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Structure and tree diversity in traditional Popoluca coffee agroecosystems in the Los Tuxtlas biosphere reserve, Mexico/Estructura y diversidad de arboles en agrosistemas cafetaleros popoluca, reserva de biosfera de Las Tuxlas, Mexico/Estrutura e diversidade de arvores em agroecossistemas cafeeiros popoluca na reserva de biofera Das Tuxlas, Mexico.

In Mexico, coffee is cultivated on the mountain slopes of the Sierra Madre Oriental facing the Gulf of Mexico, mainly in Hidalgo, Puebla, San Luis Potosi, Veracruz states and some districts in Tabasco; in the Pacific, it is cultivated in Chiapas, Colima, Jalisco, Nayarit and Oaxaca atates (Nolasco, 1985; Regalado-Ortiz, 2006) between 300 and 1,800masl. Coffee is grown on mountain slopes and in locations where northern, tropical and subtropical elements are found (Moguel and Toledo, 1999). According to Bartra (2003) 280,000 peasants produce coffee at smallholder scale in Mexico; 65% of the coffee peasants are indigenous, 183,000 of which own 2ha or less. In addition, there are 74,000 farms < 5ha. Particularly in indigenous areas, 41% of the area occupied by coffee agroecosystems is present in tropical rain forests, 23% in pine and oak forest, 21% in low deciduous forest and 15% in deciduous forest. Traditional coffee agroecosystems are considered to help maintain diversity because they conserve different forest strata (Miranda and Hernandez, 1963; Bartra, 2003). Moreover, the use of shade trees, such as 'solerillo' or 'xochicoahuitl' (Cordia alliodora) and different species of 'chalahuite' (Inga spp.), allows peasants to exploit several forest products and helps conserve orchids and other vascular epiphytes, along with birds and arthropods (Perfecto et al., 1996; Moguel and Toledo, 1999; Villavicencio and Valdez, 2003; Cruz et al., 2004; Hietz, 2005; Solis-Montero et al. 2005; Bandeira et al., 2005; Soto-Pinto et al., 2007). Similarly, within different coffee agroecosystems, environmental factors such as soil and water, together with shadow management, diversification of the tree canopy and use of cover legumes can improve coffee yields, while tree density can adversely affect coffee quality (Skovmand Bosselman et al., 2009). Also, as native trees are preserved, the role of natural regeneration could be important for the structure, floristic composition, richness and diversity of tree species (Godinez-Ibarra and Lopez-Mata, 2002; Philpott et al., 2008).

The state of Veracruz is second, after Chiapas, in coffee production in Mexico, by number of peasants and yield. Around 30% of the area dedicated to coffee is located between 300 and 800masl; these areas are considered marginal because they lie outside of the ideal agroecological zone for coffee production and yield, and quality are low (Moguel and Toledo, 1999). In the Sierra of Santa Marta, under the above mentioned conditions, management by the Popoluca peasants is similar to the diversified poly-culture structure (Franco, 2007; Hernandez-Martinez, 2008; Williams-Linera and Lopez-Gomez, 2008), which can increase [beta] diversity. However, the prolonged coffee production crisis (Martinez, 1997) has forced these peasants to eliminate many coffee agroecosystems and replace them with cattle farms, which has had a negative impact on the soil, biological diversity, production and productivity, as well as having an impact on processes such as the water, carbon and nitrogen cycles (Sanchez et al., 2003; Bandeira et al., 2005). Due to its ecological importance, the tree structure and diversity in this type of agroecosystem must be studied in greater detail, as has been done for birds and insects (Gould and Guerrero-Rivera, 2006; Lopez-Gomez et al., 2007; Oijen et al., 2010). This knowledge is essential to understand how the system operates to achieve a sustainable use of the natural resources associated with coffee production. This information is particularly relevant given the fast decline of natural resources at the local and global level, because these types of agroecosystems constitute important diversity reserves that have only recently been studied with the level of scientific rigour that they deserve (Vandermeer, 2011). The goal of this study was to analyse the tree structure and biological diversity of coffee agroecosystems established along an altitudinal gradient between 450 and 1,100masl within the buffer area of Los Tuxtlas Biosphere Reserve, Veracruz.

Materials and Methods

Study area

The study area is located in the Popoluca community of Ocotal Chico, Soteapan, Veracruz, at 18[degrees]18'31"N and 94[degrees]52'26"W, and covers 1361ha (Graciano, 2004). It is part of the buffer area of Los Tuxtlas Biosphere Reserve in the Sierra of Santa Marta (Siemens, 2004; Figure 1) and has a volcanic origin, with igneous rocks and andesitic or alkaline basaltic lava from the quaternary period. Its physiography includes five morphoedaphological units that were formed by mountains with slopes covered by volcanic cones (Siemens, 2004). The area is located in the sub-basin of the Huazuntlan River, within the Coatzacoalcos river basin. The vegetation includes 1) tropical pine forest, which is dominated by Pinus oocarpa and five oak species; 2) tropical semideciduous forest (TSF) dominated by Brosimum alicastrum, Cedrela odorata, Inga leptoloba and Luehea speciosa, among others; 3) tropical rainforest (TRF) dominated by Omphalea oleifera, Quercus sp., Terminalia amazonia and Calophyllum brasiliense; and 4) deciduous forest (DF) dominated by Alfaroa mexicana, Liquidambar styraciflua, Quercus sp. and Ulmus mexicana (Castillo-Campos and Laborde, 2004).

