Successional vegetation patterns in abandoned

Phytocoenologia, 42 (3 – 4), 101 – 132 Stuttgart, December 31, 2012

Successional vegetation patterns in abandoned pastures of the lower montane cloud forest zone in the Venezuelan Andes by Néstor GUTIÉRREZ BELTRÁN, Mérida, Venezuela, Stefanie GÄRTNER, Freiburg, Germany, Juan C. GAVIRIA R., Mérida, Venezuela, Winfried MEIER and Albert REIF, Freiburg, Germany with 8 figures, 5 photographs and 7 tables Abstract: The recolonization of abandoned pastures by secondary forest in the lower montane cloud forests belt, between 1800 and 2400 m a.s.l, was studied in the Mucujún watershed of the Venezuelan Andes. Using a side-byside approach, 83 relevés were surveyed based on a chronosequence of about 50 years of succession on recently abandoned pastures to advanced secondary forest. A total of 368 vascular plant species were recorded belonging to 205 genera and 82 families. The phytosociological classification based on the estimated species cover discriminated eight successional communities representing a gradient of successional development. The secondary forest communities were characterized by the species group Viburnum tinoides – Palicourea leuconeura which are species that were established during early successional stages and form the main structure of the secondary forest. Time was the best predictor of forest recovery, while land use intensity determined two successional pathways based mainly on floristic differences. The forest recovery is made evident by an increase in biodiversity and structure; however, the secondary forest communities are still in an intermediate successional stage compared to old growth cloud forest in other locations especially when considering the floristic composition. Keywords: Phytosociology, forest classification, lower montane forest recovery, ecological restoration, Andes, Venezuela.

by agricuture and pastures (Sarmiento et al. 1971, Ataroff & Rada 2000). Even though rates of deforestation are decreasing (FAO 2011), primary forest replacement is expected to continue in tropical ecosystems thereby reducing biodiversity and degrading soils. On the other hand the migration of people to cities and socio-economic changes has resulted in the abandonment of degraded low-yielding fields in some regions where secondary forests are now allowed to grow mitigating some of the consequences of deforestation (Aide & Grau 2004, Wright & Muller-Landau 2006). However, the recovery of tropical forest is not always certain. Depending on ecosystem resilience and disturbance intensity an area could either: recover through natural succession to become secondary forest that over time develops structural and functional features compared to the remplaced forest ecosystem (e. g. Kappelle et al. 1994, Aide et al. 1995, Rivera et al. 2000, Chinea 2002); Contrary wise, the area can degenerate into a fire-prone savanna or remain in an arrested successional stage (Sarmiento 1997, Cavelier et al. 1998). The initial biotic and abiotic conditions play an important role in the forests recovery. For example, the availability of seed and other propagules, predation, competition and soil properties generally determine the rate of recovery, the biodiversity, the composition and structure of the secondary forest and its successional development (e. g. Aide & Cavelier

Abbreviations AB ACF ASF DBH GS LMCF LMRF MCF NMDS TSF UMCF UMRF

Basal area Andean cloud forest Advanced secondary forest Diameter at breast height Grassland and Shrubland Lower montane cloud forest Lower montane rain forest Montane cloud forest Non-metric multidimensional scaling Transitional secondary forest Upper montane cloud forest Upper montane rain forest

Introduction Deforestation and land use change are major threats to tropical forest ecosystem biodiversity (Dirzo & Raven 2003). This is particularly evident in Andean montane forests (Myers et al. 2000). The area of the Andean cloud forest has been reduced to less than 10% of its original distribution (Hamilton 2001). In Venezuela large areas of cloud forest are still relatively well preserved in protected areas (e. g. national parks), and in other areas rugged topography hinders anthropogenic disturbances (Ataroff 2001, Cuello & Cleef 2009). However, the cloud forest at lower elevations, on gentle slopes, has already been replaced © 2012 Gebrüder Borntraeger, 70176 Stuttgart, Germany DOI: 10.1127/0340-269X/2012/0042-0528

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tional gradient in the upper montane rain forest and Kelly et al. (1994) provided complete descriptions of the flora and diversity in a montane forest stand of about one hectare at 2600 m a.s.l. These studies show the high diversity and variability of cloud forests depending on the local environmental conditions. The conservation of Andean biodiversity requires not only the preservation of pristine areas but also appropriate land use techniques and the development of restoration strategies to recover deforested areas (Sarmiento 2000). A prerequisite for the restoration of diverse ecosystems, like montane cloud forests, requires knowledge about the flora and ecological processes and the response of the ecosystem to disturbance. This study of plant communities established on abandoned pastures contributes to the classification and understanding of floristic changes and forest recovery in human influenced ecosystems in Andean lower montane cloud forests. The main objective was to determine specific phytosociological associations in relation to successional age and land use patterns thus improving the knowledge about options and limitations of passive forest restoration.

