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Article

Floristic Diversity and Green-Tree Retention in Intensively Managed Temperate Forests: A Case Study in Puebla, Mexico

by
Brenda E. Pérez-Pardo
1,
Alejandro Velázquez-Martínez
1,
Mireya Burgos-Hernández
2,* and
Valentín J. Reyes-Hernández
1
1
Programa de Posgrado en Ciencias Forestales, Colegio de Postgraduados, Km 36.5 Carretera México-Texcoco, Texcoco 56264, Mexico
2
Programa de Posgrado en Botánica, Colegio de Postgraduados, Km 36.5 Carretera México-Texcoco, Texcoco 56264, Mexico
*
Author to whom correspondence should be addressed.
Forests 2024, 15(6), 920; https://doi.org/10.3390/f15060920
Submission received: 14 April 2024 / Revised: 20 May 2024 / Accepted: 21 May 2024 / Published: 25 May 2024
(This article belongs to the Section Forest Biodiversity)
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Abstract

:
Clearcutting has tended to simplify forest structure and species composition, with potential negative consequences for biodiversity. Retention forestry emerged as an alternative to minimize this concern in intensively managed forests. In central Mexico, this approach was first implemented a decade ago in temperate forests, but an examination of its effects on floristic diversity and composition is non-existent. We evaluated and compared richness, diversity, and floristic composition among harvested, retention, and conserved areas in a conifer-dominated forest of central Mexico, with various parametric and non-parametric analyses. Species’ ecological and structural importance was also evaluated. We recorded 138 species, 95 genera, and 44 families of plants. Harvested areas listed the most species (99) with herb dominance, tree homogeneity, and the highest number of exotics. Retention and conserved areas’ floristic composition was similar, with the presence of epiphytes and terrestrial orchids, considered indicators of conservation. The retention areas recorded rare and endangered species, tree heterogeneity, and juveniles of structural species of temperate forests. Conserved areas showed a greater number of structural mature elements and exclusive species, though there was a smaller species number (75). We demonstrated that retention forestry is achieving its goal by maintaining the structural elements, habitats, and plant diversity of the temperate forests analyzed.

1. Introduction

In Mexico, temperate forests cover approximately 33 million hectares [1] and they are mostly located in the country’s main mountain systems, such as the Sierra Madre Occidental, Sierra Madre Oriental, and the Trans-Mexican Volcanic Belt. These ecosystems record a diversity of more than 8000 species of vascular plants [2,3] with large representation from the following genera: Pinus, Quercus, Abies, and Juniperus [4,5,6]. Despite this floristic richness, these ecosystems register an annual deforestation rate of 0.8%, being the second-most affected type of vegetation only after tropical forests [7].
Temperate forests are important in the Mexican forestry sector since they represent 73% of the managed forest area, providing more than 80% of timber production [8]. Silvicultural management is carried out under two technical schemes. The first is the Mexican Method of Management of Irregular Forests (MMOBI), a continuous cover system managed through selective cutting. The other is the Silvicultural Development Method (MDS), based on the cultivation of even-aged stands, and it is generally mono-specific [9,10,11]. The latter has become popular in pine forests since it provides technical facilities and a greater increase in volume and profitability [12,13]. However, a number of studies [14,15,16,17] have shown that these forest harvesting schemes modify the microenvironment within the stand. Furthermore, depending on the intensity of forest practices, the dynamics and structure of the forests are affected [18,19,20], due to changes in the upper canopy that influence the microclimate, availability of light, soil moisture regime, the solar radiation that reaches the forest floor, and, hence, processes such as the decomposition of organic matter and the availability of nutrients. This has a strong influence, both directly and indirectly, on the suitable conditions for the establishment of flora species and therefore their composition, altering the dynamics of the vegetation [21,22,23,24,25].
Nowadays, Mexican forests are facing different challenges, such as a decline in productive quality, constant pressure for land use change, and a growing demand for forest goods and services [10,12,23]. In response to the rapid transformation and homogenization of forests and to balance the objectives of production and biodiversity conservation, especially in complex social environments such as the Mexican forestry sector, variable retention forestry arises as a sustainable strategy, which recognizes the importance of structural complexity to maintain ecosystem functions and diversity [25,26,27,28].
The practice of forestry retention aims to maintain the dynamics and structural complexity of stands during forest harvesting through the retention of late-successional species and structures that emulate natural disturbance processes within the harvesting areas (e.g., live trees of different species and sizes, standing dead trees, understory vegetation, coarse woody debris, and gaps in the forest) [28,29,30]. The goal is not only to maintain the species diversity but also to enrich stands, restore impoverished or degraded ecosystems, as well as improve the connectivity of the forest landscape [31,32,33], and it also contributes to improving the forest aesthetics, which increases public acceptance of forest management [34].
Although variable retention forestry has been shown to have a positive influence on floristic richness and diversity, such evidence corresponds to studies in North America (Canada and the United States of America) [28,32,34,35,36,37], Europe [30,32,33,34,36,38], and the Argentine Patagonia [39,40]. In Mexico, this silvicultural practice is fairly recent, and studies that evaluate its effects on floristic diversity and composition conservation are scarce. This information is essential in the temperate forests of central Mexico, ecosystems that contain a high diversity of plants owing to the environmental conditions that result from the convergence of the Nearctic and Neotropical regions and their position in the Trans-Mexican Volcanic Belt [41]; also, this region has experienced one of the highest land use change rates in the country during recent decades [42,43], which indicates urgent attention. On the other hand, Mexican ejidos offer good examples of community-based forest management geared toward timber production that have also managed to conserve more than half of the country’s forests and improve rural livelihoods, providing timber and other non-timber forest resources in many parts of Mexico, including the central region [10,44,45].
The lack of research evaluating the effectiveness of the retention forestry constitutes an important limitation for forest management and biological conservation in central Mexico. The objectives of this study were as follows: (i) to evaluate the richness, diversity, and composition floristics in a conifer-dominated forest of central Mexico; (ii) to compare this information among harvest (i.e., clearcut areas), retention (i.e., forest retention or green-tree retention), and conserved areas (i.e., untouched areas designated within this managed forest); and (iii) to analyze the importance of this silvicultural practice in managed forests. Under this context, we propose the following hypotheses: taking into account that the management forests considering conservation can support some floristic diversity, and this type of management tends to generate a greater species richness [46,47], we expect a high richness and diversity in harvest areas, followed by green-tree retention areas. Also, we hope that retention areas will present a close floristic composition with conserved areas, mainly related to woody and structural elements of temperate forests. Based on previous studies in other countries of the Northern Hemisphere [28,30,32,33,34,35,36,39,46], we hypothesize that retention areas positively influence harvest areas, mainly in those with the highest successional state, which will be reflected primarily in the plant species composition but also in floristic richness and diversity.

