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Article

Importance of Traditional Vanilla Cultivation in the Conservation of Plant Diversity in Tropical Forests in Northern Veracruz, Mexico

by
Noé Velázquez-Rosas
1,*,
Santiago Sinaca Colin
1,
Guillermo Vázquez-Domínguez
1,2,
Abril Velasco-Murguía
1,
Evodia Silva Rivera
1,
Betsabé Ruiz-Guerra
3,
Fabio Levi Friedrich
4,
Rosenda Cortés Galindo
5,
Samaria Armenta-Montero
1 and
Rodolfo Martínez-Mota
1,*
1
Centro de Investigaciones Tropicales, Universidad Veracruzana, Xalapa 91000, Mexico
2
Facultad de Ciencias Químicas, Universidad Veracruzana, Xalapa 91000, Mexico
3
Red de Interacciones Multitróficas, Instituto de Ecología A.C., Xalapa 91073, Mexico
4
Institute of Ecology, University of Berlin, 10623 Berlin, Germany
5
Coordinación Territorial, Subsecretaria de Inclusión Productiva y Desarrollo Rural, Secretaria de Bienestar, Mexico City 06600, Mexico
*
Authors to whom correspondence should be addressed.
Sustainability 2025, 17(6), 2598; https://doi.org/10.3390/su17062598
Submission received: 29 January 2025 / Revised: 3 March 2025 / Accepted: 13 March 2025 / Published: 15 March 2025
(This article belongs to the Section Sustainability, Biodiversity and Conservation)
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Abstract

The significance of traditional agroforestry systems in preserving and enhancing tropical forest biodiversity in landscapes dominated by human activities has recently been recognized. We assessed the role of traditional vanilla cultivation on sustaining plant diversity in the tropical forests of northern Veracruz, Mexico. We analyzed the composition, alpha (Shannon and Simpson exponential) and beta diversity, the structure (stem density and basal area) and types of regeneration of woody plants across different vanilla production systems, including traditional vanilla plots, the agroforestry production of vanilla, the citrus–vanilla system, and fragments of tropical rain forest. Our findings revealed that traditional vanilla plots preserve 67% of the woody plants’ richness, with an alpha diversity similar to that of the forest fragments. The similarity between vanilla production systems and vegetation fragments was less than 30%. Traditional vanilla plots accounted for 34% of the basal area and had a stem density similar to that of the forest, while retaining 25% of shade-tolerant species. These results suggest that traditional vanilla plots are key landscape elements for conserving plant diversity and supporting the ecological functions of tropical forests.

1. Introduction

Biodiversity conservation is a priority action to safeguard our planet and humanity [1] and represents one of the primary current challenges for civilization. In tropical regions, rain forests have been extensively altered; it is estimated that more than 70% of their global extent has been transformed and they are currently immersed in a matrix composed of forest remnants, agricultural fields, and secondary forests in different states of conservation [2,3]. Although research efforts and actions to expand protected natural areas have increased, they are unfortunately insufficient to conserve the biodiversity of these ecosystems due to the intensity and magnitude of human disturbances [4,5,6,7]. Therefore, a paradigm shift in conservation has been proposed, dating back some decades, to consider the conservation potential of current landscapes that have been largely modified but that preserve elements of biodiversity [8,9]. This includes the consideration of cultural landscapes generated by indigenous peoples as a conservation unit [10]. The increase in indigenous managed territories has demonstrated their importance in biodiversity conservation [10,11,12]. Within these territories, agroforestry systems are one of the elements that have received most attention [4,13]. These systems play a key role in the conservation of biodiversity in tropical ecosystems because (i) they safeguard a significant amount of the diversity of adjacent tropical ecosystems, providing multiple benefits to human populations; (ii) at regional scales, for example, at the watershed level, they have a high heterogeneity of plant communities, due to the different management practices of local people; (iii) they maintain vital ecological processes such as water cycling and soil biotic integrity, among others; and (iv) the forest resources (timber, edible, firewood, etc.) provided by agroforestry systems can help reduce extraction pressures on protected natural areas [4,13]. Therefore, these systems can potentially contribute to the design of public policies for landscape sustainable management and biodiversity conservation [10,11].
In Mexico, a wide range of traditional agroforestry systems are associated with the main indigenous cultures of the country [13,14]. These systems integrate worldviews, knowledge, practices, and social norms of use. Their ecological importance lies in their key role in conserving native, endemic, and bioculturally important species; they also represent spaces for technical innovation in the management and domestication of species and landscapes [13]. One of the agroforestry systems that have received little attention, despite their great cultural and economic importance, is traditional vanilla cultivation (TV). Vanilla (Vanilla planifolia Andrews) is the second most important aromatic species in the world, mainly used in the food and cosmetics industry [15]. Although vanilla is now cultivated in several countries around the world, historical records indicate that vanilla was domesticated by the Totonacan people and is part of the biocultural heritage of the Totonacapan region [16]. At the beginning of the 20th century, the region had an extensive forest cover integrated with agricultural areas producing corn and beans, vanilla, tobacco, and rubber [17]. However, from 1940 onwards, new crops (coffee, citrus, and banana) were introduced and extensive cattle ranching expanded, which led to 500,000 ha of tropical forests being deforested and transformed into agricultural areas and cattle pastures [10,18]. However, TVs remain an essential element of the Totonac cultural landscape, and their management is associated with ecological succession dynamics, since vanilla is traditionally grown in secondary forests [10,17].
Currently, vanilla production has been modified and is preferably grown in agroforestry systems with a reduced number of “tutor” host trees, associated with citrus crops, in shade house systems, and, to a lesser extent, in TVs. Despite this, traditional vanilla production persists and recently many Totonacapan producers are interested in returning to traditional production practices, motivated by the problems facing this crop due to extreme weather phenomena (increased rainfall and longer dry season) caused by climate change, as well as the increase in crop diseases. The revitalization of this traditional agroforestry ecosystem would favor the preservation of Totonac knowledge and traditions, which will allow the maintenance and management of more resilient agroecosystems and the conservation of the biocultural diversity of this region. In this context, our objective was to evaluate the importance of traditional vanilla plots in the conservation of plant diversity in tropical forests in northern Veracruz, one of the most deforested areas of Mexico, as well as to analyze their relevance for the restoration and conservation of plant and biocultural diversity in the region.