1994, Holl et al. 2000, Guariguata & Ostertag 2001, Myster 2004, Aide et al. 2011). These postdisturbance conditions are determined by land use intensity, field size and length of use (Purata 1986, Aide et al. 1996). Short term disturbances, such as slash-and-burn agriculture have lower impact on the seed bank, the soil and other factors allowing for a relatively fast recovery while long term pastures and areas frequently burned have the lowest recovery rates (Uhl et al. 1988, Aide et al. 1995, Sarmiento 1997, Cavelier et al. 1998). Neotropical secondary forest succession can develop on abandoned anthropogenic ecosystems such as pastures and fallow lands. Beginning with low tree species diversity and homogenous structure, they can grow to become highly complex, species rich forests (Finegan 1996, Myster 2004). This process was summarized by Guariguata and Ostertag (2001 p. 200) into four phases: 1) the initial colonization whereby pioneer grass, herb and fern species establish and begin growing; 2) the early forest development dominated by short-lived woody pioneers; 3) the late forest development which includes some long-lived pioneers and 4) the old-growth forest dominated by shade tolerant trees. However, species composition replacement patterns and recovery times are not predictable. During secondary succession forest structure and ecosystem function seem to recover faster than does floristic composition (Guariguata & Ostertag 2001). Tropical forest successional models are dominated by findings from lowland forest types where most successional studies have been carried out. The few studies examining successional chronosequences in montane tropical ecosystems (e. g. Aide et al. 1995, Kappelle et al. 1996, Howorth & Pendry 2004) show slightly different recovery patterns. Montane tropical forests have slower recovery rates and might have different initial phases characterized by the absence or low frequency of short-lived woody pioneers (Ewel 1980, Aide et al. 1995). In addition to the low number of successional studies done in tropical mountain forests, vegetation classification for the cloud forest of the Andes cordillera of Venezuela has not yet been completed. However, for the Ramal de Garamacal, a sub-region of the Andes cordillera, a phytosociological classification was recently completed (Cuello & Cleef 2009). This classification groups the montane forest of the Ramal de Guaramacal within the order of Meliosma tachirensis – Alchornea grandiflora which includes a total of seven forest communities, four of them are classified as Andean and three as Subandean montane rain forest communities. Related studies in Andean cloud forests have generally focused on ecological and silvicultural aspects. Hetsch & Hoheise (1976) described soil units, and related seven forest types of primary cloud forest in the Venezuelan Andes Cordillera. Also, for the Andes Cordillera there are a few very detailed but small scale studies of the original cloud forest. Schneider (2001) studied an eleva-

Study area The research was conducted in the Mucujún Watershed located north of the city of Mérida (8°.65 N; 71.12 W), which belongs to the Andean cordillera of Venezuela (Fig. 1). Five ecological units were distinguished in the watershed (Ataroff & Sarmiento 2004), they are: the upper and lower Páramo above 3000 m a.s.l., the upper and lower montane cloud forest (MCF) between 1800 and 3000 m a.s.l., and the seasonal montane rainforest below 1800 m a.s.l. Currently, about 30% of the watershed is covered by forest, mainly cloud forest (Gavidia & León 2004). The tropical montane cloud forest, also classified as Andean and Subandean forest (Cuatrecasas 1958), montane rain forest (Grubb 1977) and tropical montane rain forest (Ewel et al. 1976), can be structurally and floristically differentiated into two altitudinal belts, the upper and lower montane cloud forest (LMCF) (Sarmiento et al. 1971, Hamilton 1995, 2001, Kappelle & Brown 2001). The exact altitudinal distribution varies due to precipitation, wind regimes and topography (Ataroff 2001). In the study area the LMCF lies between 1800 and 2400 m a.s.l. (Ataroff & Sarmiento 2003). Recently, major anthropogenic changes and forest recovery have taken place in the LMCF, therefore the study is focused within this altitudinal belt. Climatically the study area is characterized by an annual mean temperature between 12 and 19 °C and mean annual precipitation between 1600 and 1965 mm, with two peaks, one in April and the other in October (Ataroff & Sarmiento 2003, CIDIAT unpublished data). In this area different geological formations converge, however, most of this altitudinal belt consists

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Successional vegetation patterns in abandoned pastures in the Venezuelan Andes

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Fig. 1. Location of the study area in the Mucujún watershed, Mérida state Venezuela.

of Tertiary sediments from the Mucujún and San Javier geological formations. The Mucujún formation is mainly made up of a cyclic sequence of sandstones, siltstones and shales, while the San Javier formation is composed of thick shale, glauconitic and bioturbated sandstones and calcareous fossiliferous marine beds (Ghosh & Odreman 1987). The most common soil types in the study area, according to the classification of the Soil Survey Staff (1975), are humitropepts and troporthents, they are predominantly acidic (pH 4.6 – 5.0), with a loamy to clay loam texture (Urbina 1982). By the beginning of the last century the upper slopes of the watershed, up to 2700 m, were predominantly used as cattle pasture (Gutiérrez in prep.),

while in the valley bottoms and on the lower páramo small scale traditional agriculture was most common. Livestock densities on the land were low. Fallow land and the periodic rotation of pastures with crops was common. The use of fire for weeding pastures and forest clearing remains a common practice in the Andes region (Sarmiento 2002). In the Mucujún watershed, as in other montane regions of the neotropics (e. g. Grau & Aide 2008), land traditionally used for grazing has been abandoned in the last 50 years as people migrated to urban areas mainly due to higher incomes (Gutiérrez in prep.). Since 1985 the watershed was also designated a protected area because of its importance as a source of fresh water for the nearby city of Mérida. In addi-

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tion, since 1989 the upper part of the watershed lies within Sierra de la Culata National Park (Aldana & Bosque 2008). The regulations imposed by the different protection categories do not totally exclude land use, but differentiate between areas with priority for conservation (e. g. forest on steep slopes with inclinations above 30%) and those where traditional land use regimes can be continued (e. g. valley bottoms). Land abandonment allowed for the establishment of secondary vegetation, including shrublands and secondary forests, which nowadays cover an important area of this watershed, especially between 1800 and 2400 m a.s.l. (Gutiérrez 1999, Gavidia & León 2004). Relatively large areas are undergoing succession and different successional stages have developed over several decades. This situation provides the option to study the regeneration potential of the LMCF.