2. Materials and Methods

2.1. Study Site

The Sierra Norte de Puebla (SNP) is a mountain range that corresponds to the transition zone between the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt [48]. The region is characterized by a rugged topography, with elevations reaching 3400 m [49]. We used information from a temperate forest dominated by Pinus patula Schiede ex. Schltdl. & Cham., which has been managed for 40 years and is located in the southwest of the SNP, in the municipality of Chignahuapan. This municipality is located between the coordinates 19°40′ and 19°59′ North latitude and 97°57′ and 98°19′ West longitude (Figure 1a). Chignahuapan is immersed between mountains and hills, with elevations ranging from 1720 to 3400 m. The average annual temperature ranges between 10 and 16 °C, the average annual precipitation is 600 to 900 mm, and the climate varies from temperate subhumid C(w) in most of the municipality to semi-cold subhumid C(E) in the high mountainous parts [50], with temperate forests being the dominant ecosystem, covering 37% of the municipal area [51,52]. Consequently, forest management and the industry associated with this sector are the most important economic activities of the region [53].
The Ejido Llano Grande (Figure 1a) is a common forest area located south of Chignahuapan, where forestry has been the dominant land use type and economic activity type since 1983, which is mainly performed in the main vegetation type, the Pinus patula forest. The Ejido has 1700 hectares of forest area, of which 19,258 m3 of round wood is legally extracted annually. Forest management is carried out with the Forestry Development Method (MDS by its Spanish acronym, Método de Desarrollo Silvícola), with a 10-year cutting cycle and a 50-year rotation cycle. The treatments include plantation maintenance, thinning, and regeneration cutting, which is applied as total cuts (i.e., clearcutting) with immediate plantation [51]. Llano Grande is the ejido with the largest area and volume authorized within UMAFOR 2108 “Chignahuapan-Zacatlán” and contributes up to 55% of the total timber production of Puebla State [51,54].
To protect areas considered of High Conservation Value (HCV), such as relict oak forests, wildlife refuges, and springs, making it a national reference, the Ejido implemented diversity conservation practices on 199.91 ha through the Biodiversity in Production Forests and Certified Markets project [55]. Within these practices, Llano Grande was the first to implement retention forestry, which has been practiced since 2013 in a variable scheme, that is, by combining aggregate retention in the form of strips and dispersed retention, with structures such as live trees, dead trees, and coarse woody debris distributed within the cutting areas [51].

2.2. Floristic Inventory

The plant collections were carried out through stratified random sampling [56] by harvested areas (HAs) (i.e., areas where clearcuts with immediate plantation were applied), structural tree retention (STR) (i.e., 20 m wide retention strips, with retention of trees, woody debris, and understory vegetation), and conservation areas (CAs) (i.e., areas designated for conservation in which no management is conducted). The HAs consisted of three polygons, harvested in the years 2014 (HA2014), 2016 (HA2016), and 2018 (HA2018). The STRs were integrated by three retention strips, each located within the HA polygons and named from now on as STR2014, STR2016, and STR2018. The CAs consisted of three polygons known as El Gambito (CAEG), La Encinera (CALE), and El Campanario (CAEC), all of them dedicated to conservation, therefore presenting similar ecological conditions and altitudinal profiles among them (Figure 1b).
A total of 71 sampling points were randomly established, in which an inventory of the vascular flora was performed by the plot method of 100 m2 (10 m × 10 m), which was distributed as follows: 30 in the HAs, 29 in the STRs, and 12 in the CAs (Figure 1c). Within each plot of 100 m2, all the woody vegetation was recollected, and three subplots of 4 m2 (2 m × 2 m) were randomly established to inventory the herbaceous stratum; hence, all the different forms of growth of the vascular flora were included [56,57,58].
For each taxon collected, we recorded its habit according to what is described by Simpson [59], its height which was measured following the recommendations in Aranda et al. [60], and coverage-abundance using the Braun-Blanquet scale modified by van der Maarel [61]. In the case of trees, the number of individuals with normal diameter (DN) > 5 cm by species was counted [62]. Rare species or species under a risk category were recorded only photographically. The herborization process was according to what was proposed by Lot and Chiang [63].
The taxonomic classification of the botanical material was completed through the use of monographs [5,64,65,66,67,68,69,70], dichotomous keys in regional floras [71,72,73,74,75,76,77,78,79,80,81,82,83,84,85,86], and floristic lists [87,88,89]. Scientific names and taxonomic authorities for each taxon were verified and corrected based on the International Plant Names Index (https://www.ipni.org, accessed on 6 April 2024), TROPICOS (https://www.tropicos.org/home, accessed on 6 April 2024), Catalogue of Life (https://www.catalogueoflife.org, accessed on 7 April 2024), and Plants of the World Online (https://powo.science.kew.org, accessed on 7 April 2024) websites.
The specimens identified were deposited in the herbarium CHAPA at the Colegio de Postgraduados, Mexico. The list of species followed the order of the most recent classifications published for each taxonomic group. For ferns and related species, we used the system proposed by PPG I [90]; for gymnosperms, the system of Christenhusz et al. [91]; and for angiosperms, the one proposed by APG IV [92]. The conservation status of the species was identified according to the NOM-059-SEMARNAT-2010 [93] and the red list of the International Union for Conservation [94]. Endemism was determined following Rodríguez-Acosta et al. [88], Villaseñor [3], and Rzedowki [6,85], and for synanthropic and exotic species, Villaseñor and Espinosa-García [95], as well as the Mexican weeds website [96], were consulted.