2. Materials and Methods

2.1. Study Area

The study was carried out in the Totonacapan indigenous region, which is a historical and cultural unit that began its configuration 2500 years ago. It is located between the high elevation zone of the Sierra Madre Oriental and the coastal region of the Gulf of Mexico, including the central-northern zone of the state of Veracruz, the Sierra Norte of Puebla, and the municipality of Acaxochitlán, Hidalgo [19,20]. In the coastal plain of Totonacapan Veracruz, sampling sites were located in three municipalities: Papantla de Olarte, Gutiérrez Zamora, and Cazones. The climate is of the warm humid, subhumid type with rains in summer and autumn. The average annual temperature and precipitation are 24 °C and 1160.4 mm, respectively [21]. Northern Veracruz state is one of the most deforested regions in the country; more than 90% of the tropical forests have been transformed into cattle and oil exploitation zones, where citrus, corn, and cattle pastures predominate [22]. The vegetation is represented by fragments of tropical rain forest in different states of conservation [22,23].

2.2. Characterization of Vegetation in Elements of the Totonac Biocultural Landscape

In total, 35 study sites were selected for vegetation sampling, 25 of which correspond to vanilla production systems: 11 TV, 5 agroforestry systems (AF), and 9 vanilla systems associated with citrus (CV; Figure 1). In addition, five fragments of medium semi-deciduous forest (F) and five fragments of secondary forest (SF) derived from the same type of vegetation were considered. At each site, consent was obtained from the producers and landowners, explaining the research activities and the objectives of the project. The selection criteria for the sites were as follows: in the case of the F sites, the communities should conserve similar structural characteristics (a tree height greater than 20 m and the presence of trees of diameter classes greater than 30 cm), the primary species characteristic of the region should be frequent, and there should be no evidence of recent extraction or logging. The selected SF sites had at least 10 years without timber extraction or logging activities, a tree height greater than 10 m, dominant tree diameter classes less than 10 cm, and an abundance of secondary species. In both cases, it was necessary to ensure that sampling units could be placed within the site area.
The vanilla production systems were categorized into three groups according to the criteria proposed by Castelan Culebro [24]. TV systems are established mainly in secondary forests, where young and adult trees are used as vanilla tutors, and there is generally a great variety of primary and secondary species. Management practices are mainly pruning of the canopy to regulate shade and humidity, allowing the development of the vanilla, as well as managing leaf litter to form mulch around the vanilla tutors. In the CV production system, orange, mandarin, or lemon trees are used as tutors. Management practices in these sites commonly include cutting weeds to make compost around the tree trunks to conserve moisture and provide nutrition for the vanilla plants. In the AF system, stakes are planted, and a reduced number of preferably fast-growing species are used (Erythrina sp., Gliricidia sepium (Jacq.) Kunth, Damburneya salicifolia (Kunth) Trofimov & Rohwer, and Hammelia patens Jacq., among others). In some cases, irrigation systems are introduced, and leaf litter is managed.
In each of the vanilla production types, a 500 m2 circle was placed in the center of the plot. This sample size was selected because the extent of traditional parcels often does not exceed 1000 m2. In the case of forest and secondary forest fragments, circles of 1000 m2 were placed in the center of the fragment; in some cases where the fragments had a reduced extent, the circles were 500 m2. In each circle, the stems of all woody plants (trees and shrubs) with a diameter at breast height (DBH) > 2.5 cm were measured and botanical samples were taken for taxonomic determination. The specimens collected were herborized and determined in the field and with taxonomic keys and the support of specialists. The specimens were compared with the specimens in the XAL Herbarium of the Instituto de Ecología A. C., Xalapa, Mexico. All vegetative and fertile specimens were stored at the Tropical Research Center of the Universidad Veracruzana, Xalapa, Mexico.
To evaluate the structure of each site, stem density and basal area were estimated according to Mostacedo and Fredericksen [25]; both variables were standardized to one hectare. All species recorded were classified by their regeneration type considering the criteria of Hill and Curran [26] and Martínez Ramos [27] as light-demanding species (pioneers), generalists (intermediate successional species), and shade-tolerant species. The frequency of each type of regeneration per site was estimated by considering the sum of the relative density of all species belonging to each group.