In each relevé all terrestrial growing vascular plants were recorded using a nested sampling design with three different subplot sizes for each height layer (Table 1): for the ground vegetation layer < 1 m in height, subplots of 40 m2 (20 × 2 m) were used, located in the middle of the relevé; for shrublands and understory vegetation between 1 to 3 m in height, 80 m2 (20 × 4 m) subplots were used; and for the tree layers (over 3 m) 200 m2 (20 × 10 m) subplots were used, the tree layers were divided into sub-layers of 5 m each to detect changes in the vertical structure (Table 1). In each subplot and layer the ground cover of individual species was estimated using the modified Braun-Blanquet scale (r, +, 1, 2a, 2 m, 2b, 3, 4, 5) (Glavac 1996). In each plot the following site characteristics were recorded: inclination, aspect, average height of the uppermost vegetation layer and geographical position. Indicators of recent disturbance were measured using an ordinal scale (1 = low: browsing and livestock trails absent; 2 = medium: trails present but not frequently used, browsing present, few or dry cow dung piles; 3 = high: presence of trails, clear evidence of browsing and cow dung, recently felled trees or branches; and 4 = fire: trees and shrubs with fire scars and fire killed trees). Fires are frequently set to prevent tree colonization. Local people were interviewed to cross-check observed indicators with actual land use intensity. Nine relevés were randomly selected within the age classes < 14, 15 – 23 and > 44 years for creating structural profiles. In these relevés all woody plants with a diameter at breast height (DBH = 1.3 m) greater than 5 cm were recorded in 200 m2 (20 × 10 m) plots. For each woody plant, the species, its relative position within the relevé, DBH, crown projection (measured in the four cardinal directions), and height were recorded. In cases where the trees had multiple stems each stem was considered a single individual. Plant samples were identified in the Botanical Garden of the Faculty of Sciences at the Universidad de Los Andes (ULA). These samples were compared with herbarium samples from the Faculties of Sciences (MERC), and at the Forest and Environmental Sciences (MER) of the Universidad de Los Andes in Mérida and the Venezuelan National herbarium (VEN) in Caracas. The samples were left at the MERC herbarium under the numbers NG002-588, for general

Methods Sampling design Based on a sequence of aerial photographs from 1952, 1979, 1987 and 1996 stands of different successional ages were identified. The time of abandonment is not known exactly, therefore it was estimated for every stand using the mean age between two photographs within the series (Aide et al. 1995). Based on this information four structural vegetation formations representing successional stages were identified: 1) grassland about 5 year after abandonment, 2) shrubland, 3) transitional forest (between 14 and 23 years old) and 4) advanced secondary forest (more than 44 years old). A geographical database was developed with GRASS 6.4 geographical information system (GRASS Development Team 2010) for geo-processing the aerial photographs and to delineate the study area. The data base was loaded with the following cartographic information: geology, soil and precipitation (CORPOANDES unpublished data), slope, aspect and altitude. These data were obtained by digitalization and processing of topographic maps (1:25000) (Dirección de cartografía nacional 1975). Based on this cartography information the study area was stratified according to time after abandonment, inclination, and the predominant geology in order to have relatively homogeneous areas needed to assess the successional development using the side-by-side approach where different locations in a gradient after abandonment were assessed to reconstruct the successional pathway (Mueller-Dombois & Ellenberg 1974). Stands with different successional stages were more frequent on slopes with inclinations between 20 to 60 % and in the Mucujún and San Javier geological formation. The stands with these characteristics were loaded on a GPS and located in the field. Between 2006 and 2007, 83 relevés were established in all successional stages trying to record at least 10 relevés for all combinations of successional ages.

Table 1. Height layer used in the estimation of plant cover.

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collection, and relevé numbers and sub-samples for relevé vouchers (e. g. NGP01-nij). The taxonomical classification of the plants followed the nomenclature of the Catalog of Venezuelan Flora (Hokche et al. 2008).

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species and plots. The final differential species in the classification table were those with an F-value > 1.948 (with 10, 67 d.f). The dominant or most frequent species were used to name the species and relevé groups, following phytosociological principles. Gradient analysis

Data analysis

An ordination analysis was performed to relate the successional communities to site variables and time since abandonment. A Non-metric Multidimensional Scaling (NMDS) approach was used in PC-ORD 5.0 (McCune & Grace 2002), using species as variables and the same preprocess as for the tabular classification already described. The Soerensen distance measure was selected (McCune & Grace 2002). For the NMDS axes solution was used considering the lowest stress. The NMDS axes were correlated using the Kendall Tau coefficient, with aspect, inclination, altitude, distance to forest border in 1952, distance to the closest farm houses and time since abandonment.