2.3. Floristic Richness and Diversity

The sampling effort was analyzed using species accumulation curves based on the non-parametric estimator Chao 2 of species richness. This estimator has little sensitivity to the distribution of species, making it suitable for application in modified communities such as managed forests [97,98]. Accumulation curves were generated using EstimateS v9.1.0 [99]. The completeness level was calculated as C = (S obs)/(S Chao 2) × 100 [100], for which a minimum level of 70% was established to be considered acceptable sampling [99].
Richness was determined as the number of species, genera, and families recorded. Floristic diversity was analyzed by stratum [101] using Simpson’s dominance index ( D = ∑ p i 2 ) and the Shannon–Wiener entropy index ( H ′ = − ∑ p i ×   n p i ) [56,102]. These indices express a measure of the community’s complexity based on the species’ proportional abundance, so their use is recommended both in natural communities and in modified environments [103,104]. The ecological importance of each species by plant community was also considered by estimating an index of the adjusted ecological importance value (AEIV) [105].

2.4. Floristic Composition

Based on the floristic inventory, a binary presence–absence matrix was developed to evaluate the floristic similarity between HAs, STRs, and CAs, using Jaccard’s similarity coefficient [102], and we constructed a dendrogram with the hierarchical clustering method that uses the unweighted pair group method with arithmetic mean (UPGMA) [57,102]. To graphically visualize the similarity among the groups, the non-parametric multidimensional scaling (NMDS) ordination method was used [106,107,108]. An analysis of similarities (ANOSIM) was also performed to obtain the significance of these similarities [106,109].
The percentage contribution of species to the dissimilarity between groups was calculated by a similarity percentage analysis (SIMPER) based on Euclidean distance. Finally, the turnover of species between areas was represented using a Venn diagram, in which the exclusive and shared species between the HAs, STRs, and CAs were identified [110]. All analyses were completed in PAST v3 [111].

3. Results

3.1. Richness and Diversity Floristics

A total of 138 species, 95 genera, and 44 families were recorded in the study site (Table S1), over a total sampling area of 7100 m2. Asteraceae, Poaceae, Orchidaceae, Rosaceae, and Ericaceae were the most representative families, contributing 48.5% of the species registered. At the genus level, Ageratina, Quercus, Pseudognaphalium, Roldana, and Salvia L. showed the highest richness, accumulating 17% of all species (Table 1).
Overall, the most frequent species in the tree stratum were P. patula, Abies religiosa, Quercus castanea, Q. crassipes, and Prunus serotina. The common elements in the shrub stratum were Lonicera mexicana, Ribes affine, Cestrum fasciculatum, Salvia elegans, Roldana angulifolia, Vaccinium leucanthum, Monnina ciliolata, and Smilax moranensis. In the herbaceous stratum, the dominant species were Erigeron longipes, Alchemilla procumbens, Didymaea alsinoides, and Adiantum andicola. Herbs were the best represented (83 species), followed by shrubs (27) and trees (17), while climbers (6) and epiphytes (5) were the least represented. Of the flora inventoried, three new records stand out for the state of Puebla: Geum canadense, Rubus pumilus, and Pedicularis canadensis (Table S1).
The flora of Llano Grande was mostly composed of native species, with 16% endemism. The exotic component consisted of seven species: Pseudognaphalium luteoalbum, Sonchus asper, Taraxacum officinale, Rumex acetosella, Senecio inaequidens, Cirsium vulgare, and Hedera helix. These last three are reported as invasive. Finally, no species was found in an important category according to the IUCN Red List of Threatened Species, and only Monotropa hypopitys (Figure S1) is protected by national legislation according to NOM-059-SEMARNAT-2010.

3.1.1. Harvested Areas (HAs)

The species accumulation curve (Figure 2a) indicated that it was possible to have 80% of the richness expected in the harvest areas, represented by 3% of monilophytes, 2% of gymnosperms, and 95% of angiosperms. Of the latter, 87% are eudicots and 13% are monocots. Here, a total of 99 species, 69 genera, and 36 families were recorded, with Asteraceae, Poaceae, Fagaceae, and Rosaceae being the most diverse families, since together they represented 39% of the genera and 50.5% of the species recorded. The most representative genera were Quercus, Pseudognaphalium, Roldana, and Ageratina (Table 2), which contain 18.18% of the species in these areas. According to the habit, 62% of the species are herbs, 19% shrubs, 15% trees, and 4% climbers. Most of the flora is native (82%), 17 species are endemic to Mexico (Table S1), and 6 exotic species recorded in Ejido Llano Grande were registered here (except H. helix).
The HAs (Figure 3) showed floristic homogeneity, particularly in the tree stratum, where P. patula is the dominant species (AEIV = 27.31), followed by P. serotina (AEIV = 22.30). The diversity was medium according to the diversity indexes obtained (IS = 0.22; IS-W = 1.83). Individuals of Q. castanea, Q. laurina, Buddleja cordata, and Salix paradoxa. were found, although they were juveniles and had low frequency. Among the shrub elements, L. mexicana (AEIV = 23.70), R. affine (AEIV = 15.23), and C. fasciculatum (AEIV = 13.11) stand out. This stratum recorded a diversity of SI = 0. 20 and IS-W 2.04. The herbaceous stratum is poorly represented, with E. longipes (AEIV = 26.30) and A. procumbens (AEIV = 10.77) as the common species with a diversity of IS = 0.26 and IS-W = 2.08.