2.3. Statistical Analysis

To analyze the alpha diversity among vegetation fragments and vanilla production systems, rarefaction and extrapolation curves were created with the iNext Online package, version (30 March 2024) [28,29], based on Hill numbers (q0 = richness; q1 = Shannon’s exponential index; q2 = Simpson’s inverse index). To carry out beta diversity analysis, using the abundance data for each species at each site, a matrix of 35 sites × 178 species was constructed. The data from this matrix were square-root-transformed to generate an initial triangular resemblance matrix using Bray–Curtis similarity. Clustering was achieved through group averaging, and the similarity profile was tested using similarity profile analysis (SIMPROF) [30] and permutation tests (999 iterations). SIMPROF tests the statistical significance of every node within a dendrogram, starting from the top (all points within a single group) and highlighting only those groups that show within-group multivariate structure. Non-metric multidimensional scaling (NMDS) [31,32] was used to generate a 2D model of plot dispersion to test group affiliation created by SIMPROF, assess multidimensional stress levels, and evaluate the effects of outliers. Similarity percentage analysis (SIMPER) [33] determines the contribution of each species to the differences in community composition between groups. SIMPER employs the Bray–Curtis similarity measure to identify positive and negative diagnostic species across vegetation types. Plant species with a combination of high frequency and abundance were also identified and listed for diagnostic purposes and type delineation.
The average values of density and basal area between vegetation fragments and vanilla production systems were compared with an analysis of variance, the assumptions of homogeneity of variance and normality were tested with the Levene and Shapiro–Wilk tests, respectively, and Tukey’s post hoc test was used. The frequency of regeneration types was compared through generalized linear models, using a Poisson distribution and Log link function; when significant differences were recorded, a Bonferroni post hoc test was performed in the Jamovi program, version 1.1.9.0.

3. Results

3.1. Richness and Diversity of Woody Plants

A total of 178 species were recorded across vanilla production systems, forest fragments, and secondary forests, and were grouped into 42 families and 125 genera (Table A1). The families with the highest number of species were Fabaceae (n = 29), Rubiacae (n = 15), Euphorbiaceae (n = 10), and Myrtaceae (n = 9). The highest number of species was recorded in F (n = 115), followed by SF (n = 82) and TV (n = 78). In contrast, the lowest richness was recorded in AF (n = 4) and CV (n = 21).
Specifically, the richness estimators used revealed that the sampling coverage for all vanilla production systems, forest fragments, and secondary forests was higher than 96%, with the highest values found in the forest fragments and traditional vanilla plots. The highest values of diversity q1 and q2 (Shannon and Simpson exponential, respectively) were recorded in the forest fragments, followed by traditional vanilla plots and secondary forests (Table 1). The beta diversity analysis showed that there is a high turnover of species within and between vanilla production systems and vegetation fragments (forest and secondary forests). The groups were integrated into the cluster (SIMPROF; Pi = 5.45; p = 0.01; number of permuted statistics greater than or equal to Pi = 0) and NMDS (stress of 0.16).
Vegetation fragments and the different vanilla production systems were grouped except for the CV system, which was clustered almost independently as a single group (Figure 2). These groups were arranged in a gradient from communities with the highest richness and density of individuals (groups C, B, and A) to less diverse and less dense communities (groups F, E, and D). The similarity between groups was below 19.0% (Table 2), with the highest values among groups C and B, which mainly integrated the fragment communities of F, SF, and TV. The lowest similarities were recorded between group C and D; the latter group, which integrated almost all the communities of the CV system, presented the lowest similarity in relation to the majority of the groups (Table 2).
The importance of species to the integration of each group, in terms of abundance, was differential. In the case of group C (which integrated forest and secondary forest communities, mainly), the most important species were Brosimum alicastrum Sw., Bursera simaruba (L.) Sarg., D. salicifolia, Tabernaemontana alba Mill., Protium copal (Schltdl. & Cham.) Engl., and Aphananthe monoica (Hemsl.) J.F. Leroy, which contributed 47% of the density of individuals in this group. In group B (composed of secondary forest communities and traditional vanilla plots), B. simaruba, D. salicifolia, and T. alba were the most relevant species, comprising together 52% of the abundance. In the remaining groups, the number of species containing more than 50% of the abundance was lower. In group A, represented by traditional vanilla plots and secondary forest, three species contributed 65% of the abundance, including B. simaruba, Nectandra rubriflora (Mez) C.K. Allen, and Guazuma ulmifolia Lam. In group F, Erythrina americana Mill., G. sepium, and Cedrela odorata L. contributed 94%; in group E, Citrus × aurantium L. and H. patens contributed 66%; and in group D, C. × auratium contributed 91% of the abundance.
When analyzing the number of unique species between vegetation fragments and vanilla production systems (Figure 3), 51 species were recorded in F. This value was three times higher compared to SF and TV, and between 10 and 25 times higher compared to AF and CV, respectively.

3.2. Structural Characteristics of Vegetation Fragments and Vanilla Cultivation Systems

Structurally, changes were recorded between vegetation fragments and vanilla production systems (Figure 4). The average basal area of the vegetation fragments (forest and secondary forest) were significantly higher compared to the three vanilla production systems (F(4, 28) = 56.59; p = 0.001), which had between 34 and 50% of the basal area of forests and secondary forests, respectively (Figure 4a). In terms of stem density, the vegetation fragments had a similar average density with two production systems (TV and AF), while the CV system had significantly lower values compared to F, SF, and AF (F(4, 28) = 5.50; p = 0.002; Figure 4b).
In vanilla production systems and secondary forest fragments, light-demanding species were significantly more frequent (χ2 = 682; df = 8; p < 0.01; Figure 5); in contrast, shade-tolerant species were significantly more frequent in forest fragments. There was a tendency for light-demanding species to increase from conserved forests to secondary forests and vanilla production systems, with the highest values in AF and CV. Conversely, the presence of shade-tolerant and shade-intolerant species decreased in the same direction (Figure 5).