Classiﬁcation A phytosociological vegetation table according to Braun-Blanquet or the Zürich Montpellier school (Mueller-Dombois & Ellenberg 1974, Glavac 1996) was developed based on a statistical analysis sequence proposed by Wildi (1989) and Wildi & Orlóci (1996) using MULVA 5.0. Before the analysis the Braun-Blanquet coverabundance scale was transformed to an ordinal (0 – 9) scale as proposed by van der Maarel (1979, 2007) and Wildi & Orlóci (1996). An outlier analysis was carried out for the species and relevés, five relevés were consequently excluded because of their floristic separation leaving 78 relevés to be used for the analyses. Species with very low constancy (present in less than 5 % of the relevés) were also excluded (McCune & Grace 2002). When the forest communities reach the intermediate successional stages, the species turnover in secondary forest stands is expected to be relatively slow and major changes are expected in the forest structural features (Finegan 1996, Guariguata & Ostertag 2001). Therefore, numerical analysis was based on a combination of floristic and structural criteria. The occurrence of a species in different layers was used as a separate species/layer unit in the analysis (Table 1). The analyses sequence for developing the phytosociological table proposed by Wildi (1989) is based on two cluster analyses, one analyzes the similarity of the relevés and the other clusters species based on their common occurrence. With a subsequent concentration analysis the cluster groups of species and relevés that were related to one other were arranged along the main gradient detected by a correspondence analysis. Using a discriminant analysis (Jancey’s ranking method Wildi 1989, Wildi & Orlóci 1996)) differential species were selected for the relevé groups. Specifically, the options chosen were: relevé data transformation using the fourth root transformation, followed by a general relativization by species in order to down weight the influence of dominant species (van der Maarel 1979). All species data, other than the zeros were transformed using the logarithmic transformation (Wildi & Orlóci 1996), also applying the general relativization of total species frequency in the relevés. For the cluster analysis, the van der Maarel coefficient, and minimum variance clustering was used in both cases. The first vector of the correspondence analysis explained about 11.9 % of the total variance and was selected as the main gradient used to arrange the groups as well as the individual

Species richness and structure Species richness was assessed for the main successional groups and for the successional communities resulting from the phytosociological classification calculated as the average of the total number of species of the plots in each category. The total species number was further subdivided into the following growth forms: grass-like species (grasses and sedges), forbs, lianas and vines (all climbers), pteridophytes (herbaceous ferns and fern-allies), tree ferns, shrubs, treelets, trees, palms and bamboo. Structural profiles were drawn for selected plots to visualize the physiognomic characteristics of the main community groups resulting from the classification (shrubland, transitional and advanced forest). Also, structure diagrams were drawn showing the average cover of each height layer in Table 1.

Results Plant communities The results of the classification revealed two main relevé groups that represent successional development from abandoned pastures to secondary forest (Fig. 2). The first one includes recently abandoned areas still dominated by grasses and shrubs: the Grassshrubland (GS) group. This group is characterized by the grasses Melinis minutiflora, Axonopus compresssus and the bracken fern Pteridium aquilinum (see Table 2). The second group is the secondary forest dominated by the Viburnum tinoides – Palicourea leuconeura species group (see Table 2). These species establish in early successional stages and become a fundamental part of the forest structure. Within the secondary forest group there are two subgroups: the

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transitional secondary forest (TSF: communities 2.1 and 2.2.1, Fig. 2), which represents the initial forest stages; and the advanced secondary forest (ASF: 2.2.2 Fig. 2). Based on the floristic composition and structural development, eight seral plant communities have been identified. The plant communities were related to the time since abandonment and the previous and present land use representing the recovery process of the lower montane cloud forest. Descriptions of the communities are presented below:

The grass species Axonopus compressus and Melinis minutiflora are differential species of this community group. M. minutiflora was introduced to improve the forage quality of pastures and nowadays is considered an invasive species in the mountain grasslands in the neotropics (Parsons 1972). These two species cover between 5 and 60 % of the surface, depending on the community. The ground layer of this community group is composed of the following grass and sedge species Paspalum notatum, Panicum trichidiachne; Scleria hirtella, Rhynchospora tuerckheimii; the forbs Trimezia martinicensis, Hyptis spp., Aeschynomene elegans; the shrubs Tibouchina geitneriana, Clidemia ciliata, Chromolaena laevigata, Chromolaena voglii, Baccharis trinervia; and the fern Anemia villosa. Frequently there are remnant trees of the previous forest or natural tree regeneration that was left in the pasture as a source of firewood or shade for cattle. These individuals belong to the following species: Calycolpus moritzianus, Myrsine coriacea, Vismia baccifera (Fig. 3). Another important component of this community group is the bracken fern Pteridium aquilinum that grows intermingled with the grass and shrub species and also persists in more advanced successional stages.

Group 1: Axonopus compressus – Melinis minutiflora Grass- and Shrubland (Table 2/1 – 27)

This community group includes herbaceous or grasslike vegetation with scattered shrubs and trees. The grass layer is 0.20 to 0.50 m tall, but dense patches of thick grasses mixed with small shrubs can reach heights of up to 1.2 m. The herbaceous layer covers, on average, more than 60 % of the surface. Under the taller grasses and shrubs an inconspicuous moss layer can occur covering less than 5 %. In some relevés there is up to 20 % bare soil as a result of cattle trampling. Litter covers up to 40 % of the the ground, especially when the herbaceous layer is tall (> 50 cm).