Agroecosystem selection and measurements

Based on participatory workshops, a list of 69 peasants was compiled. Their agroecosystems were located in areas previously occupied by 1) TSF (TSF coffee) between 450 and 600masl, with warm humid climate, summer precipitation (Garcia, 1988) and Acrisols; 2) TRF (TRF coffee) between 600 and 800masl, with warm humid climate, rainfall throughout the year and Acrisols; and 3) DF (DF coffee) between 800 and 1000masl, with semi-warm wet climate, rainfall throughout the year and Andosols. All soil types are highly susceptible to erosion (Mariano and Garcia, 2010). All coffee agroecosystems studied are located in slopes that vary between 15 and 60%, and within them some of the trees from the original vegetation were preserved. Using a random number table, 30 agroecosystems were chosen along the altitudinal gradient (Scheaffer and Ott, 1987), 10 from each section of the altitudinal gradient. Farm size varied based on the requests that each farmer made to the PROCEDE (Ejido and Community Right Program) of the National Agricultural Records. On each farm, a 400 [m.sup.2] (20 x 20m) site was marked and divided into four 10 x 10m (100 [m.sup.2]) quadrats that, in turn, were subdivided into eight 5 x10m (50 [m.sup.2]) quadrats. Four of these rectangles were randomly chosen and the height and cover of shrub and herbaceous strata were measured. For all the trees in the sampling area, the diameter at breast height (DBH) was measured at 1.3m above soil surface, and the total height and trunk height (up to the first branch) were measured using a Haga altimeter. Based on these data, basal area was calculated as BA = ([pi] x [D.sup.2])/4, where BA: basal area and D: DBH. The cover was quantified based on perpendicular measurements of the vertical projection of tree crowns, and the corresponding area was calculated as CC= ([([D.sub.1] + [D.sub.2]/4).sup.2])[pi] (MueUer-Dombois and Ellenberg, 1974). The distance between trees was measured with a measuring tape in order to know the horizontal distribution of species. The vegetation structure was analysed based on the relative density values (RDVs), frequency (FR) and relative dominance (DOR) based upon DBH. All relative values were calculated by dividing the number, frequency and dominance of a species by the total number, frequency and dominance of all species. The importance value was calculated as the sum of the three values (IV = RDV + DOR + FR), and this value was divided by three to obtain the relative importance value (RIV) (MuUer-Dombois and Ellenberg, 1974; Moreno, 2001). To quantify the floristic similarity, the S0rensen coefficient (MueUer-Dombois and Ellenberg, 1974) was calculated with the formula IS = (2C/A + B) x 100, where A is the number of species in community A, B is the number of species in community B, and C is the number of species in both communities. Similarly, the complementarity index was calculated (Moreno, 2001). First, the total richness was calculated for all sites with the formula [S.sub.AB] = a + b - c, where a: number of species in site A, b: number of species in site B and c: number of species common to both sites. Next, the number of species unique to each site was calculated as [U.sub.AB] = a + b - 2c. The complementarity index was calculated based on the values obtained above with the formula [C.sub.AB] = [U.sub.AB]/[S.sub.AB], where [U.sub.AB] is the species unique to each site and [S.sub.AB] is the total richness of all sites. The value of the index varies between 0 and 1, where 0 represents identical sites, and 1 indicates entirely different sites. By multiplying the value by 100, a percentage was obtained. Species richness and diversity was analysed with the ShannonWiener, Simpson and Fisher diversity indexes using the software Estimates 8.2.0 (Colwell, 2009).

Coffee agroecosystems structure

The vegetation structure was graphically represented with vertical and horizontal profile diagrams. To recognize the floristic composition, voucher specimens for all the plant species that were present on the coffee agroecosystems were collected. Species that were not at the sites but had flowers and/or fruit were also collected, although they were not included in the analysis. As the elevation increased, only plants that had not been previously observed were collected. Voucher specimens were deposited in the herbarium at the Instituto de Investigaciones Biologicas, Universidad Veracruzana in Xalapa, Veracruz, Mexico.

Results and Discussion

General structure and floristic composition of coffee agroecosystems

Coffee agroecosystems had four strata: herbaceous, shrub, low trees and tall trees, one layer less than those observed by Soto-Pinto et al. (2000). Due to peasant management the herbaceous layer had a low cover, which favoured the presence of some species with economic value and abundant leaf litter; additionally, weed control is carried out mainly by machete (66.6%), only 16.6% with herbicide, while another 16.6% use both (Franco, 2007). In this stratum, the dominant plants were shrub hot pepper (Capsicum annuum var. annuum), 'barbasco' (Dioscorea composita), cucumber (Cucumis sativus), 'tomatillo ' (Solanum pimpinellifolium), bean (Phaseolus spp.), hot pepper fruits (Capsicum annuum), goosefoot (Chenopodium sp.), Caladium bicolor, Colocasia sp., Ceratozamia sp. and 'camedor' palm (Chamaedorea spp.), which was introduced through government programs and the Sierra de Santa Marta A. C. project.

In TSF coffee agroecosystems the shrub stratum was dominated by different varieties of Coffea arabica, including Mundo Novo (80.7%), Robusta (8.7%), Caturra (6.4%) and Criolla (4.1%). In TRF coffee plantation, Mundo Novo (79.8%), Caturra (7.5%), Robusta (6.8%) and Criolla (5.9%) were present. Finally, in DF coffee agroecosystems, Caturra (50%), Garnica (28.1%), Mondo Novo (10.7%) and Criolla (11.23%) dominate. Coffee plants were planted in 2.5 x 2.5m and 2.0 x 2.0m grids, for a density of 1600-2,500 shrubs/ha, similar to what was found by Soto-Pinto et al. (2000) and Peeters et al. (2003) in different places of Chiapas, Mexico. However, accordingly to Descroix and Wintgens (2004), density for coffee plantations under shade must be 1250-1600 plants/ha with distances of 2.8 x 2.8 to 3.0 x 3.0 for Robusta varieties, and 1100-1600 plants/ ha for Arabica; that is to say, 3 x 3 to 2.5 x 2.5m. In this stratum, some species, such as Mexican pepper leaf (Piper sanctum) and 'platanillo' (Heliconia curtispatha) were not eliminated because their economic importance.

The floristic composition at the 30 study sites comprised 51 tree species. The most important were I. vera Willd (RIV = 26.42), Cordia alliodora (RIV = 10.59), Cecropia obtusifolia (7.40), Heliocarpus appendiculatus (6.85) and 23 herbaceous species. Forty-four families were identified (Table I); the most numerous were Mimosaceae (seven species), Asteraceae (six species), Fabaceae (six species) and Myrtaceae (four species). I. vera had the highest RIV along the altitudinal gradient because peasants consider it to be a tree with multiple uses: it does not lose its foliage in the dry season, produces firewood and provides more cover. Romero-Alvarado et al. (2002) found that the presence of Inga species does not improves the quality of coffee. Furthermore, using a parameterisation model, VanOijen et al. (2010) found that coffee yield tends to decrease with tree density in different coffee plantations in Central America, even in the presence of N-fixing trees, a similar phenomenon as was observed by Skovmand Bosselman et al. (2009) in Colombia. Importantly, although all species provide shade, the peasants conserve species like Vochysia guatemalensis (it has three different uses), C. odorata and Swietenia macrophylla because they sell the wood or use them for construction (they cover between 37-45% of the sites). Fruit trees cover 26-31% of the sites, outstanding among them Annona reticulata, Inga jinicuil and Byrsonima crassifolia (this one with three different uses). This Activity is similar to that observed by Rice (2011) in Peruvian and Guatemalan coffee plantations. It is noteworthy that, similar to Peruvian and Guatemalan peasants survival, Popoluca peasant survival depends not only on coffee agroecosystems (22%), but also other incomes such as government programs (52%), off-farm labor (17%) and livestock sales (9%) (Franco, 2007). In San Fernando, near the study area, socioeconomic variables influence ecological ones and modernization might have a negative effect in traditional coffee agroecosystems diversity (Potvin et al., 2005).