4. Discussion

In this study, we evaluated the role of different vanilla cultivation systems in the conservation of plant diversity in Totonacapan, a bioculturally relevant region of Mexico, which has been highly fragmented by anthropogenic activities. Traditional vanilla plots were shown to be a relevant landscape element in the conservation of plant diversity, compared to the other two vanilla production systems evaluated: agroforestry and citrus with vanilla. In addition, its structure may favor the maintenance of ecological functions such as carbon sequestration. These results are consistent with the idea that agroforestry systems are key to the conservation of plant diversity in tropical forests in sites dominated by human activities and can play a relevant role in maintaining the ecological functions of tropical forests [9,14,34].
The TVs studied harbor two-thirds of the woody species richness recorded in the forest fragments, which is similar to the results reported in vanilla eco-plantations in forests in Madagascar [35]. For other agroforestry systems, the percentages vary between 35 and 68% [4,14,36]. In traditional agroforestry systems, high plant richness has been reported, similar to that of conserved forests in the humid and dry tropics [14,34,35], which is promoted by the different goods that the plants provide to the population [10,13,14,36]. In the TVs of Papantla, the farmers use between 70 and 100% of the tree and shrub species for different uses: as shade and vanilla tutors, firewood, food, for medicinal purposes, in construction, etc. [37]. Traditional vanilla plantations promote plant richness and diversity due to four main aspects: (i) these agrosystems maintain part of the structure and composition of the preserved forests, and the medium canopy trees are used for shade and tutors for the vanilla plants [20,37]; (ii) they promote species diversification using useful plants; (iii) these systems are free of agrochemicals, which help facilitate the regeneration of plant species; and (iv) a significant factor in conserving plant diversity in traditional vanilla plantations is the strong connection producers still maintain with nature through their belief system. For example, local people protect trees and the forest in reverence of Kwikgolo (“old tree” in Totonac), a deity that protects the forest and provides goods to the villagers [38].
The composition and structure of the woody plant communities hosted by the vanilla production systems differed from the vegetation fragments. The results of the beta diversity analysis showed that, among the different groups composed of vanilla production systems and vegetation fragments, there is a low similarity (<20%), i.e., each group harbors a specific composition and unique species that do not occur in any other system. The changes in the composition of TVs are linked with the producers’ traditional knowledge and practices. Based on their experience, skills, and needs, they make decisions in the design of TVs considering the local environmental conditions. This approach promotes a heterogeneous use of the territory, leading to an increase in the plant β-diversity. For example, forest fragments showed the highest richness and diversity, in addition to the highest frequency of unique species recorded, where 31% of these were represented by one individual, a pattern of rarity similar to that reported in other tropical forests in Mexico [39]. This reveals that the conservation of woody plant diversity in the study area depends on the preservation of as many landscape elements as possible to maintain environmental heterogeneity and plant diversity, where vegetation fragments (conserved and secondary forests) and traditional agroforestry systems are essential [14,34,35].
In TV, there was a low similarity between the different plots evaluated (30%), which is a result of differing management by the farmers in this agroforest system. The owners of the plots make decisions in the design, the selection of species (e.g., use of shade plants, tutors, edibles, and firewood), and the organization of the vanilla cultivation system, according to their needs and knowledge, resulting in the diversification of species used in these systems [10,37,38]. Conversely, in the more intensive vanilla production systems (AFS and CV), the diversity of other useful species is reduced, because priority is given to the intensification of vanilla production.
In terms of structure, TV and AFS presented a stem density similar to that of the vegetation fragments; this tendency has been recorded in other agroforestry systems [14,34]. However, when the basal area was analyzed, the vegetation fragments had higher values, three and two times higher compared to vanilla production systems. This is because, in conserved and secondary forests, the abundance of trees with diameter classes > 30 cm is higher [40]; this pattern was recorded in the vegetation fragments studied. In contrast, in vanilla production systems, the trees used as tutors and for shade are mainly of diameter classes less than 10 cm DBH.
This structural change has important implications for the ecosystem functions of vanilla cultivation systems and their benefits to people, for example, the ability of agroforestry systems to function as carbon sinks [41,42]. In this sense, the carbon sequestration capacity of agroforestry systems depends, among other factors, on environmental conditions and management [41]. In our study, although TVs present a basal area similar to that of the other two vanilla production systems (AFS and CV), they could have a higher carbon sequestration capacity because they have a greater abundance of shade-tolerant species. Generally, these species are slow-growing and have a high wood density, which is associated with a higher carbon storage potential [41]. In practical terms, carbon sequestration could be another source of economic resources, through the payment of carbon credits [42], for farmers in the Totonacapan region to encourage vanilla production in a traditional agroforestry system.
Anthropic pressures on tropical forests have led to a decrease in their richness and diversity, the homogenization of their composition, and changes in their functions [36]. One of the trends recorded in tropical forests is that as deforestation and fragmentation increase, the dominance of light-demanding native plants increases [2,39,43,44]. In our study, we recorded that the dominance of light-demanding species increases in a gradient from vegetation fragments (forest and secondary forest) to more intensive vanilla production systems (CV). In the forest fragments of this study, shade-tolerant species were dominant; this pattern is similar to that recorded in other tropical forests of Mexico [2,39]; on the other hand, in the secondary forest fragments and vanilla production systems, light-demanding species were dominant. In particular, in the CV system, C. × auratium contributes 91% of the abundance of stems, which is characteristic of intensive agricultural production systems in the tropics, where conditions are unfavorable for the development of species with other types of regeneration [44]. In TV, shade-tolerant species have a relative density that represents 50% of that recorded in the forest fragments. These species typically face dispersal limitations and environmental restrictions in disturbed sites [2,44]. The high presence of shade-tolerant species in traditional vanilla plantations suggests that these agrosystems, together with secondary forest fragments, offer conditions that promote the establishment and growth of such species. Additionally, they serve as sites for germplasm conservation and a source of propagules for tolerant species. Increasing traditional vanilla plots and maintaining secondary forests could favor succession processes and the reestablishment of tropical forests in the Totonocapan region.
The role of TV in the conservation of the plant diversity of the tropical forests of Totonocapan must be analyzed from a landscape perspective [4,14,44]. The conservation of the region’s plant diversity depends largely on the preservation of the remaining vegetation fragments in a highly transformed habitat mosaic. These areas, along with secondary forests, support the largest number of tropical forest species. However, the traditional vanilla plots in the municipality of Papantla are an alternative that contributes to the sustainable management of vanilla, the supply of useful plants (providing, e.g., food and firewood), and the maintenance of plant diversity. This farming system is a central element of the Totonac cultural landscape (sensu Toledo et al. [10]), in which the management of these systems was integrated with the successional processes of the forests, resulting in a heterogeneous landscape configuration that preserves significant biocultural diversity (more than 350 useful species were managed in these systems; [10]).
Currently, extensive citrus production is promoting the deforestation of tropical forest fragments, the simplification of the floristic structure and composition, and the elimination of traditional agrosystems in Totonacapan. Therefore, the promotion of vanilla production with the traditional system and the preservation of and increase in vegetation fragments under the traditional Totonac landscape management and planning scheme represent an opportunity for the conservation of the plant diversity of tropical forests in this region. Other studies have shown that the protection and growth of regions managed by indigenous peoples, based on their traditional ecological knowledge, worldview, and practices, have favored the conservation of the diversity of tropical forests in other regions of the world [6,11,12].
The promotion and growth of agroforestry systems that favor biodiversity conservation and the improvement of the economic conditions of farmers is one of the great challenges to be resolved in the tropical region [45]. In the Papantla area, vanilla production under the traditional scheme has fallen into disuse for different reasons, including low production, substitution by more intensive and technical systems, and the loss of traditional ecological knowledge associated with these vanilla cultivation systems [38]. However, producers have recently valued the importance of revitalizing the knowledge of traditional management because they have verified that these systems are more resilient to the environmental problems of the region, associated with climate change, for example, extreme droughts, the increased frequency of extreme weather events, pests, and diseases [46,47]. In addition, the production under this scheme is of higher quality in terms of the size and concentration of vanilla in the fruits [24]. In international markets, there is a high commercial demand for natural vanilla; therefore, the promotion of traditional production systems could favor more economically favorable marketing schemes (access to organic markets, gourmets, and fairer prices) and payments for carbon credits, as well as achieving environmental benefits that favor the well-being of peasant populations.
Our results reinforce the idea that traditional agroforestry systems and the associated ecological knowledge should be considered in the design of coherent public policies for sustainable management and nature conservation, in accordance with Mexico’s biocultural reality [13]. In recent years, the Mexican government has implemented the ‘Sembrando Vida’ program, which aims to contribute to the social welfare of rural sectors and to recover forest cover through the development of agroforestry systems. However, this program has in many cases ignored indigenous peoples’ practices, knowledge, and forms of governance [48,49], which have been shown to be consistent with sustainable development [10,13,14]. In the study area, the Totonacapan region, TVs can not only favor the conservation of plant diversity and the restoration of ecosystem processes in tropical forests, but they can also play a key role by consolidating vanilla production under sustainable management schemes that will improve the economy and the quality of life of local populations in a highly degraded and overexploited Mexican region that paradoxically shelters cultural and biological diversity essential for people’s well-being.