3.1.2. Structural Tree Retention Stripes (STRs)

According to the species accumulation curve, 76% of the estimated richness was captured in the retention areas (Figure 2b), represented in 97 species, 73 genera, and 38 families. Of the latter, Asteraceae, Poaceae, Ericaceae, Rosaceae, and Lamiaceae included 44% of the genera and 48% of the recorded species. The genera with the highest number of species were Quercus, Roldana, and Ageratina, which together represent 13.4% of the inventoried species (Table 2).
The STRs (Figure 3) presented a developed tree stratum, reflected in high diversity values (IS = 0.14; IS-W = 2.07) and with Q. castanea (AEIV = 17.92), A. religiosa (AEIV = 16.46), S. paradoxa (AEIV = 14.99), P. patula (AEIV = 13.03), and P. serotina (AEIV = 12.51) as the most common species and with the highest values of AEIV. The shrub stratum was variable, as it presented both areas with abundant vegetation and others with clearings. The above was reflected in the levels of diversity that varied from high to medium (IS = 0.11 and IS-W = 2.47). The dominant species were V. leucanthum (AEIV = 16.45), L. mexicana (AEIV = 11.26), and Roldana barba-johannis (AEIV = 8.97). In the herbaceous stratum, A. andicola (AEIV = 12.64), followed by Piptochaetium fimbriatum (AEIV = 6.44) and A. procumbens (AEIV = 6.22), were the most frequent species. This stratum presented diversity values of IS = 0.10 and IS-W = 2.76.
A total of 13% of the species registered were trees, 22% were shrubs, and 59% were herbs. A total of 1% of the species were epiphytes and 6% were climbers. A total of 87% of the flora was native, with 15 endemic to Mexico. Three species were exotic (C. vulgare, T. officinale, and H. helix), of which H. helix is invasive. Monotropha hypopitis was conspicuous among the species recorded in this area, and it is subjected to special protection (Pr) (Table S1).

3.1.3. Conservation Areas (CAs)

The species accumulation curve (Figure 2c) indicated that in the conservation areas, it was possible to have 84% of the estimated richness represented in 75 species, 58 genera, and 34 families. Among the latter, Asteraceae, Rosaceae, Ericaceae, and Polypodiaceae stand out, which together represented 36.21% of the genera and 37.33% of the registered species. Ageratina, Roldana, Quercus, and Salvia were the genera with the most species, representing 17.33% of the species (Table 2).
The conservation areas presented greater variability in environmental conditions (Figure 3) and in the tree flora that resides in them, reflected in high-to-medium global diversity values (IS = 0.16; IS-W = 2.07). El Gambito (CAEG) is located at an altitude of 2785 m, with an area of 18,825 m2, and the dominant tree species were A. religious and P. patula. La Encinera (CALE) has an area of 62,462 m2 of remnant oak forest, at an altitude of 2733 m. Here, the main arboreal elements are Q. castanea and Q. laurina. For its part, El Campanario (CAEC) is the largest conservation area (227,003 m2) and is located in the highest altitude forest, standing above 3000 m. In this area, A. religiosa, Q. castanea, Q. laurina, and P. patula were the dominant species.
The tree stratum in the CAs reflected high diversity (IS = 0.16; IS-W = 2.07), and Q. castanea (AEIV = 22.38) was the most common specie. The shrub stratum in the CAs was dominated by L. mexicana (AEIV = 13.82), C. fasciculatum (AEIV = 13.10), and S. elegans (AEIV = 12.96). The diversity values were IS = 0.12 and IS-W = 2.36. Meanwhile, the herbaceous stratum presented a diversity of SI = 0.17 and IS-W = 2.36, with Geum canadense (AEIV = 17.23) and A. andicola (AEIV = 13.86) as the structural elements. Herbs were the main life form (56%), followed by shrubs (23%), while trees were the least represented (16%). A total of 1% of the species were epiphytes and 5% were climbers. A total of 85% of the flora is native, with 11 species endemic to Mexico. No exotic or threatened species were recorded here.