The structure: floristic composition, vertical strata, spatial distribution and diversity of the coffee agroecosystems studied followed similar patterns to those observed by Perfecto et al. (1996) and Soto-Pinto et al. (2000) in Chiapas; Bandeira et al. (2005) in the Chinantec region, Oaxaca; and Hernandez-Martinez (2008) in Coatepec, Veracruz. Moreover, local management and knowledge of agroecosystems play a fundamental role in the selection of the species that will be part of these systems because each peasant follows a different strategy to structure the coffee agroecosystem, altogether with a vast knowledge of local environmental conditions. We found 51 different tree species (345 individuals) in the studied sites, 60 to 85% fewer than reported in similar agroecosystems and vegetation types studies in Veracruz (Sanchez et al., 2003; Villavicencio and Valdez, 2003; WilliamsLinera et al., 2005; Lopez-Gomez et al., 2007). We collected 44 different families of plants in the whole study area, representing 84 different plant species, of which 64 are trees. That is, twice the plant families and 28% more trees than reported by Peeters et al. (2003) in Paredon, Chiapas. Additionally, the coffee agroecosystems studied conserved 25% more species, or at least the same number of species, as compared with some TSFs in Puerto Rico (Bandeira et al., 2005; Gould and Guerrero-Rivera, 2006).

The horizontal structure of all the coffee agroecosystems studied was similar; 80% of the tree species displayed a random distribution, and only 20% displayed a uniform one (Figure 2). Height ranges 5-35m, and it can be deduced that the more or less complex tree structure of the agroecosystems can help as a refuge for a diversity of birds, insects, and microorganisms (Philpott and Bichier, 2012; Jacinto, 2012; Retama et al., 2014). It is also important that the age of coffee plantations is 16-40 years old, the older being located at higher elevations, while coffee agroecosystems closer to villages are the younger ones, generally with a better management.

For TSF coffee agroecosystems (Table II), height was 0.6-26.0m. The tallest species were Acosmium panamense ('guayacan', 12m), Cecropia obtusifolia (trumpet tree, 26m), Cedrela odorata (cedar, 19m), Cordia alliodora ('solerillo', 20m), Gliricidia sepium (13m), Heliocarpus appendiculatus ('jonote', 15m), Inga jinicuil (22m), I. vera ('chalahuite', 26m) and Trema micrantha ('mupi' or 'ixpepe', 26m). Seventeen tree species (97 individuals) were identified on these coffee agroecosystems. The species with the highest RIVs were A. panamense, C. obtusifolia, C. odorata, Cojoba arborea ('canamazo'), C. alliodora, H. appendiculatus, I. vera, Pimenta dioica (allspice) and T. micrantha. The importance value for I. vera was twice as large as the importance value of C. alliodora. The species with the lowest RIVs were Citrus sinensis, Chrysophyllum cainito, Carica papaya, Pachira aquatica and Tephrosia sp. (introduced). The species with the highest cover were I. jinicuil, with 80.3 [m.sup.2], greater than that of I. vera (69.3 [m.sup.2]) despite having a lower density, B. crassifolia (68.7 [m.sup.2]), C. alliodora (64.5), G. sepium (63.4) and A. panamense (45.6 [m.sup.2]). A total of 37 species were identified from the different strata.

In the TRF coffee agroecosystems (Table III), 18 tree species (115 individuals) were identified. The maximum height was 35m, and the minimum 4.5m. The tallest species were Apeiba tibourbou (18m), Calophyllum brasiliense (35m), C. alliodora (32m), Hirtella triandra (26), I. jinicuil (25m), I. vera (26, Luehea speciosa (17), Pimenta dioica (20) and V. guatemalensis (18). The species with the highest RIVs were Apeiba tibourbou ('palo gusano' or 'papachote'), Citrus sinensis (sour orange), C. alliodora, Inga jinicuil (pod), I. vera, P. dioica, T. micrantha and Vochysia guatemalensis ('corpo'). The species with the lowest importance values were Coccoloba uvifera (sea grape), Citrus aurantifolia (lime) and Swietenia macrophylla (mahogany). The species with the greatest cover were A. tibourbou (151.66 [m.sup.2]), C. brasiliense (103.86), C. alliodora (51.54), Hirtella triandra (55.41), I. jinicuil (59.20) and L. speciosa (77.47 [m.sup.2]). These coffee agroecosystems had a total of 36 species.

In the areas with DF coffee agroecosystems (Table IV) 16 tree species (133 individuals) were observed, with a minimum height of 4.2 and a maximum of 32m. The tallest trees were A. reticulata (20m), Cecropia obtusifolia (18), H. appendiculatus (18), H. triandra Sw (14), I. jinicuil (30), I. vera (32), T amazonia (31), T micrantha (18) and V guatemalensis (18). The species with the highest RIVs were I. vera, T. micrantha, T amazonia, I. jinicuil, C. obtusifolia, V. guatemalensis, C. odorata and L. guatemalensis. The species with the lowest RIVs were Bursera simaruba (copper wood), L. guatemalensis ('gusanillo' or 'palo blanco'), Spondias mombin (yellow mombin) and Tectona grandis (introduced). The species with the greatest cover were A. reticulata L. (93.3 [m.sup.2]), T. amazonia (75.9), T. micrantha (55.4) and I. vera (50.7 [m.sup.2]). On these coffee agroecosystems, 31 species were collected from the different strata.

Structurally, the species with the highest importance value along the altitudinal gradient were I. vera, A. tibourbou, C. alliadora and T. micrantha. The first two species also dominate coffee agroecosystems in the Chinantec region in Oaxaca (Bandeira et al., 2005). The type II structural pattern of these species suggests the existence of disturbed areas in an advanced phase of tree gap planting (Martinez-Ramos and Alvarez-Buylla, 1995). As observed in the study by Lopez-Gomez and Williams-Linera (2006) on the coffee agroecosystems of Ocotal Chico, no important structural differences existed because the peasants were interested in species composition, not in increasing the height or basal area of the trees. In addition to I. vera, other species that were highlighted in Lopez-Gomez and Williams-Linera (2006) are Citrus spp., Mangifera indica, Psidium guajava and Persea schiedeana. The first three were found in the present study. However, B. crassifolia, C. alliadora, I. jinicuil, L. speciosa and T. micrantha displayed greater cover and lower density.

Population structure

Based on the diameter class distribution of species with a higher importance value, some structural patterns (sensu Martinez-Ramos and AlvarezBuylla, 1995) were distinguished. For TSF coffee agroecosystems, I. vera and C. alliodora displayed a type II pattern, which is characterised by a higher frequency of intermediate size individuals and a lower frequency of older individuals. T. micrantha follows a type III pattern, with small, intermediate and large individuals. C. obtusifolia and A. panamense did not display any defined structural patterns (Figure 3). In TRF coffee agroecosystems, I. vera and C. alliadora followed a type II pattern, but V. guatemalensis was characterised by a type III pattern, with small, intermediate and large individuals. I. jinicuil and A. tibourbou did not show a defined structural pattern (Figure 4). In DF coffee agroecosystems, I. vera, T. micrantha and I. jinicuil displayed a type II pattern, and T. amazonia, and C. obtusifolia did not have a defined structural pattern (Figure 5). The horizontal tree distribution was heterogeneous along the gradient as a result of the topological arrangement and management conducted by peasants (Figure 2). The population structure of C. alliadora and V. guatemalensis is due because their use is centered on diameter classes for home construction and planks, respectively.