5. Conclusions

The traditional vanilla plots of Totonacapan protect the richness, diversity, and structure of woody plants that can favor the conservation of plant diversity and the restoration of the ecological processes of the tropical forests in the northern region of the state of Veracruz, which is one of the most deforested zones in Mexico. In the Totonacapan region of Veracruz, traditional ecological knowledge still persists that can be very relevant to restoring the Totonac cultural landscape, where traditional vanilla plots and agrosystems and forest fragments (in different states of conservation) are essential elements for the conservation of the biocultural diversity and well-being of peasant populations. Therefore, planning the sustainable development of this important biocultural region must start from within the Totonac communities, taking advantage of their knowledge, practices, and worldview of nature, with the aim of conserving the biodiversity of tropical forests and the well-being of human populations.

Author Contributions

Conceptualization, N.V.-R., G.V.-D. and R.M.-M.; Methodology, N.V.-R., G.V.-D., R.M.-M., S.A.-M. and S.S.C.; Validation, N.V.-R. and G.V.-D.; Formal Analysis, N.V.-R., S.A.-M. and B.R.-G.; Investigation, N.V.-R., G.V.-D., S.S.C., R.M.-M., E.S.R., A.V.-M., S.A.-M., F.L.F. and R.C.G.; Resources, N.V.-R. and R.M.-M.; Data Curation, N.V.-R., B.R.-G. and S.S.C.; Writing—Original Draft Preparation, N.V.-R., B.R.-G. and R.M.-M.; Writing—Review and Editing, all authors; Visualization, N.V.-R. and S.A.-M.; Supervision, N.V.-R. and R.M.-M.; Project Administration, R.M.-M.; Funding Acquisition, R.M.-M. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by Secretaría de Ciencia, Humanidades, Tecnología e Innovación (SECIHTI), México, [CF-2023-G-559] and the APC was funded by Centro de Investigaciones Tropicales, Universidad Veracruzana.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

The raw data supporting the conclusions of this article will be made available by the authors on request.