Fig. 2. Plant communities of the secondary succession of the LMCF on abandoned pastures. Result of a cluster analyses of 78 relevés, based on Soerensen distance and flexible beta (ß = – 0.25), species cover was transformed to an ordinal scale. The numbers represent the groups and communities of the phytosociological classification (Table 2).

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The Axonopus compressus – Melinis minutiflora group is divided into four communities: the Dichanthelium aciculare grassland (subgroup 1.1 Fig. 2), the anthropogenic Ischaemum latifolium savanna (subgroup 1.2 Fig. 2), the Rhynchospora rugosa pasture (subgroup 1.3 Fig. 2) and the Baccharis nitida shrubland (subgroup 1.4 Fig. 2).

Melinis minutiflora community group such as Pteridium aquilinum, Paspalum notatum and Scleria hirtella. Ecology and distribution

The Dichanthelium aciculare community can be found relatively close to farmsteads and trails. The relevés represent recently abandoned pastures or fallow fields. However, when land owners want to increase the lands production by weeding or burning, this vegetation community can become an active pasture or a crop field again. Cattle can be found foraging on those lands. The Dichanthelium aciculare community was found on mid to upper slopes, generally in places having an inclination of more than 20 %. Two subtypes can be distinguished within this community: the Oxalis latifolia and the Psidium guineense subtype.

Dichanthelium aciculare Grassland (Table 2/1 – 10) Structure and composition

The Dichanthelium aciculare community is a grassland type with a dense, patchy ground layer intermingled with shrubs (Photo 1). The ground layer generally covers more than 70 % of the surface and is about 1 m tall. Scattered small trees commonly cover 10 to 20 % and reach heights of 5 to 7 m, the most frequent species are Calycolpus moritzianus, Myrsine coriacea, Vismia baccifera, Miconia theaezans and Fraxinus uhdei. Diagnostic species are Dichanthelium aciculare, Kyllinga pumila, Chamaecrista nictitans, Chaptalia nutans, Lantana maxima. They are accompanied by the dominant species of the Axonopus compressus –

Oxalis latifolia Subtype: pastures and old crop ﬁelds (Table 2/1 – 2) Structure and composition

The ground layer of the Oxalis latifolia–subtype is between 0.6 and 1.5 m tall with a homogeneous coverage of 70 to 90 %. The shrub layer is about 1.5 m

Photo 1. Dichanthelium aciculare (relevé 11) abandoned grassland. In the background Miconia theaezans is densely flowered.

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dus, Clidemia ciliata var. elata, Baccharis trinervia. Also frequent are small trees and juveniles of Vismia baccifera, Calycolpus moritzianus, Myrsine coriacea, Fraxinus uhdei, Palicourea leuconeura and Miconia theaezans, covering up to 10 % of the area and reaching 3 to 6 m in height.

tall and has coverage of 10 to 30 %. A 4 to 7 m high tree layer mostly composed of Calycolpus moritzianus commonly covers 20 to 25 % (Fig. 8a). This community subtype is floristically richer than the other grassland types. The abundance and dominance of the grass-like species is lower compared with the other grasslands, and forbs and small shrubs are dominant, Oxalis latifolia, Sida rhombifolia, Clibadium surinamense and Castilleja arvensis are diagnostic species of this subtype. Companion species are the grass-likes and the forbs Paspalum lentiginosum, Carex longii; Desmodium molliculum, Elephantopus mollis, Borreria laevis; the shrub species Piper aduncum, Monochaetum meridense, Monnina pubescens, Rubus floribundus, and the ferns Blechnum occidentale and Thelypteris balbisii.

Ecology and distribution

The Psidium guineense subtype represents old pastures that have not been maintained nor cleared for several years. From the aerial photos it’s been at least five years. Although they are occasionally grazed by cattle (personal observation) the isolated dense patches of shrubs and grasses make access for cattle difficult. These patches of shrubs serve as important “nurse” patches facilitating the establishment of shrubs and trees such as Calycolpus moritzianus, Myrsine coriacea and Vismia baccifera, Palicourea leuconeura. Additionally, the abundance of grasses decreases under them. The Psidium guineense subtype has many juvenile trees with the potential to become forest in the medium term. However, because these pastures have relatively easy access and are close to farmsteads it is common that become active pastures again by being burned.


The Oxalis latifolia subtype is found on steep slopes (30 – 33°), with a north northwest exposition, moist soils with a loamy to sandy clay loam texture. The stands are close to farmsteads and farm roads. Due to their steepness the sites cannot be permanently used as pasture or cropland and they often occur as early successional grasslands or fallow. According to information provided by the local farmers, the stands of this subtype were used as crop fields as recently as 5 – 10 years ago. This is also indicated by the occurrence of common cropland weed species like Oxalis latifolia (e. g. Royo-Esnal & López 2008). Although the sites covered by the Oxalis latifolia subtype are no longer intensively managed, they must be regarded as a successional stage, periodically arrested, by disturbances caused by cropping or by cattle grazing.