Floristic similarity

According to the Sorensen index, the coffee agroecosystems that were established in TSF and DF had 21% similarity and shared seven species: C. obtusifolia, C. odorata, H. appendiculatus, I. jinicuil, I. vera, P dioica and T micrantha. The agroecosystems that were located in TRF and DF were 21% similar and had seven species in common: H. triandra, I. jinicuil, I. vera, P. dioica, S. mombin, T. micrantha and V. guatemalensis. Coffee agroecosystems located in TSF and TRF displayed 30% similarity and had 11 common species: C. annum var. glabriusculum, C. sinensis, C. alliodora, Erythrina americana, I. jinicuil, Inga punctata, Inga marginata, I. vera, P. dioica, T. micrantha and Willardia schiedeana. The indexes of floristic similarity were low; that is to say, the different coffee agroecosystems have high replacement rates due to the decisions peasants made about plants they used in each section of the altitudinal gradient, a phenomenon also reported by Williams-Linera and Lopez-Gomez (2008) and by Rice (2011) for fruit species. This observation is remarkable for the case of TSFs, which are located closest to dwellings. In other areas of Veracruz, the values were even lower (Williams-Linera and Lopez-Gomez, 2008). The mean floristic similarity was 12%, more than twice that found by Guiracocha et al. (2001) in cacao agroforestry systems in Costa Rica. Likewise, Godinez-Ibarra and Lopez-Mata (2002) reported an intermediate similarity, with a low number of shared species, for three TSF samples.

Species richness, diversity and complementarity index

Along the altitudinal gradient, 345 individuals were recorded (60 tree and 23 herbaceous species) within 12000 [m.sup.2]. The greatest tree richness (44.5%) occurred on coffee agroecosystems that were located in TSFs. For these agroecosystems, the Shannon-Wiener diversity index varied between 3.39 and 1.89, the Simpson index ranged between 61.95 and 31.1 and Fisher's alpha varied between 57.8 and 27.35. The coffee agroecosystems that presents higher diversity values are those located near dwellings. These values confirm the greater biological diversity of these systems (Table V).

The complementarity in species composition for the coffee agroecosystems that were located in TSFs and DFs was 88%; those located in TRFs and DFs had the same value. For agroecosystems located in TSFs and TRFs, complementarity was 82%, similar to those obtained by Williams-Linera et al. (2005) and Lopez-Gomez et al. (2007) in deciduous forest and coffee agroecosystems of central Veracruz. Similarly, Villavicencio and Valdez (2003) found a 58% floristic similarity and 42% different species for coffee agroecosystems established in TSFs and TRFs in San Miguel, near Cordoba, Veracruz. In this same area, these authors observed greater evenness in the tree structure of rustic coffee agroecosystems established in TSF. Our results indicate a high replacement rate and, therefore, a high p diversity, which confirms that moderate disturbances resulting from human management, may have increased the species richness, although the original vegetation diversity was not reached (Williams-Linera et al., 2005; Philpott et al., 2008a).

Furthermore, the exclusive species found in each coffee agroecosystem studied herein also indicate a high diversity (Table VI) and confirm the influential role of traditional peasants in preserving and even increasing diversity. Their management practices seem to be fundamental for conservation of natural resources in the area. It should be noted that, contrary to what was found by Philpott et al. (2008b) in Sumatra, Popoluca peasants conserve more native species along the altitudinal gradient (of those mandatory to be certified by programs like the Smithsonian Migratory Bird Center or 'Bird Friendly'). This diversity could be the basis for local programs aimed to conserve trees, but also birds, insects, microorganisms, biogeochemical cycles and give more resilience to the agricultural matrix (sensu Perfecto and Vandermeer, 2008). For instance, tree species such as A. panamense, C. brasiliense, T. amazonia, T. micrantha and V. guatemalensis in the lower and upper tree strata can diversify the productivity of coffee agroecosystems, giving emphasis to the use of evergreen species. This diversity contributes to soil structural stability because of the high susceptibility to erosion (Juarez, 2008; Cruz, 2009). In the lower tree stratum, C. alliodora, B. crassifolia, C. papaya, C. sinensis, C. cainito, I. jinicuil, P. dioica and S. mombin are important species. In the herbaceous stratum, some species, such C. annuum var. annuum, Chenopodium sp., C. sativus and S. pimpinellifolium, could be used as garden produce, and species such as Colocasia bicolor, Colocasia sp., Chamaedorea sp. and Ceratozamia sp. could be used as ornamentals.

Conclusions

Four strata were found in the 30 coffee agroecosystems studied. Inga vera had the highest importance value; however, we found 84 different plants, 64 of which are trees. Of those whose uses could be documented, we found one to three different uses, timber, fruits and medicinal being remarkable. Coffee agroecosystems located near dwellings (TSD coffee) have higher diversity values; however, its tree density is lower (97 individuals) than in TRF coffee (115 individuals) and in DF coffee (133 individuals). Tree height ranges 5-35m. Results show high diversity indices, even higher than in other areas of Chiapas, which is confirmed by the few species that all the coffee agroecosystems share, by the high replacement rate, and by the great number of exclusive species found at each coffee agroecosystem. All these confirm the fundamental role of peasant's knowledge and management in the selection of species and the structure of the agroecosystem, but also in increasing and in some cases improving diversity. Popoluca peasants conserve native species instead of exotics, of which only three species were found. With the information obtained, diversification and restoration programs could be organized based upon native tree richness and the participation of the Popoluca people. This will allow to structure agroecological matrices to improve production and productivity of agroecosystems, but also conserve birds, mammals, insects, microorganisms and the essential biogeochemical cycles.

Received: 09/02/2013. Modifies: 07/28/3014. Accepted: 07/29/2014.

Guadalupe Castillo Capitan. M.Sc. in Agricultural Sciences, Universidad Autonoma Metropolitana-Xochimilco (UAM-X), Mexico. Professor, Universidad Veracruzana (UV), Mexico.

Carlos H. Avila-Bello. Ph.D. in Agroecology, Colegio de Postgraduados (COLPOS), Mexico. Professor, Address: Facultad de Ingenieria en Sistemas de Produccion Agropecuaria, UV Acayucan, Veracruz. 96000, Mexico. e-mail: carlavila@uv.mx

Lauro Lopez-Mata. Ph.D. in Botany, University of North Carolina, USA. Professor, COLPOS. Montecillo, Mexico.

Fernando de Leon Gonzalez. Ph.D. in Soil Sciences, Institut National Agronomique (ParisGrignon), France. Professor, UAM-X, Mexico.

