos H. Ávila‑Bello1 · Ángel Héctor Hernández‑Romero1 ·

Received: 11 May 2022 / Accepted: 1 February 2023

Keywords Biocultural diversity · Complex systems · Indigenous seeds · Sustainability

2 Materials and methods

2.2 Diagnosis of the coffee agroecosystems

2.3 Estimation of carbon sequestration

Y = (exp(-3.78 + (0.95*log(DBH) + log(H)) Hughes et al. (1999)

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Y = exp(− 1.46 + 1.69 ln (DBH)) Gómez-Castro et al. (2010)

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

2.4 Species valuation by peasants

3.1 General characteristics of the agroecosystems

3.2 Carbon sequestration by stratum and species

3.3 Valuation of tree species

4.1 Agroecosystems, their management and value for peasants

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

*Species that can be deciduous or evergreen according to climate conditions

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

4.2 Valuation of tree species and C sequestration

Content courtesy of Springer Nature, terms of use apply. Rights reserved.

4.3 Other functions of trees for agroecosystem design: melliferous potential

Authors and Affiliations

Carlos H. Ávila‑Bello1 · Ángel Héctor Hernández‑Romero1 ·

You might also like

2 Materials and methods

2.2 Diagnosis of the coffee agroecosystems

2.3 Estimation of carbon sequestration

2.4 Species valuation by peasants

3.1 General characteristics of the agroecosystems

3.2 Carbon sequestration by stratum and species

3.3 Valuation of tree species

4.1 Agroecosystems, their management and value for peasants

4.2 Valuation of tree species and C sequestration

4.3 Other functions of trees for agroecosystem design: melliferous potential