Ischaemum latifolium Savannas (Table 2/11 – 15) Structure and composition

The Ischaemum latifolium community is a grassland or herbaceous community with a very dense ground layer up to 1.5 m in height (Fig. 8c). Ischaemum latifolium together with Pteridium aquilinum are the dominant species. Companion species are the grasses Setaria geniculata, and Isachne rigens, and sedges Rhynchospora rugosa, and R. tuerckheimii. The shrub layer is composed of emergent shrubs up to 3 m tall (Fig. 8c), mainly dominated by Clidemia ciliata, Palicourea leuconeura and Macleania rupestris. The tree layer is made up of sparse treelets of Vismia baccifera, Myrsine coriacea, Piptocoma niceforoi, Myrcia fallax and Fraxinus uhdei. Common are the climbers Blepharodon grandiflorum and Oligactis volubilis.

Psidium guineense Subtype: abandoned pastures (Table 2/3 – 10) Structure and composition

The Psidium guineense subtype is characterized by a homogeneous meadow-like grass layer shorter than 0.5 m, dominated by the grass species Axonopus compressus, Dichanthelium aciculare, Paspalum notatum and the sedges Rhynchospora aff. rugosa, R. nervosa, Kyllinga pumila, Scleria hirtella, among others. Less abundant are the forbs Chaptalia nutans, Chamaecrista nictitans, Trimezia martinicensis, Elephantopus mollis, Stylosanthes mexicana. There are, however, other species in this layer that are usually associated with shrubby patches reaching 0.6 to 1.2 m in height (Fig. 8b). These are Pteridium aquilinum and Melinis minutiflora, accompanied by Andropogon bicornis, and Paspalum lentiginosum. The shrub layer is generally lower than 2 m, covering 10 – 50 (– 70) % of the surface. Psidium guineense occurs with high constancy together with shrubs like Chromolaena laevigata, C. voglii, Rubus floribun-


The Ischaemum latifolium community seems to be in a stage of arrested succession. As can be seen in the aerial photographs (1952 – 1996), over 40 years there have been few structural changes in the stands currently dominating this community. Undoubtedly the recurrent use of fire by the local farmers prevents further successional development. Sites with this vegetation type and close to farms have a long history of being burned, probably by the locals. Snags and shrubs with fire scars are evidence of past fires. On

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the other hand cattle grazing seems to be scarce. This community is mainly found on clay, loamy clay or sandy clay loam soils at mid slopes with inclinations between 20 – 40 % and mainly southwest expositions.


This pasture community is more frequently visited by cattle than the others and their selective grazing maintains the structure of this vegetation type. The aerial photographs show few changes in areas with these characteristics even though farmers indicated that they do not “clear” (weed) the fields. The fields also seem to be in an arrested successional stage as a result of regular grazing. These pastures are found at mid slopes with an inclination between 27 to 36 (– 65) %, and south to northwest aspects, on sandy clay to sandy clay loam soils. They are frequently located at considerable distance from farms.

Rhynchospora rugosa Pasture (Table 2/16 – 22) Structure and composition

The Rhynchospora rugosa community mainly differs from the other grassland communities mentioned by its structure: the ground layer is shorter (0.2 to 0.4 m height), and Pteridium aquilinum and Melinis minutiflora are rarely found growing over 1 m. The shrub layer is also shorter and has less coverage (5 – 45 %, Fig. 8d). There are some scattered trees reaching 4 m in height with less than 5% coverage (Photo 2). The species composition is similar to the communities described above although there is a higher constancy and abundance of Rhynchospora rugosa, Paspalum notatum and of the shrubs Clidemia ciliata var. ciliata, Monochaetum bonplandii and Macleania rupestris. Common tree species are Vismia baccifera, Piptocoma niceforoi, and Myrcia fallax. The liana Blepharodon grandiflorum shows high constancy.

Baccharis nitida Shrubland (Table 2/23 – 27) Structure and composition

The herbaceous layer of the Baccharis nitida community is irregular and patchy but still forms a main component of this vegetation type, it covers 15 to 80 % of the surface and is about 0.5 m in height (Fig. 8e); Pteridium aquilinum, Melinis minutiflora and other grasses have high constancy, but their dominance is not as great as in the communities already described. In the ground layer Rubus floribundus, Coccocypse-

Photo 2. Rhynchospora rugosa grassland (relevé 16). In this case the grass layer is shorter (compare with the others grasslands communities).

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about 4 m in height covering 20 to 40 % (Fig. 8e). The dominant species are C. moritzianus, M. coriacea, B. nitida, Miconia theaezans, V. baccifera, V. tinoides and Heliocarpus americanus (Fig. 3). There are some cases where C. moritzianus, M. coriacea and Fraxinus uhdei occur as emergents. The abundance of lianas increases with increases in structure; common are Clematis populifolia, Ditassa aff. albonerva, and Blepharodon grandiflorum. On the ground there is a moss layer covering about 5 % of the surface, the reduction of the herbaceous cover layer is accompanied by an increase in litter accumulation.

lum lanceolatum, Arenaria lanuginosa, Elephantopus mollis are frequent as well as the ferns Blechnum occidentale, Nephrolepis pendula and Anemia villosa. The shrub layer is dense with a cover between 15 – 65 % and reaches up to 2.5 m in height (Fig. 8e); the diagnostic species Baccharis nitida has a high constancy and coverage in this layer. Other common shrubs are Lepidaploa canescens, Clidemia ciliata var. elata, Macleania rupestris, Palicourea leuconeura and Miconia aeruginosa. Relevant in the shrub layer is the cover of the tree species Calycolpus moritzianus, Vismia baccifera, Viburnum tinoides and Myrsine coriacea. The tree layer is made up of an incipient canopy

Fig. 3. Structural profile of a typical grass-shrubland community (relevé 86: community of Baccharis nitida).