ACKNOWLEDGMENTS

The authors acknowledge the authorities and inhabitants of Ocotal Chico, Los Tuxtlas Biosphere Reserve, for permission and support, to A. Matias Santiago, G. Matias Gonzalez, P Gutierrez Albino and B. Matias Gonzalez; to J.L. Villasenor, Biology Institute, UNAM, for nomenclature update and revision of the floristic list; to the Program for Professorship Improvement (PROMEP), Secretary of Public Education, for funding project 103.5/04/1411 (PTC-59); and to Olga Ricalde Moreno for suggestions to improve the English language.

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TABLE I
FLORISTIC COMPOSITION OF THE COFFEE
AGROECOSYSTEMS IN OCOTAL CHICO, SOTEAPAN,
VER, MEXICO *

Family             Scientific name                  Use

Anacardiaceae      Astronium graveolens Jacq.       Timber
                   Mangifera indica L.              Fruit
                   Spondias mombin L.               Fruit
Annonaceae         Annona reticulata L.             Fruit, medicinal *
                   Rollinia mucosa (Jacq.)          Not documented
                     Baill.
Asteraceae         Ageratella sp.                   Not documented
                   Baltimora recta L.               Not documented
                   Critonia daleoides (DC.)         Medicinal
                   Montanoa sp.                     Medicinal
                   Sinclairia discolor              Not documented
                     Hook. & Arn.
                   Vernonia patens Kunth            Not documented
Bignoniaceae       Spathodea campanulata Beauv.     Shade
Bombacaeae         Pachira aquatica Aubl.           Medicinal
Boraginaceae       Cordia alliodora                 Timber
                     (Ruiz & Pav.) Oken
Burseraceae        Bursera simaruba (L.) Sarg.      Hedge, shade
Caricaeae          Carica papaya L.                 Fruit
Cecropiaceae       Cecropia obtusifolia Bertol.     Shade
Chrysobalanaceae   Hirtella triandra Sw             Medicinal
Combretaceae       Terminalia amazonia              Timber
                     (J. F. Gmel.) Exell
Cucurbitaceae      Sechium edule (Jacq.) Sw.        Edible
Euphorbiaceae      Acalypha microstachya Benth.     Medicinal
Fabaceae           Acosmium panamense               Timber
                     (Benth.) Yakovlev
                   Erythrina americana Mill.        Hedge, edible
                                                      (flowers)
                   Gliricidia sepium Stend.         Hedge, firewood
                   Lonchocarpus                     Shade
                     guatemalensis Benth.
                   Tephrosia sp.                    Temporal shade
                   Willardia schiedeana             Shade
                     (Schltdl.) F. J. Herm
Guttiferaceae      Calophyllum brasiliense          Timber,
                     Cambess.                         construction
Haemodoraceae      Xiphidium caeruleum Aubl.        Not documented
Hamamelidaceae     Liquidambar styraciflua L.       Shade
Heliconiaceae      Heliconia curtispatha            Not documented
                     Petersen
Hypericaceae       Vismia baccifera (L.)            Medicinal
                     Triana & Planch.
                   Vismia camaguey Sparague         Not documented
                     & L. Riley
Lamiaceae          Hyptis mutabilis                 Not documented
                     (L. Rich.) Briq.
Lauraceae          Ocotea verticillata Rohwer       Shade
Lasistemataceae    Lacistema aggregatum Rusby       Not documented
                     (P. J. Bergiev)
Malpighiaceae      Byrsonima crassifolia            Shade, fruit,
                     (L.) Kunth                       medicinal
                   Malpighia glabra L.              Not documented
                   Tetrapterys schiedeana           Not documented
                     Schltdl. & Cham.
Malvaceae          Sida acuta Burm. f.              Medicinal
                   Sida cordiflolia L.              Not documented
                   Sida rhombifolia L.              Medicinal
Maranthaceae       Stromanthe acrochlamys           Not documented
                     (Woodson & Standley)
                     H. A. Kenn. & Nicolson
Melastomataceae    Adelobotrys adscendens           Not documented
                     (Sw.) Triana
                   Miconia argentea (Sw.) DC.       Handles for
                                                      tools, shade
Meliaceae          Cedrela odorata L.               Timber, shade
                   Swietenia macrophylla            Timber, shade
                     G. King
                   Trichilia havanensis Jacq.       Timber, handles
                                                      for tools
Mimosaceae         Zapoteca sp.                     Medicinal
                   Cojoba arborea (L.)              Timber, shade
                     Britton & Rose
                   Inga jinicuil Schltdl.           Shade, fruit
                     & Cham.
                   Inga punctata Willd.             Shade, firewood
                   Inga marginata Willd.            Shade, firewood
                   Inga vera Willd.                 Shade, firewood
                   Leucaena leucocephala            Shade, fruit
                     (Rose) S. Zarate
Myrtaceae          Calyptranthes lindeniana         Shade
                     O. Berg.
                   Eugenia acapulcensis Steud.      Shade, fruit,
                                                      medicinal
                   Eugenia capuli (Schltdl.         Fruit, shade
                     & Cham.) O. Berg.
                   Pimenta dioica (L.) Merr.        Spice, shade
Orchidaceae        Catasetum integerrimum Hook.     Ornamental
                   Sacoila lanceolata A. Rich       Ornamental
                   Vanilla planifolia G. Andrews    Ornamental
Palmae             Astrocaryum mexicanun            Edible
                     Liebm ex Mart.
Primulacaceae      Rapanea sp.                      Not documented
Polygonaceae       Coccoloba uvifera L.             Medicinal
Rubiaceae          Alibertia edulis (L. Rich)       Medicinal
                     A. Rich. ex. DC.
                   Chiococca alba (L.) Hitchc.      Not documented
Rutaceae           Citrus aurantifolia Swingle      Fruit, Shade
                   Citrus sinensis (L) Osbeck       Fruit, Shade
                   Zanthoxylum caribaeum Lam.       Shade
Salicaceae         Zuelania guidonia (Sw.)          Not documented
                     Britton & Millsp.
Sapindaceae        Allophylus cominia (L.) Sw.      Medicinal
                   Cupania glabra Sw.               Firewood
Solanaceae         Capsicum annum Var.
                     glabriusculum (Dunal)
                   Heiser & Pickersgill             Edible
                   Solanum pimpinellifolium L.      Edible
Sapotaceae         Chrysophyllum cainito L.         Fruit
                   Chrysophyllum mexicanum          Fruit, handles
                     Brandegee & Standl.              for tools
Surianaceae        Suriana maritima L.              Not documented
Thelypteridaceae   Thelypteris blanda C. F. Reed    Not documented
Tiliaceae          Apeiba tibourbou Aubl.           Medicinal
                   Heliocarpus appendiculatus       Not documented
                     Turcz.
                   Luehea speciosa Wild.            Timber, shade
Ulmaceae           Trema micrantha (L.) Blume       Bird feed
Verbenaceae        Tectona grandis L. f.            Timber
Vochysiaceae       Vochysia guatemalensis           Construction,
                     Donn. Sm.                        timber, shade

Family             Life form    Original
                                vegetation
                                type

