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.
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
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.
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.
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.
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: firstname.lastname@example.org
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.
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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|Title Annotation:||texto en ingles|
|Author:||Capitan, Guadalupe Castillo; Avila-Bello, Carlos H.; Lopez-Mata, Lauro; De Leon Gonzalez, Fernando|
|Date:||Sep 1, 2014|
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