3.2. Floristic Composition

According to the Jaccard similarity analysis, two large groups were identified (Figure 4, groups 1 and 2) at a distance of Jaccard = 0.28. Group 1 concentrates all the CA sampling plots, all of the STRs, and almost all of the HAs of the polygon 2014 (HA2014). Within subgroup 1A, STR2016 and STR2018 show floristic compositions that are similar to each other (group 1Aa), except plot STR201803, which showed greater floristic similarity with some plots from the CAs of El Campanario (group 1Ab). Meanwhile, three STRs from the 2016 polygon were more similar to the CAs corresponding to El Gambito (ACEG) and La Encinera (ACLE) (group 1Ac). Another subgroup (1B) within group 1 shows greater floristic similarity between the plots corresponding to STRs (1Ba) and HAs of polygon 2014 (1Bb) at a Jaccard distance of 0.35. Meanwhile, group 2 integrated all the HA plots of polygons 2016 and 2018, except plot HA201602 which was integrated into group 1Aa, as well as plot HA201405.
The NMDS analysis showed differences between HAs and CAs. An overlap was observed between the STRs and the CAs, with a poor stress value (0.2894) (Figure 5). The ANOSIM (Table 3) showed significant differences (p < 0.05) between the three areas. The greatest floristic difference detected occurred between the HAs and the CAs (R = 0.719) and the smallest between the STRs and CAs (R = 0.256), coinciding with what was observed in the dendrogram and NMDS. The SIMPER analysis detected 10 species with the largest contribution to the total dissimilarity which accumulated 19.5% of the total dissimilarity (Table 4).
A total of 49 species shared between the three types of areas were identified. A total of 24 species are shared between the HAs and the STRs, and only 4 are shared between the HAs and CAs. The HAs are the ones that had the highest number of unique species, while the CAs recorded 15 (Figure 6).
Regarding the areas evaluated, in the HAs, only the Asteraceae family represented 32.32% of the inventoried flora, followed by Poaceae with 7%, so the difference between the two most diverse families is 25.32%. These numbers are important since they explain the high-to-medium diversity values obtained according to the indices evaluated. Although in the STRs these two families were also the most numerous in terms of species, the gap between them was smaller (14.4%) and the diversity values recorded were higher than those of the HAs. The CAs shared the Compositae as the most representative group, but the gap with the second-most diverse family (Rosaceae) was rather small (9.3%), showing greater heterogeneity of taxonomic groups although with fewer species concerning the two areas previously.
Although at the level of the number of species, Quercus dominates in the first two areas (HAs and STRs), and juvenile individuals were mainly found in the HAs, without mature elements. Meanwhile, P. patula, which is the species with the greatest ecological importance in the area, together with Prunus serotina, were the elements that dominated the tree stratum.
In the STRs and CAs, the results were different; unlike in the HAs, the arboreal stratum was heterogeneous. In the STRs, Q. castanea, A. religiosa, S. paradoxa, P. patula, and P. serotine were the dominant components and recorded the highest ecological importance values. In the CAs, the ecologically important species with the greatest presence were A. religiosa, Q. castanea, Q. laurina, and P. patula.
Lonicera mexicana was the structural species of the shrub stratum that the three areas share as the one of greatest ecological importance. In the herbaceous stratum, they differ in composition. In the HAs, E. longipes is the most common species, and in the STRs, it is A. andicola. This last species together with G. canadense stands out in the CAs.

4. Discussion

4.1. Floristic Composition, Richness, and Diversity

The Pinus patula forest of the Ejido Llano Grande protects an important floristic diversity, registering 138 species in an inventoried area of 7100 m2. This number represents 0.55% of the vascular plants estimated for Mexico [112] and 2.36% of those reported for the temperate forest biome in the country [3]. The recorded flora is equivalent to 2.76% of the vascular plants reported for the State of Puebla [113], as well as 14.5% [2,3] and 14% of the flora reported for temperate forests at the state level and in the “Chignahuapan-Zacatlán” Supply Basin at the regional level [89], respectively.
As far as plant diversity in managed forests is a subject of public concern, the floristic richness recorded in the forest of the Ejido Llano Grande is larger than that reported in other similar works in the country [114,115]. For example, Luna-Bautista et al. [114] reported 43 species in an area of 29,200 m2 in a Pinus oaxacana forest, in the State of Oaxaca, with 45 years of management under the forestry conservation and development system (SiCoDeSi). Meanwhile, Rendón-Pérez et al. [115] in a Pinus and Pinus-Quercus forest in the state of Hidalgo, with more than 100 years of management under the MDS, recognized 88 species of vascular plants in an area of 31,600 m2. These differences in the number of species may be due to the effect of the type of forest management and the sampling method used. It should be noted that these floristic studies are among the few completed in managed forests in Mexico, with the most numerous being those focused just on the tree layer [11,116,117,118,119,120]. This indicates the existence of important gaps in information regarding other life forms and strata and highlights the importance of carrying out this type of study.
The best-represented plant families correspond to the most diverse ones recorded in Mexico [3], particularly Asteraceae, Poaceae, and Orchidaceae are among the 10 most diverse in Puebla State [113]. The high richness of the Asteraceae family agrees with that reported by Luna-Bautista et al. [114] and Rendón-Pérez et al. [115] in temperate forests under management, as well as with Velasco-Luis et al. [121] in an unmanaged disturbed forest. However, our results differ by reporting a high number of species of grasses and orchids, groups that were not among the most numerous in the aforementioned studies. This draws attention since it is known that most orchid species are distributed in environments with little disturbance [122].
The floristic dominance that we obtained agrees with Rzedowski [6], who mentions that the most cited families are the richest in temperate forests, and Villaseñor [3], who reports Asteraceae as particularly rich in temperate forests. According to Handal-Silva et al. [113], 20% of Asteraceae in Puebla are attributed to their presence in coniferous and oak forests and, secondly, to the disturbance of areas with vegetation cover.
In this study, two genera predominated in number of species, Quercus and Ageratina, and both genera are among the 25 most diverse in the country [3]. Oak forests are a characteristic element of the Mexican temperate forests and are frequently mixed with Pinus forests and other conifers. The highest diversity of this genus is concentrated in the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt [5,6,113], the geographical area where the Ejido Llano Grande is located, which explains why this group of plants occupies the first place regarding the number of species. Ageratina is reported as a genus with a high presence in temperate forests [123], therefore, our result is not surprising. Pseudognaphalium, Roldana, and Salvia are genera that are also recorded with a greater presence in the temperate forests of the country with distribution along the Trans-Mexican Volcanic Belt [72,124], so they have a wide presence in the state of Puebla and were recorded with high richness in this work.
The dominance of herbs in the Llano Grande forest, followed by shrubs, is consistent with reports by Rendón-Pérez et al. [115] and Luna-Bautista et al. [116] in areas similar to the study site in managed temperate forests. The high representativeness of herbaceous and shrub elements is related to the high number of Compositae and grasses recorded in the study site. Meanwhile, the lower representation of epiphytes and climbers is characteristic of coniferous forests, since these groups register their greatest diversity and abundance in tropical ecosystems [113,125].
The SIMPER analysis showed that the understory is responsible for most of the floristic dissimilarity between areas, since 9 of the 10 species with the largest contribution to the total dissimilarity were herbs and shrubs, coinciding with Luna-Bautista et al.’s [114] results in a temperate forest in Mexico. This outcome is not unusual in managed forests, since the main use is carried out in the tree layer, which in this study is reported as more homogeneous in HAs with respect to STRs and CAs, as mentioned above.
The floristic richness of Puebla increases to 5235 species with the incorporation of three new records through this study and 465 for the Chignahuapan-Zacatlán Supply Basin. The endemism reported here is greater than those recorded in other managed forests, which is related to the location of the ejido, within the Trans-Mexican Volcanic Belt, which stands out for its high richness and endemism [126,127]. It should be noted that the figure reported here is equivalent to 1.17% of the 1935 national endemic species that the state of Puebla protects reported by Villaseñor [3] and 0.19% of the reported species from the entire country by the same author.
In terms of exotic species, three are prominent given their invasive nature and the potential damage that they can cause to the native flora in the medium term: S. inaequidens, C. vulgare, and H. helix [128,129]. Meanwhile, M. hypopitys, a saprophyte subject to special protection (Pr) according to national legislation [93], was rare in the study area but reported as typical of the Abies forest [130]. The mycoheterotrophic genus Monotropa L. is commonly distributed in temperate coniferous forests [77]. Particularly, M. hypopitys has an exclusive presence in mature forests [131]. The red form of M. hypopitys (here recorded) indicates high mycorrhizal specificity [132].
The results mentioned above highlight the importance of replicating this type of study, especially in managed forests, since it is possible to find several typical structural elements of these ecosystems and also shows the limited botanical exploration made so far in the region, which is necessary to correctly manage these natural resources. This is in addition to showing possible management lines such as the control and monitoring of exotic species. Therefore, we consider that the present work constitutes a contribution to floristic knowledge in managed temperate forests, emphasizing the findings of new records for the State of Puebla.