Anacardiaceae      Tree         DF
                   Tree         TRF
                   Tree         TRF-DF
Annonaceae         Tree         DF
                   Tree         TRF

Asteraceae         Herb         DF
                   Herb         DF
                   Shrub        TRF
                   Herb         TRF
                   Herb         TRF

                   Shrub        TRF
Bignoniaceae       Tree **      TRF
Bombacaeae         Tree         TSF
Boraginaceae       Tree         TSF-TRF

Burseraceae        Tree         DF
Caricaeae          Tree         TSF
Cecropiaceae       Tree         TSF-DF
Chrysobalanaceae   Tree         TRF-DF
Combretaceae       Tree         DF

Cucurbitaceae      Herb         TSF-TRF-DF
Euphorbiaceae      Tree         TRF
Fabaceae           Tree         TSF

                   Tree         TSF-TRF

                   Tree         TSF
                   Tree         DF

                   Shrub **     TSF
                   Tree         TSF-TRF

Guttiferaceae      Tree         TRF

Haemodoraceae      Herb         TRF
Hamamelidaceae     Tree         DF
Heliconiaceae      Herb         TSF

Hypericaceae       Tree         TSF

                   Tree         DF

Lamiaceae          Herb         TRF

Lauraceae          Tree         DF
Lasistemataceae    Tree         DF

Malpighiaceae      Tree         TSF

                   Shrub        TSF
                   Woody vine   DF

Malvaceae          Shrub        TSF
                   Shrub        TRF
                   Shrub        TRF
Maranthaceae       Herb         TSF

Melastomataceae    Vine         DF

                   Tree         TRF

Meliaceae          Tree         TSF-DF
                   Tree         TRF

                   Tree         TSF

Mimosaceae         Tree         TSF
                   Tree         TSF

                   Tree         TSF-TRF-DF

                   Tree         TSF-TRF
                   Tree         TSF-TRF
                   Tree         TSF-TRF-DF
                   Tree         TRF

Myrtaceae          Tree         DF

                   Tree         TSF

                   Tree         TSF

                   Tree         TSF-TRF-DF
Orchidaceae        Epiphyte     DF
                   Herb         TSF
                   Epiphyte     TRF
Palmae             Tree         DF

Primulacaceae      Tree         DF
Polygonaceae       Tree         TRF
Rubiaceae          Tree         TSF

                   Tree         TSF
Rutaceae           Tree         TRF
                   Tree         TSF-TRF
                   Tree         TSF
Salicaceae         Tree         DF

Sapindaceae        Tree         DF
                   Tree         TSF
Solanaceae

                   Herb         TSF-TRF
                   Herb         TSF
Sapotaceae         Tree         TSF
                   Tree         TSF

Surianaceae        Shrub        TRF
Thelypteridaceae   Herb         DF
Tiliaceae          Tree         TRF
                   Tree         TSF-DF

                   Tree         TRF
Ulmaceae           Tree         TSF-TRF-DF
Verbenaceae        Tree **      DF
Vochysiaceae       Tree         TRF-DF

* Medicinal uses were documented based upon Leonti (2002).
** Introduced.

TABLE II
TREE STRUCTURE OF COFFEE AGROECOSYSTEMS
LOCATED IN THE TROPICAL SEMIDECIDUOUS RAINFOREST
(450-600M) IN OCOTAL CHICO *

Species                  Number of       Cover      Height (m)
                        individuals   ([m.sup.2])

Acosmium panamense           3           45.6          10.6
Byrsonima crassifolia        1           68.6           15
Carica papaya                1           6.61           3
Cojoba arborea               2            0.1          0.7
Cecropia obtusifolia         5           23.93         14.1
Cedrela odorata              3           17.7          16.2
Citrus sinensis              3            4.4          4.9
Cordia alliodora            11           64.5          17.3
Chrysophyllum cainito        1           2.14           3
Gliricidia sepium            2           63.4          12.5
Heliocarpus                  3           23.9          8.6
  appendiculatus
Inga vera                   45           69.3          16.4
Inga jinicuil                4           80.4          13.3
Pachira aquatica             1            2.0          2.5
Pimenta dioica               4           11.4          7.1
Tephrosia sp.                1           0.33           2
Trema micrantha              7           30.8          8.7
n=17                        97

Species                 Basal area    Absolute    Relative
                        ([m.sup.2])   frequency   density

Acosmium panamense        218.16      0.3 (30%)     0.03
Byrsonima crassifolia     283.52      0.1 (10%)     0.01
Carica papaya              19.63      0.1 (10%)     0.01
Cojoba arborea             0.12       0.3 (30%)     0.02
Cecropia obtusifolia      263.59      0.3 (30%)     0.05
Cedrela odorata           245.13      0.2 (20%)     0.03
Citrus sinensis            84.94      0.1 (10%)     0.03
Cordia alliodora          234.32      0.5 (50%)     0.11
Chrysophyllum cainito      50.26      0.1 (10%)     0.01
Gliricidia sepium         188.69      0.2 (20%)     0.02
Heliocarpus               333.29      0.2 (20%)     0.03
  appendiculatus
Inga vera                 261.74      1 (100%)      0.46
Inga jinicuil             176.71      0.2 (20%)     0.04
Pachira aquatica           7.06       0.1 (10%)     0.01
Pimenta dioica             44.76      0.3 (30%)     0.04
Tephrosia sp.              7.06       0.1 (10%)     0.01
Trema micrantha           157.73      0.3 (30%)     0.07
n=17                      2576.80        4.4        1.00

Species                 Relative    Relative
                        frequency   dominance

Acosmium panamense        0.06        0.08
Byrsonima crassifolia     0.02        0.11
Carica papaya             0.02        0.00
Cojoba arborea            0.06        0.00
Cecropia obtusifolia      0.06        0.10
Cedrela odorata           0.04        0.09
Citrus sinensis           0.02        0.03
Cordia alliodora          0.11        0.09
Chrysophyllum cainito     0.02        0.02
Gliricidia sepium         0.04        0.07
Heliocarpus               0.04        0.12
  appendiculatus
Inga vera                 0.22        0.10
Inga jinicuil             0.04        0.06
Pachira aquatica          0.02        0.003
Pimenta dioica            0.06        0.017
Tephrosia sp.             0.02        0.003
Trema micrantha           0.06        0.06
n=17                      1.00        1.00

Species                 IV.     RIV

Acosmium panamense      0.18    6.12
Byrsonima crassifolia   0.14    4.76
Carica papaya           0.04    1.35
Cojoba arborea          0.08    2.96
Cecropia obtusifolia    0.22    7.40
Cedrela odorata         0.17    5.71
Citrus sinensis         0.08    2.88
Cordia alliodora        0.31   10.59
Chrysophyllum cainito   0.05    1.75
Gliricidia sepium       0.13    4.64
Heliocarpus             0.20    6.85
  appendiculatus
Inga vera               0.79   26.42
Inga jinicuil           0.15    5.17
Pachira aquatica        0.03    1.19
Pimenta dioica          0.12    4.22
Tephrosia sp.           0.03    1.19
Trema micrantha         0.20    6.71
n=17                    3.00   100.00

* Reference area 4,000[m.sup.2] (10 sampling sites
of 400[m.sup.2]).