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Group 2: Viburnum tinoides – Palicourea leuconeura Secondary forest (Table 2/28 – 78)

This community represents the shrubland stage on abandoned grasslands. Floristically it is an intermediate stage between grasslands and secondary forest. There are species from the grassland communities described above and species from more advanced successional stages. The lower frequency of disturbances (fire, weeding, cattle grazing) allows the establishment of shrubs and trees promoting higher species richness and the development of a more complex vertical structure (Fig. 7), compared to the stages before (1.1.2, 1.2 and 1.3, Fig. 2) and after (2.1 and 2.2, Fig. 2). However, these areas are not completely abandoned as evidenced by the presence of cattle trails, browsing and firewood collection. According to the aerial photographs the stands with these characteristics have been undergoing succession for about 14 to 20 years. This community is commonly found on east to south facing slopes with inclinations between 20 to 55 %. The soil texture is generally loamy to sandy clay. The stands are located relatively close to farms and within a matrix of grassland-shrubland communities.

This forest group is composed of woody pioneer species occurring in the grassland communities as seedlings or sprouts in the ground and a shrub layer. These pioneer species become the dominant species of the canopy and understory in the subsequent secondary forest communities. The species within this group remain in the advanced secondary forest until they are replaced by more shade tolerant cloud forest species. The main species of this community group are the trees and treelets Viburnum tinoides, Miconia theaezans, Palicourea leuconeura, Myrsine coriacea, Calycolpus moritzianus and Alchornea triplinervia (in more advanced stages); the ferns Blechnum occidentale and Nephrolepis pendula, the orchid Malaxis aff. excavata; and the lianas Oligactis volubilis, Baccharis decussata and Calea septuplinervia. Individuals of the woody species more or less form a closed canopy reducing the light availability for grasses and other herbaceous pioneers facilitating the establishment of more shade-tolerant species.

Fig. 4. Structural profile of Escallonia paniculata – Clethra fagifolia transitional secondary forest (relevé 13).

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nant canopy species are V. tinoides, M. theaezans, Myrsine coriacea, Vismia baccifera, P. leuconeura and Calycolpus moritzianus. Some shrub species such as M. rupestris and sometimes the fern P. aquilinum can reach the canopy usually by leaning on neighbouring trees. Above the canopy are frequent emergents or remnant individuals of the tree species C. moritzianus, M. coriacea, F. uhdei and V. baccifera (Fig. 4). In all layers, the lianas Oligactis volubilis, Passiflora edulis var. edulis and Calea septuplinervia can be found.

Transitional secondary forest (Table 2/28 – 48)

There are two communities classified as transitional secondary forest. They represent forests in an intermediate stage between shrub-grassland and advanced secondary forest as they have floristic elements of both. They belong mainly to the Viburnum tinoides – Palicourea leuconeura group although they retain species like Pteridium aquilinum, Macleania rupestris, Chromolaena voglii, and small individuals of the pioneer tree species Vismia baccifera. All of these species have a higher constancy in shrub-grassland communities. These forests do not have a distinctive floristic composition as they experience continuous species turnover as a result of succession. However, these communities differ from the advanced secondary forest because of the absence of several species and a less well developed structural diversity. The transitional forests are composed of three main vertical layers: the ground layer which covers about 25 % of the surface reaches 0.70 m in height; the understory is 1 to 2.5 m in height and covers about 30 %, and the tree layer which reaches a height of 3 to 6 m and is the main canopy and has a uniform cover (50 to 70 %); frequently above this layer there are emergent trees 8 to 12 m tall covering 5 to 30 % (Fig. 4, Fig. 8f,g). Terrestrial mosses cover between 1 to 20 % of the ground. A thick layer of litter covers about 70 % of the ground surface.


This is a transitional forest community and according to the aerial photos the stands were abandoned about 20 to 40 years ago. The structural development is evident at this stage; nevertheless, there is still a relatively dense layer of heliophytes like Pteridium aquilinum and other grassland species remaining from earlier successional stages due to the low height and openness of the canopy. The establishment of additional woody species together with the remnant herbaceous species resulted in the relatively high species richness found in this group (Fig. 7). This vegetation type is generally found on NE to SW slopes with an inclination of 30 to 60 %. These sites have mostly clay loam to loamy sand soils. They are located in places relatively close to farms, but have limited access. Cattle use occasionally this forest to travel through and as cover.