4.2. Importance of Forest Retention in Conservation

The physiognomic characteristics and taxa found in the Llano Grande forest corresponded with the description of the Pinus forest of the Trans-Mexican Volcanic Belt [130]. Of the genera and structural species described by Rzedowski [130] for this vegetation, A. religiosa, Q. laurina, P. serotina, S. paradoxa, L. mexicana, R. affine, V. leucanthum, S. elegans, and C. fasciculatum were identified as the species with a high value of ecological importance in the different studied areas included.
Although Ejido Llano Grande is a managed forest, the implementation of retention forestry and conservation areas as a management strategy is highly functional, since this forest conserves a significant number of native species, of which 24 are part of the synanthropic flora of Mexico. The number of exotic elements reported here (i.e., seven species) agrees with the results of Santibáñez-Andrade et al. [124] in the Magdalena River Basin, with proportions between 5 and 9% in a forest in the center of the country.
The HAs and STRs showed a greater number of species than the conservation areas, concentrated mainly in the herbaceous elements, which respond favorably to the increase in luminosity and generally present high reproductive success [22,133]. These results are consistent with various studies that found an increase in the richness of harvested stands [22,133] and in retention areas [34,36,134]. In harvested stands, this occurs due to the influx of ruderal and exotic plants [21,25,133] and in retention areas, by native forest species, in addition to ruderal and exotic plants [36,37,39,134]. This could be verified in Llano Grande since the largest number of exotic species were concentrated in the HAs and to a lesser extent in the STRs; STRs also recorded juvenile species from mature forests. The presence of species characteristic of mature forests may be due to the habitat provided by the retained structures, such as trees and woody debris, typical of advanced successional states [27,28].
The homogeneity of the tree stratum in the HAs by individuals of P. patula is also noticeable, since this was the species with the greatest ecological importance. This is the result of the type of forest management that is performed in these areas and that induces the dominance of this species of pine for commercial purposes [11,115,117]. In contrast, in the STRs and the CAs, the arboreal stratum was heterogeneous, although in the STRs, more juvenile individuals were found, while in the CAs, mainly mature elements were observed.
The diversity values obtained also agree with what was observed by Luna-Bautista et al. [114] in south Mexico but differ from other authors who claim that diversity increases after harvest [25,133,135,136]. However, plant diversity may not present a pattern of direct response to silvicultural management, but it may be inversely related to species richness, as it has been noted by other authors [137].
Jaccard’s analysis showed greater floristic similarity between the STRs and CAs, which demonstrates that the STRs are meeting the objective of protecting floristic elements of mature forests typical of these ecosystems. This was verified by recording plants associated with shade, humid, or mesic conditions and specialists such as orchids, epiphytes, and Monotropa hypopitys. Other examples are Orthosia angustifolia, Symphoricarpos microphyllus, R. pumilus, Monotropa uniflora, Corallorhiza maculata, Goodyera striata, and Arbutus xalapensis, which were not found in the logging areas. The presence of epiphytes (Pleopeltis polylepis, Tillandsia sp.) and terrestrial saproxylic orchids (C. maculata, G. striata) indicates that the forest still maintains a good state of conservation since these groups are considered indicators of vegetation conservation [138,139]. For example, Jiménez-Bautista [140] found that epiphyte richness and diversity showed a marked reduction in secondary forests compared to primary forests, especially in the case of Orchidaceae and Polypodiaceae. Other studies have shown the importance of STRs in other parts of the world such as refuges for bryophytes, epiphytes, and late seral species [28,32,37].
It is remarkable to have found a greater floristic similarity of older HAs (AA2014) with nearby STRs (STR2014), which suggests that STRs affect the floristic composition of the adjacent HAs in the short term. This edge effect exerted by STRs has not been studied in depth; studies suggest that the influence of STRs is limited to short distances [35,37,141]. However, this study shows that at a distance of 50 m, the understory vegetation may be enriched by the STRs. In managed stands, Luna-Bautista et al. [114] found that the species shared between stands increased with the age of the stand, which could explain why only the HA polygon with the longest succession time showed similarity with the nearby STRs, given the influence of the pioneer species [142].
As a result of its forest management history, the Llano Grande forest has few structural pine forest species and some species associated with disturbance conditions. However, the practice of conserving floristic diversity has resulted as important since, thanks to this activity, the forest protects the habitat suitability that allows for the establishment of species such as M. hypopitys, as well as Orchidaceae and Bromeliaceae elements and whose shade conditions and greater humidity are essential for their establishment [23,131,140]. These conditions are present in STRs and CAs for forest cover, tree stratum heterogeneity, and understory vegetation. Furthermore, epiphytes are one of the botanical groups that are most affected by forest management [134], and the practice of retention allows for maintaining some of these elements and their habitat, which in turn serves as a habitat for other living organisms and allows for the proper functioning of these ecosystems.