TABLE III
TREE STRUCTURE IN COFFEE AGROECOSYSTEMS
LOCATED IN THE TROPICAL RAINFOREST (600-800M)
IN OCOTAL CHICO

Species                    Number of       Cover      Height (m)
                          individuals   ([m.sup.2])

Apeiba tibourbou                2          151.6          18
Calophyllum brasiliense         1          103.9          35
Citrus aurantifolia             2           12.3          4.5
Citrus sinensis                 3            7.7          7.8
Coccoloba uvifera               1           12.3          26
Cordia alliodora               15           51.5         23.9
Hirtella triandra               1           55.4           6
Inga jinicuil                   5           59.2         15.6
Inga vera                      59           42.3         14.4
Leucaena leucocephala           3            6.8          6.2
Luehea speciosa                 3           77.5         12.6
Mangifera indica                1           17.3          7.5
Pimenta dioica                  4           42.3         10.5
Spathodea campanulata           1            3.9           5
Spondias mombin                 2            6.0           5
Swietenia macrophylla           2           11.2           6
Trema micrantha                 3           41.6          8.2
Vochysia guatemalensis          7           18.7          9.6
n=18                          115

Species                   Basal area    Absolute    Relative
                          ([m.sup.2])   frequency   density

Apeiba tibourbou           15614.54     0.1 (10%)     0.02
Calophyllum brasiliense      855.30     0.1 (10%)     0.01
Citrus aurantifolia           63.61     0.1 (10%)     0.02
Citrus sinensis              263.98     0.3 (30%)     0.03
Coccoloba uvifera            176.71     0.1 (10%)     0.01
Cordia alliodora             776.01     0.4 (40%)     0.13
Hirtella triandra           1017.87     0.1 (10%)     0.01
Inga jinicuil                589.64     0.4 (40%)     0.04
Inga vera                    376.10     1 (100%)      0.51
Leucaena leucocephala         34.90     0.1 (10%)     0.03
Luehea speciosa              732.21     0.2 (20%)     0.03
Mangifera indica             295.59     0.1 (10%)     0.01
Pimenta dioica               226.98     0.3 (30%)     0.03
Spathodea campanulata         95.03     0.1 (10%)     0.01
Spondias mombin               78.54     0.2 (20%)     0.02
Swietenia macrophylla        116.89     0.1 (10%)     0.02
Trema micrantha              143.13     0.3 (30%)     0.03
Vochysia guatemalensis       173.36     0.4 (40%)     0.06
n=18                       21630.47        4.4         1

Species                   Relative    Relative
                          frequency   dominance

Apeiba tibourbou            0.02        0.72
Calophyllum brasiliense     0.02        0.04
Citrus aurantifolia         0.03        0.00
Citrus sinensis             0.07        0.01
Coccoloba uvifera           0.02        0.01
Cordia alliodora            0.09        0.04
Hirtella triandra           0.02        0.05
Inga jinicuil               0.09        0.03
Inga vera                   0.23        0.02
Leucaena leucocephala       0.02        0.00
Luehea speciosa             0.05        0.03
Mangifera indica            0.02        0.01
Pimenta dioica              0.07        0.01
Spathodea campanulata       0.02        0.00
Spondias mombin             0.05        0.00
Swietenia macrophylla       0.02        0.01
Trema micrantha             0.07        0.01
Vochysia guatemalensis      0.09        0.01
n=18                        1.01        1.00

Species                    IV     RIV

Apeiba tibourbou          0.76    25.33
Calophyllum brasiliense   0.07     2.33
Citrus aurantifolia       0.05     1.78
Citrus sinensis           0.11     3.67
Coccoloba uvifera         0.04     1.33
Cordia alliodora          0.26     8.67
Hirtella triandra         0.08     2.67
Inga jinicuil             0.16     5.33
Inga vera                 0.76    25.33
Leucaena leucocephala     0.05     1.73
Luehea speciosa           0.11     3.67
Mangifera indica          0.04     1.33
Pimenta dioica            0.11     3.67
Spathodea campanulata     0.03     1.00
Spondias mombin           0.07     2.34
Swietenia macrophylla     0.05     1.70
Trema micrantha           0.11     3.58
Vochysia guatemalensis    0.16     5.33
n=18                      3.02   100.80

* Reference area 4,000[m.sup.2] (10 sampling sites
of 400[m.sup.2]).

TABLE IV
VEGETATION STRUCTURE OF COFFEE AGROECOSYSTEMS
LOCATED IN THE DECIDUOUS FORESTS (800-1000M) IN OCOTAL
CHICO *

Species                  Number of       Cover      Height
                        individuals   ([m.sup.2])    (m)

Annona reticulata             1          93.3        20
Astrocarium mexicanun         1          13.5         5
Bursera simaruba              2           1.7         2.8
Cecropia obtusifolia          3          51.6        14.6
Cedrela odorata               4           3.1         4.2
Heliocarpus                   2          17.7         6.9
  appendiculatus
Hirtella triandra             3          48.7         4.5
Inga jinicuil                 6          45.4        13.5
Inga vera                    86          50.8        11.8
Lonchocarpus                  3           0.3        12
  guatemalensis
Pimenta dioica                2           7.6         6
Spondias mombin               1           4.5         5.8
Tectona grandis               1          49.1         6
Terminalia amazonia           2          75.9        31
Trema micrantha              11          55.4        12.4
Vochysia guatemalensis        5          20.9        14.3
n = 16                      133

Species                 Basal area    Absolute    Relative
                        ([m.sup.2])   frequency   density

Annona reticulata          764.53     0.1 (10%)     0.01
Astrocarium mexicanun      314.16     0.1 (10%)     0.01
Bursera simaruba             8.81     0.1 (10%)     0.02
Cecropia obtusifolia       481.75     0.2 (20%)     0.02
Cedrela odorata             46.86     0.2 (20%)     0.03
Heliocarpus                 95.03     0.2 (20%)     0.02
  appendiculatus
Hirtella triandra          838.10     0.1 (10%)     0.02
Inga jinicuil              373.25     0.2 (20%)     0.05
Inga vera                  351.52     1 (100%)      0.65
Lonchocarpus                73.39     0.1 (10%)     0.02
  guatemalensis
Pimenta dioica              94.17     0.2 (20%)     0.02
Spondias mombin            314.16     0.1 (10%)     0.01
Tectona grandis            415.47     0.1 (10%)     0.01
Terminalia amazonia       1541.34     0.1 (10%)     0.02
Trema micrantha            341.87     0.6 (60%)     0.08
Vochysia guatemalensis     264.75     0.2 (20%)     0.04
n = 16                    6319.22     3.6           1