Acknowledgments

We would like to thank the vanilla producers of the Totonacapan region of Veracruz and the company GAYA, Vanilla y Especias S. A. de C. V., for allowing the project to be carried out on their fields.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study.

Abbreviations

The following abbreviations are used in this manuscript:
FForest (medium semi-deciduous)
SSecondary forest
TVTraditional vanilla cultivation
AFVanilla agroforestry system
CVCitrus–vanilla system
DBHDiameter at breast height
NMDSNon-metric multidimensional scaling
SIMPROFSimilarity profile analysis
SIMPERSimilarity percentage analysis

Appendix A

Table A1. List of woody species recorded in fragments of medium semi-deciduous forest (F), secondary forest (SF), and vanilla production systems (traditional vanilla cultivation = TV; agroforestry vanilla cultivation = AF; citrus–vanilla = CV) in the Totonacapan region.
Table A1. List of woody species recorded in fragments of medium semi-deciduous forest (F), secondary forest (SF), and vanilla production systems (traditional vanilla cultivation = TV; agroforestry vanilla cultivation = AF; citrus–vanilla = CV) in the Totonacapan region.
FamilySpeciesFSFTVAFCV
AcanthaceaeOdontonema callistachyum (Schltdl. & Cham.) Kuntze X
AnnonaceaeAnnona scleroderma Saff. X
Cymbopetalum baillonii R.E.Fr.X
Desmopsis trunciflora (Schltdl. &Cham.) G.E.Schatz ex Maas, E.A.Mennega & WestraXX
Desmopsis uxpanapensis G.E.SchatzX
Mosannona depressa (Baill.) Chatrou X
ApocynaceaeCascabela ovata (Cav.) Lippold X
Cascabela thevetioides (Humb.& Bonpl.) LippoldXX
Tabernaemontana alba Mill.XXXX
AraliaceaeDendropanax arboreus (L.) Decne. & Planch.XX X
ArecaceaeAttalea butyracea (Mutis ex L.f.) Wess:Boer X X
Chamaedorea microspadix Burret X
Chamaedorea oblongata Mart. X
AsparagaceaeYucca gigantea Lem.X
AsteraceaeCritonia morifolia (Mill.) R.M.King & H.Rob.XX
Flaveria trinervia (Spreng.) C.MohrXXXX
BignoniaceaeParmentiera aculeata (Kunth) Seem.XXX
Tabebuia rosea (Bertl.) DC. XXX
BoraginaceaeEhretia tinifolia L. X
BurseraceaeBursera simaruba Sarg.XXXXX
Protium copal (Schltdl. & Cham.) engl.XXX
CannabaceaeAphananthe monoica (Hemsl.)J.-F.LeroyX
Celtis iguanaea (Jacq.) Sarg.X
CapparaceaeMorisonia quiriguensis (Standl.) Christenh. & Byng X
CaricaceaeCarica papaya L.XX X
CelastraceaeEvonymopsis mexicanus Benth. X
Crossopetalum parviflorum (Hemsl.) LundellX
Wimmeria concolor Schltdl. & Cham.XXX
Wimmeria obtusifolia Standl.X XX
ChrysobalanaceaeCouepia polyandra (Kunth) RoseX
CleomaceaeCleome uniglandulosa Cav.X
EuphorbiaceaeAcalypha macrostachya Jacq.XX
Adelia barbinervis Cham.& Schltdl.XXX
Alchornea chiapasana MirandaX
Alchornea latifolia Sw.X
Balakata luzonica (S.Vidal) EsserXXXX
Bernardia dodecandra (Sessé ex Cav.) GovaertsX
Cnidoscolus multilobus (Pax) I.M.Johnst.XXXX
Croton draco Schltdl.X XX
Croton soliman Cham. & Schltdl.X
Jatropha curcas L. XX
FabaceaePseudalbizzia tomentosa (Micheli) E.J.M.Koenen & Duno X
Pseudalbizzia tomentosa var. purpusii (Britton & Rose) E.J.M.Koenen & Duno X
Bauhinia divaricata L.XX
Bauhinia ungulata L. XXX
Cajanus cajan (L.) Millsp. X
Calliandra houstoniana var. anomala (Kunth) Barneby X
Calliandra trinervia var. arborea (Standl.) Barneby X
Cathormion umbellatum subsp. moniliforme (DC.) Brummitt X
Cojoba arborea (L.) Britton & RoseX
Diphysa carthagenensis Jacq.X
Diphysa microphylla Rydb. X
Diospyros nigra (J.F.Gmel.) Perr.X
Ehretia tinifolia L. X
Erythrina americana Mill. XX
Erythrina folkersii Krukoff & Moldenke X
Erythrina macrophylla DC. X
Gliricidia maculata (Kunth) Walp.XXX
Gliricidia sepium (Jacq.) Kunth XXXX
Inga acrocephala Steud. X
Inga punctata Willd. X
Inga sapindoides Willd. X
Leucaena leucocephala (Lam.) de WitXXXX
Lonchocarpus eriocarinalis MicheliX
Lysiloma divaricatum (Jacq.) J.F:Macbr. XX
Piscidia piscipula (L.) Sarg.XXX
Pithecellobium dulce (Roxb.) Benth. X
Pithecellobium lanceolatum (Humb. & Bonpl. Ex Willd.) Benth.XXX
Pterocarpus acapulcensis Rose X