5. Conclusions

The forest of the Ejido Llano Grande protects an important floristic richness, with 138 species of vascular flora, 95 genera, and 44 families, as well as several endemic plants. Our study presents an assessment of the diversity level and floristic composition of a managed forest where the first areas of retention forestry were implemented more than ten years ago in Mexico. We demonstrated that it is possible to reconcile the dichotomous debate between diversity conservation vs timber production since the retention areas together with conservation areas are fulfilling their role of protecting and maintaining the diversity of plant species in the short term as well as the structure of the ecosystem in accordance with the production of wood and other products to benefit local communities. The value of retained trees for biodiversity conservation will likely increase over time, as they will become older compared to production forest trees. Future studies should be performed in other areas where this silvicultural approach has been implemented since local conditions can influence the proper management of the species and the establishment of retention areas. The results could be used to improve this practice in forest management areas, to have a win–win situation in conservation and production.

Supplementary Materials

The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/f15060920/s1, Table S1: Floristic list of vascular flora recorded in the Pinus patula-dominated forest in the Ejido Llano Grande, Chignahuapan, Puebla, Mexico; Figure S1: Specimens of Monotropa hypopitys recorded in Llano Grande, Chignahuapan, Puebla, Mexico.

Author Contributions

Conceptualization, A.V.-M.; methodology, B.E.P.-P. and M.B.-H.; software, B.E.P.-P.; validation, B.E.P.-P. and M.B.-H.; formal analysis, B.E.P.-P. and M.B.-H.; investigation, B.E.P.-P., M.B.-H., A.V.-M. and V.J.R.-H.; resources, B.E.P.-P.; data curation, B.E.P.-P. and M.B.-H.; writing—original draft preparation, B.E.P.-P. and M.B.-H.; writing—review and editing, B.E.P.-P., M.B.-H., A.V.-M. and V.J.R.-H.; visualization, M.B.-H. and B.E.P.-P.; supervision, M.B.-H. and A.V.-M.; project administration, A.V.-M.; funding acquisition, B.E.P.-P. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Council of Humanities, Science and Technologies (student grant B.E.P-P.-CONAHCYT-1079483).

Data Availability Statement

The data presented in this study are available upon request from the first author.

Acknowledgments

The authors thank the ejidal authority of Ejido Llano Grande, Antonio Rodríguez Salazar, for his support in the fieldwork. We are grateful to the staff of the herbarium CHAPA; Mauricio Mora and Rolando Jiménez, for their support during the identification of specimens; as well as the three anonymous reviewers for their valuable comments that allowed us to improve this manuscript.

Conflicts of Interest

The authors declare no conflicts of interest.