Species                 Relative   Relative
                        frequency  dominance

Annona reticulata         0.03       0.12
Astrocarium mexicanun     0.03       0.05
Bursera simaruba          0.03       0.02
Cecropia obtusifolia      0.06       0.06
Cedrela odorata           0.06       0.02
Heliocarpus               0.06       0.03
  appendiculatus
Hirtella triandra         0.03       0.13
Inga jinicuil             0.06       0.05
Inga vera                 0.28       0.07
Lonchocarpus              0.03       0.02
  guatemalensis
Pimenta dioica            0.06       0.02
Spondias mombin           0.03       0.03
Tectona grandis           0.03       0.06
Terminalia amazonia       0.03       0.23
Trema micrantha           0.17       0.04
Vochysia guatemalensis    0.06       0.03
n = 16                    1          0.98

Species                 IV     RIV

Annona reticulata       0.13   4.33
Astrocarium mexicanun   0.09   3.00
Bursera simaruba        0.07   2.33
Cecropia obtusifolia    0.14   4.67
Cedrela odorata         0.11   3.67
Heliocarpus             0.08   2.67
  appendiculatus
Hirtella triandra       0.18   6.00
Inga jinicuil           0.16   5.33
Inga vera               1.00   33.33
Lonchocarpus            0.07   2.33
  guatemalensis
Pimenta dioica          0.10   3.33
Spondias mombin         0.07   2.33
Tectona grandis         0.11   3.67
Terminalia amazonia     0.28   9.33
Trema micrantha         0.29   9.67
Vochysia guatemalensis  0.13   4.33
n = 16                  3.01   100.33

* Reference area 4000 [m.sup.2] (10 sampling sites of
400 [m.sup.2]).

TABLE V
BIOLOGICAL DIVERSITY INDEX FOR COFFEE
AGROECOSYSTEMS IN OCOTAL, CHICO

Site      Type of      Fisher's   Shannon's   Simpson's
       agroecosystem    alpha       index       index

1       TSF coffee       43.4       1.89          1
2       TSF coffee       57.8       2.42        61.95
3       TSF coffee        44        2.73        42.71
4       TSF coffee        40         2.9        37.13
5       TSF coffee      39.35       3.04        34.95
6       TSF coffee      37.72       3.15        33.62
7       TSF coffee        37        3.23        32.65
8       TSF coffee      34.83       3.28        31.42
9       TSF coffee      34.71       3.35        31.45
10      TSF coffee      34.17       3.39        31.15
11      TRF coffee      34.12       3.44        31.14
12      TRF coffee      33.19       3.47        30.75
13      TRF coffee      33.02       3.51        30.85
14      TRF coffee       32.7       3.53        30.77
15      TRF coffee       32.4       3.56        30.62
16      TRF coffee       31.9       3.58        30.51
17      TRF coffee      31.83        3.6        30.57
18      TRF coffee      30.48       3.62        30.58
19      TRF coffee      30.61       3.63        30.36
20      TRF coffee       30.3       3.64        30.3
21       DF coffee      29.67       3.65        30.38
22       DF coffee      29.32       3.66        30.35
23       DF coffee        29        3.67        30.23
24       DF coffee      28.54       3.68        30.35
25       DF coffee      28.26       3.69        30.28
26       DF coffee      28.07        3.7        30.22
27       DF coffee      27.91        3.7        30.11
28       DF coffee      27.78       3.71        30.21
29       DF coffee      27.65       3.72        30.25
30       DF coffee      27.35       3.73        30.21

TSF coffee: tropical semi deciduous forest coffee
agroecosystems, TRF coffee: tropical rain forest
coffee agroecosystems, DF coffee: deciduous forest
coffee agroecosystems. Calculation made with
Estimates Version 8.2.0
(http://viceroy.eeb.uconn.edu/estimates)

TABLE VI
EXCLUSIVE SPECIES FOUND IN THE DIFFERENT
COFFEE AGROECOSYSTEMS, ACCORDINGLY WITH ORIGINAL
VEGETATION TYPE, IN OCOTAL CHICO, SOTEAPAN, VERACRUZ

TSF coffee (23)           TRF coffee (23)

Acosmium panamense        Acalypha microstachya
Alibertia edulis          Apeiba tibourbou
Byrsonima crassifolia     Calophyllum brasiliense
Calathea macrochlamys     Citrus aurantifolia
Carica papaya             Coccoloba uvifera
Chiococca Alba            Eupatorium daleoides
Chrysophyllum cainito     Hyptis mutabilis
Chrysophyllum mexicanum   Leucaena leucocephala
Cojoba arborea            Luehea speciosa
Cupania glabra            Mangifera indica **
Eugenia acapulcensis      Miconia argentea
Eugenia capulli           Montana sp.
Gliricidia sepium         Rollinia mucosa
Heliconia curtispatha     Sida cordiflolia
Malpighia glabra          Sida rhombifolia
Sacoila lanceolata        Sinclaria discolor
Sida acuta                Spathodea campanulata **
Pachira aquatica          Suriana maritima
Tephrosia sp. **          Swietenia macrophylla
Trichilia havanensis      Vanilla planifolia
Vismia camaguey           Vernonia patens
Zanthoxylum caribaeum     Vochysia guatemalensis
Zapoteca sp.              Xiphidium caeruleum

TSF coffee (23)           DF coffee (21)

Acosmium panamense
Alibertia edulis          Adelobotrys adscendens
Byrsonima crassifolia     Agerantia sp.
Calathea macrochlamys     Allophylus cominia
Carica papaya             Annona reticulata
Chiococca Alba            Astrocarium mexicanum
Chrysophyllum cainito     Astronium graveolens
Chrysophyllum mexicanum   Baltimore recta
Cojoba arborea            Bursera simaruba
Cupania glabra            Calyptranthes lindeniana
Eugenia acapulcensis      Catasetum integerrimum
Eugenia capulli           Lacistema aggregatum
Gliricidia sepium         Liquidambar styraciflua
Heliconia curtispatha     Lonchocarpus guatemalensis
Malpighia glabra          Ocotea verticillata
Sacoila lanceolata        Rapanea sp.
Sida acuta                Tectona grandis **
Pachira aquatica          Terminalia amazonia
Tephrosia sp. **          Tetrapterys schiedeana
Trichilia havanensis      Thelypteris blanda
Vismia camaguey           Vismia baccifera
Zanthoxylum caribaeum     Zuelania guidonia
Zapoteca sp.

TSF coffee: tropical semi deciduous forest coffee
agroecosystems, TRF coffee: tropical rain forest
coffee agroecosystems, DF coffee: deciduous forest
coffee agroecosystems. ** Introduced.
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Author:Capitan, Guadalupe Castillo; Avila-Bello, Carlos H.; Lopez-Mata, Lauro; De Leon Gonzalez, Fernando
Publication:Interciencia
Date:Sep 1, 2014
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