Schnella glabra (Jacq.) DugandXX
Senna papillosa (Britton & Rose) H.S.Irwin & Barneby XXX
Vachellia cornigera (L.) Seigler & EbingerXXX
IcacinaceaeMappia racemosa Jacq.X
LauraceaeDamburneya salicifolia (Kunth) Trofimov & RohwerXXXX
Licaria capitata (Cham. & Schltdl.) Kosterm.X
Nectandra hihua (Ruiz 6 Pav.) Rohmer X
Nectandra rubriflora (Mez) C.K.AllenX X
Nectandra villosa Nees & Mart. X
Ocotea puberula (Rich.) NeesXX
Persea americana Mill.XXX
Persea cinerascens S.F.BlakeX
Persea longipes (Schltdl.) Meisn.X X
MalpighiaceaeBunchosia biocellata Schltdl. X
Bunchosia lindeniana A.Juss.XXXXX
Byrsonima crassifolia Kunth X X
Malpighia glabra L.X
MalvaceaeCarpodiptera cubensis Griseb.XX
Guazuma ulmifolia Lam.XXXX
Heliocarpus appendiculatus Turcz. XXX
Heliocarpus donnellsmithii Rose X
Malvaviscus arboreus Dill. ex Cav. X XX
Pachira aquatica Aubl. X
Robinsonella mirandae Gómez PompaXX
MeliaceaeCedrela odorata L.XXXXX
Swietenia humilis Zucc. X X
Trichilia havanensis Jacq.XXXXX
Trichilia hirta L.XX
Trichilia minutiflora Standl.X
Trichilia moschata Sw.X
Trichilia septentrionalis C.DC.X
MoraceaeBrosimum alicastrum Sw.XXX
Ficus aurea Nutt.X
Ficus benjamina L. X
Ficus maxima Mill. XXX
Ficus obtusifolia KunthXXX
Ficus yoponensis Desv.X
Trophis mexicana (Liemb.) Bureau X X
Trophis racemosa Urb.XXX
MyrtaceaeCalycorectes mexicanus O.BergX
Eugenia capuli Schltdl.XXXXX
Myrcia bartlettii (Standl.) A.R.Lourenço & Sánchez-Cháv.XX
Myrcia chytraculia (L.) A.R.Lourenço & E.LucasX
Myrcia chytraculia var. pauciflora (O.Berg) G.P.Burton & E. LucasX
Myrcia schlechtendaliana (O.Berg) A.R.Lourenço & Sánchez-Cháv.X
Pimenta dioica (L.) Merr.XXX X
Psidium guajava L. XX
NyctaginaceaePisonia aculeata L. X
PentaphylaceaeTernstroemia tepezapote Schltdl. & Cham.X X
PicramniaceaePicramnia antidesma Sw.X X
Picramnia teapensis Tul.X
PiperaceaePiper aduncum L. X
Piper amalago L.X
Piper aequale VahlXX
Piper hispidum sw.XX
Piper sanctum Miq.X
PolygonaceaeCoccoloba barbadensis Jacq. X
Coccoloba hirtella Lundell X
Coccoloba hondurensis LundellX
Coccoloba montana Standl.XX
PrimulaceaeArdisia compressa Kunth X
Ardisia tuerckheimii Donn.Sm. X
Bonellia macrocarpa subsp. pungens (A.Gray) B.Stahl & Källersjö X
Myrsine coriacea (Sw.) R.Br. Ex Roem. & Schult.X
Parathesis revoluta KunthX
RosaceaePrunus brachybotrya Zucc.X X
Prunus hainanensis (G.A.Fu & Y.S.Lin) H.Yu, N.H.Xia & H.G.YeX
RubiaceaeChione venosa var. mexicana (Standl.) David W.TaylorX X
Chomelia spinosa Jacq.XX
Exostema caribaeum (Jaq.)Roem & Schult.X
Hamelia patens Jacq. XXXX
Palicourea padifolia (Roem. & Schult.) C.M.Taylor & Lorence X
Palicourea pseudinundata (Wernham) Delprete & J.H.Kirbr. X
Palicourea sousae (Lorence & Dwyer) LorenceXXX
Posoqueria latifolia (Rudge) Roem. & Schult.X
Psychotria costivenia Griseb.X
Psychotria erythrocarpa Schltdl.X
Psychotria nervosa Sw.XXX
Psychotria papantlensis (Oerst.) Hemsl.X
Psychotria sarapiquensis Standl.X
Randia armata (Sw.) DC.XX
Randia laetevirens Standl.X
RutaceaeAmyris monofoliaris P.E.SánchezX
Citrus aurantiaca Swingle X
Citrus × aurantium L.XXXXX
Citrus limon (L.) Osbeck X
Esenbeckia pentaphylla Griseb.X
Murraya paniculata (L.) Jack X
SalicaceaeCasearia corymbosa Kunth X
Casearia laetioides (A.Rich.) Warb.XXXX
Pleuranthodendron lindenii (Turcz.) SleumerXXXX
Xylosma pubescens Griseb. XX
SapindaceaeCupania glabra Sw.XXXX
Litchi chinensis Sonn. X
Matayba oppositifolia BrittonXX
Sapindus saponaria L.XX
Talisia macrophylla Radlk.X
SapotaceaeChrysophyllum mexicanum Brandegee ex Standl.X X
Manilkara chicle (Pitter) GillyXXX
Manilkara sapota Van RoyenXXX
Sideroxylon obtusifolium (Roem. & Schult.) T.D.Penn.X X
Sideroxylon obtusifolium subsp. buxifolium (Roem. & Schult.) T.D.Penn.X
Sideroxylon persimile (Hemsl.) T.D.Penn.X
SolanaceaeCestrum racemosum Ruiz & Pav. XXX
Solanum umbellatum Mill. XX
UrticaceaeCecropia obtusifolia Bertol. X
Myriocarpa longipes Liebm.XX
Urera simplex Wedd.X
VerbenaceaeLippia myriocephala Schltdl. & Cham. XX
ViolaceaeRinorea guatemalensis BartlettX