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Forests 15 00920 g001
Figure 1. (a) Geographic location of the Ejido Llano Grande in the Sierra Norte de Puebla, Puebla State, Mexico. (b) Location of the polygons of HAs (HA2016, HA2018, and HA2014), STRs (STR2016, STR2018, and STR2014), and CAs (CAEC, CALE, and CAEG) within the Ejido Llano Grande. (c) Dots show the distribution of sampling points in each HA, STR, and CA polygon.
Figure 1. (a) Geographic location of the Ejido Llano Grande in the Sierra Norte de Puebla, Puebla State, Mexico. (b) Location of the polygons of HAs (HA2016, HA2018, and HA2014), STRs (STR2016, STR2018, and STR2014), and CAs (CAEC, CALE, and CAEG) within the Ejido Llano Grande. (c) Dots show the distribution of sampling points in each HA, STR, and CA polygon.
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Figure 2. Accumulation curves of the vascular flora species recorded in the Ejido Llano Grande, Chignahuapan, Puebla, Mexico by the following: (a) harvest areas (HAs), (b) structural tree retention stripes (STRs), and (c) conservation areas (CAs). C.I. = Confidence Interval.
Figure 2. Accumulation curves of the vascular flora species recorded in the Ejido Llano Grande, Chignahuapan, Puebla, Mexico by the following: (a) harvest areas (HAs), (b) structural tree retention stripes (STRs), and (c) conservation areas (CAs). C.I. = Confidence Interval.
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Figure 3. Physiognomy of the vegetation in the evaluated areas in the P. patula-dominated forest of the Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Harvested areas (HAs), structural tree retention stripes (STRs), conservation areas (CAs), and their polygons (2014, 2016, and 2018). Photographs by Brenda E. Pérez-Pardo.
Figure 3. Physiognomy of the vegetation in the evaluated areas in the P. patula-dominated forest of the Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Harvested areas (HAs), structural tree retention stripes (STRs), conservation areas (CAs), and their polygons (2014, 2016, and 2018). Photographs by Brenda E. Pérez-Pardo.
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Figure 4. Dendrogram recovered from a hierarchical cluster analysis (UPGMA) based on the Jaccard coefficient indicating floristic similarity between the sampling plots. Green and blue colors show the two large groups according to Jaccard similarity level (Jaccard value = 0.28, marked with the blue dashed line).
Figure 4. Dendrogram recovered from a hierarchical cluster analysis (UPGMA) based on the Jaccard coefficient indicating floristic similarity between the sampling plots. Green and blue colors show the two large groups according to Jaccard similarity level (Jaccard value = 0.28, marked with the blue dashed line).
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Figure 5. Non-parametric multidimensional scaling analysis (NMDS) among harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of the P. patula-dominated forest in Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Dots represent the sampling points and the distance between them represents dissimilarity.
Figure 5. Non-parametric multidimensional scaling analysis (NMDS) among harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of the P. patula-dominated forest in Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Dots represent the sampling points and the distance between them represents dissimilarity.
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Figure 6. Venn diagram showing the species replacement between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of Ejido Llano Grande, Chignahuapan, Puebla, Mexico.
Figure 6. Venn diagram showing the species replacement between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of Ejido Llano Grande, Chignahuapan, Puebla, Mexico.
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Table 1. Families and genera with the highest species richness in the Pinus patula-dominated forest of the Ejido Llano Grande, Puebla, Mexico.
Table 1. Families and genera with the highest species richness in the Pinus patula-dominated forest of the Ejido Llano Grande, Puebla, Mexico.
FamilyNo. Genera/No. SpeciesGenusNo. Species
Asteraceae15/36Ageratina6
Poaceae9/10Quercus6
Orchidaceae7/7Pseudognaphalium4
Rosaceae6/7Roldana4
Ericaceae6/7Salvia4
Others52/71Others114
Table 2. Families and genera with the highest species richness by areas evaluated in the Ejido Llano Grande, Chignahuapan, Puebla, Mexico.
Table 2. Families and genera with the highest species richness by areas evaluated in the Ejido Llano Grande, Chignahuapan, Puebla, Mexico.
Harvested areas (HAs)FamilyNo. SpeciesGenusNo. Species
Asteraceae32Quercus6
Poaceae7Ageratina4
Fagaceae6Roldana4
Rosaceae5Pseudognaphalium4
Others53Others84
Tree structural retention (STRs)Asteraceae21Quercus5
Poaceae7Roldana4
Ericaceae7Ageratina4
Rosaceae6-
Lamiaceae6-
Others50Others84
Conservation areas (CAs)Asteraceae13Ageratina4
Rosaceae6Roldana3
Ericaceae5Quercus3
Polypodiaceae4Salvia3
Others47Others62
Table 3. Analysis of similarities (ANOSIM) between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Green cells indicate R values, blue cells indicate the p-value, and white cells indicate the total number of species per area evaluated.
Table 3. Analysis of similarities (ANOSIM) between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) of Ejido Llano Grande, Chignahuapan, Puebla, Mexico. Green cells indicate R values, blue cells indicate the p-value, and white cells indicate the total number of species per area evaluated.
HAsSTRsCAs
HAs990.00030.0003
STRs0.528970.0015
CAs0.7190.25675
Table 4. Species with the highest contribution to the total dissimilarity between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) and the mean values according to the SIMPER analysis.
Table 4. Species with the highest contribution to the total dissimilarity between harvested areas (HAs), structural tree retention stripes (STRs), and conservation areas (CAs) and the mean values according to the SIMPER analysis.
SpeciesContribution (%)Accumulated (%)Mean
HAs		Mean
STEs		Mean
CAs
Abies religiosa2.372.370.030.860.75
Chimaphila umbellata2.304.680.070.860.67
Geranium seemannii2.146.810.830.140.25
Monnina ciliolata1.918.720.200.690.50
Baccharis salicifolia1.8510.570.700.030.00
Roldana angulifolia1.8412.410.300.620.92
Baccharis conferta1.8114.220.670.380.00
Vaccinium leucanthum1.7615.980.370.930.83
Roldana barba-johannis1.7517.730.470.720.42
Bromus carinatus1.7219.450.430.590.33
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Pérez-Pardo, B.E.; Velázquez-Martínez, A.; Burgos-Hernández, M.; Reyes-Hernández, V.J. Floristic Diversity and Green-Tree Retention in Intensively Managed Temperate Forests: A Case Study in Puebla, Mexico. Forests 2024, 15, 920. https://doi.org/10.3390/f15060920

AMA Style

Pérez-Pardo BE, Velázquez-Martínez A, Burgos-Hernández M, Reyes-Hernández VJ. Floristic Diversity and Green-Tree Retention in Intensively Managed Temperate Forests: A Case Study in Puebla, Mexico. Forests. 2024; 15(6):920. https://doi.org/10.3390/f15060920

Chicago/Turabian Style

Pérez-Pardo, Brenda E., Alejandro Velázquez-Martínez, Mireya Burgos-Hernández, and Valentín J. Reyes-Hernández. 2024. "Floristic Diversity and Green-Tree Retention in Intensively Managed Temperate Forests: A Case Study in Puebla, Mexico" Forests 15, no. 6: 920. https://doi.org/10.3390/f15060920

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