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Sustainability 17 02598 g001
Figure 1. Location of vegetation fragments (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV), in the Totonapacan Veracruz, region.
Figure 1. Location of vegetation fragments (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV), in the Totonapacan Veracruz, region.
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Figure 2. Cluster (a) and non-parametric multidimensional scaling (b) for vegetation fragments (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV).
Figure 2. Cluster (a) and non-parametric multidimensional scaling (b) for vegetation fragments (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV).
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Figure 3. Unique species per vegetation type (forest = F; secondary forest = SF) and vanilla production system (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV).
Figure 3. Unique species per vegetation type (forest = F; secondary forest = SF) and vanilla production system (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV).
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Figure 4. Density of individuals (a) and basal area (b) of vegetation types (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV). Different letters on the bars indicate significant differences between vegetation types and vanilla production systems (p < 0.002).
Figure 4. Density of individuals (a) and basal area (b) of vegetation types (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV). Different letters on the bars indicate significant differences between vegetation types and vanilla production systems (p < 0.002).
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Figure 5. Relative abundance of regeneration types (light-demanding = D; intermediate successional species = I; shade-tolerant = T) in vegetation types (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV). Different letters on the bars indicate significant differences between regeneration types (p < 0.01).
Figure 5. Relative abundance of regeneration types (light-demanding = D; intermediate successional species = I; shade-tolerant = T) in vegetation types (forest = F; secondary forest = SF) and vanilla production systems (traditional vanilla cultivation = TV; vanilla agroforestry system = AF; citrus–vanilla system = CV). Different letters on the bars indicate significant differences between regeneration types (p < 0.01).
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Table 1. Richness and diversity of forest fragments and vanilla production systems in the Totonacapan region of Veracruz. q0 = species observed; q1 = exponential of Shannon; q2 = exponential of Simpson (confidence interval, 95%).
Table 1. Richness and diversity of forest fragments and vanilla production systems in the Totonacapan region of Veracruz. q0 = species observed; q1 = exponential of Shannon; q2 = exponential of Simpson (confidence interval, 95%).
SiteRichnessNumber of Individuals Estimate Sample Coverageq0q1q2
Forest11523320.99114.9 (9.4)45.4 (2.3)27.1 (1.7)
Secondary forest8210040.9681.9 (19.2)25.3 (2.6)15.2 (1.3)
Traditional vanilla cultivation7810270.9877.9 (9.5)28.7 (2.3)17.3 (1.6)
Agroforestry system 47110.9739.9 (26.8)12. 0 (1.2)7.8 (0.7)
Citrus–vanilla212870.9720.9 (13.3)5.2 (1.0)3.1 (0.5)
Table 2. Similarity values (Bray–Curtis) between groups resulting from the cluster and non-parametric multidimensional scaling.
Table 2. Similarity values (Bray–Curtis) between groups resulting from the cluster and non-parametric multidimensional scaling.
GroupABCDE
A
B17.19
C11.518.54
D7.046.542.04
E10.917.396.4727.2
F4.7512.342.619.6821.5
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Velázquez-Rosas, N.; Sinaca Colin, S.; Vázquez-Domínguez, G.; Velasco-Murguía, A.; Silva Rivera, E.; Ruiz-Guerra, B.; Friedrich, F.L.; Cortés Galindo, R.; Armenta-Montero, S.; Martínez-Mota, R. Importance of Traditional Vanilla Cultivation in the Conservation of Plant Diversity in Tropical Forests in Northern Veracruz, Mexico. Sustainability 2025, 17, 2598. https://doi.org/10.3390/su17062598

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Velázquez-Rosas N, Sinaca Colin S, Vázquez-Domínguez G, Velasco-Murguía A, Silva Rivera E, Ruiz-Guerra B, Friedrich FL, Cortés Galindo R, Armenta-Montero S, Martínez-Mota R. Importance of Traditional Vanilla Cultivation in the Conservation of Plant Diversity in Tropical Forests in Northern Veracruz, Mexico. Sustainability. 2025; 17(6):2598. https://doi.org/10.3390/su17062598

Chicago/Turabian Style

Velázquez-Rosas, Noé, Santiago Sinaca Colin, Guillermo Vázquez-Domínguez, Abril Velasco-Murguía, Evodia Silva Rivera, Betsabé Ruiz-Guerra, Fabio Levi Friedrich, Rosenda Cortés Galindo, Samaria Armenta-Montero, and Rodolfo Martínez-Mota. 2025. "Importance of Traditional Vanilla Cultivation in the Conservation of Plant Diversity in Tropical Forests in Northern Veracruz, Mexico" Sustainability 17, no. 6: 2598. https://doi.org/10.3390/su17062598

APA Style

Velázquez-Rosas, N., Sinaca Colin, S., Vázquez-Domínguez, G., Velasco-Murguía, A., Silva Rivera, E., Ruiz-Guerra, B., Friedrich, F. L., Cortés Galindo, R., Armenta-Montero, S., & Martínez-Mota, R. (2025). Importance of Traditional Vanilla Cultivation in the Conservation of Plant Diversity in Tropical Forests in Northern Veracruz, Mexico. Sustainability, 17(6), 2598. https://doi.org/10.3390/su17062598

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