Peperomia galioides Transitional secondary forest (Table 2/28 – 36)

Escallonia paniculata – Clethra fagifolia Transitional secondary forest (Table 2/37 – 48)

Structure and composition Structure and composition

The Peperomia galioides community is a relatively open forest. The ground layer covers from 10 to 40 %, and averages 0.5 m in height. The diagnostic species are Peperomia galioides, accompanied by Coccocypselum lanceolatum, Borreria laevis, Panicum aff. trichidiachne, Rubus aff. floribundus; the ferns Blechnum occidentale and Nephrolepis pendula and the orchid Ponthieva maculata, among others, grow in the ground layer. There are abundant seedlings of Fraxinus uhdei, Myrcia fallax, Rhamnus sphaerosperma and less abundant are Toxicodendron striatum and Alchornea triplinervia. The understory has a cover of 20 to 60 % and reaches a height of 2.5 m (Fig. 8f). It is composed mainly of treelets species Palicourea leuconeura, Viburnum tinoides, Miconia theaezans, Clusia aff. salvinii, R. sphaerosperma, M. fallax, and some shrubby pasture species like Macleania rupestris, Clidemia ciliata, Monochaetum bonplandii and the bracken fern Pteridium aquilinum. The canopy reaches 4 to 6 m in height and covers 30 to 50 (– 70) % (Fig. 8f). The individual tree species present in this layer generally have multiple, relatively thin stems (e. g. V. tinoides, M. theaezans) or are densely branched (e. g. Myrcia fallax) (Photo 3). The domi-

The Escallonia paniculata – Clethra fagifolia community is relatively similar to the Peperomia galioides community in terms of structure (Photo 4), however it differs slightly in the species composition. The ground layer is about 0.5 – 0.7 m in height and covers about 20 % (Fig. 8g), it is dominated by Coccocypselum lanceolatum, Monochaetum bonplandii, Psychotria aubletiana, Palicourea leuconeura, Baccharis brachylaenoides; the grasses Ichnanthus rigens and Axonopus compressus, and the sedge Rhynchospora tuerckheimii; the pteridophytes Blechnum occidentale, Nephrolepis pendula, Lycopodium complanatum, the orchids Ponthieva maculata, Malaxis excavata and Cyclopogon sp.; the lianas Oligactis volubilis, Calea septuplinervia and Blepharodon grandiflorum; and juveniles of the tree species forming the canopy. The understory layer (1.5 – 3 m in height, 25 – 50 % cover, Fig. 8g) is composed of treelets and shrubs such as P. leuconeura, Viburnum tinoides, Macleania rupestris, Myrcia fallax, Vismia baccifera, the fern Pteridium aquilinum, the lianas Oligactis volubilis, Calea septuplinervia and Blepharodon grandiflorum and in some places the tree fern Cyathea caracasana

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Photo 3. Peperomia galioides transitional secondary forest (relevé 21); numerous thin stems are still the main structure of this early successional forest.

Photo 4. Escallonia paniculata – Clethra fagifolia Transitional secondary forest (relevé 12).

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niceforoi, which could reach the emergent tree layer and become part of the main forest canopy in later successional stages. The grassland and savanna species of the ground layer are less dominant, a decline in the abundance of grass-like species and also Pteridium aquilinum, creates the conditions required for the establishment of shade-tolerant species. This community is commonly found at a mid slope position with a S to SW exposition and an inclination of over 30 %, on loam to sandy clay loam soils. Generally this secondary forest is found far away from farms or is difficult to access. Currently these areas are not utilized; however there was evidence of very extensive cattle use.

was encountered. The main canopy layer is 4 to 7 m in height and covers 40 – 70 %; the dominant species are V. tinoides, M. fallax, Piptocoma niceforoi, V. baccifera, Myrsine coriacea, Miconia theaezans, Miconia lonchophylla, and Clusia aff. salvinii, accompanied by the shrub M. rupestris, and the lianas O. volubilis, Securidaca sp. and Smilax cf. subpubescens. The diagnostic species Escallonia paniculata, and Clethra fagifolia are more abundant in this layer. There is a layer of emergent trees about 10 to 13 m tall covering 5 to 20 % (Fig. 8g), this layer is dominated by P. niceforoi, some individuals of Fraxinus uhdei, Alchornea triplinervia, V. baccifera, and the lianas Oligactis volubilis and Securidaca sp. Ecology and distribution

Advanced secondary forest (Table 2/49 – 78)

This forest type shows clear evidence of advanced development towards secondary forest in terms of structure as well as in species composition. The canopy is composed of tree species such as Piptocoma

These are well structured forest communities, generally with four vertical layers. The top layer is a thick canopy (> 50 % cover) reaching from 8 to 12 m in

Fig. 5. Structural profile of Psychotria meridensis – Hedyosmum sp. advanced secondary forest (relevé 26).

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height, dominated by Piptocoma niceforoi (Fig. 5) accompanied by other common trees such as Alchornea triplinervia, Hieronyma moritziana, Calycolpus moritzianus (Table 2). Under this main canopy there is a treelet layer (height 5 – 8 m), where the Viburnum tinoides – Palicourea leuconeura species group remain present. The third layer is the understory (averages 2.5 m in height with 25 % cover). The most relevant species are Psychotria aubletiana, P. leuconeura, Miconia theaezans, V. tinoides, Geissanthus fragrans and Eugenia sp. The ground layer reaches about 0.60 m in height and has 20 % coverage; P. aubletiana, P. leuconeura and juveniles of Geissanthus fragrans are some of the most common species. Although these forest communities are within the Viburnum tinoides – Palicourea leuconeura community group, the diagnostic species are slightly less abundant. But at the same time, species such as G. fragrans, A. triplinervia, P. aubletiana, Eugenia sp. and H. moritziana become more common.


The dominant species are mainly from the V. tinoides – P. leuconeura species group. These species, established during the grass-shrubland stages, thereafter grow into the higher vertical layers and reach their peak importance (dominance and constancy) in about 15 to 30 years after the stands were abandoned. The diagnostic species of this community group are slowly being replaced by more shade-tolerant species such as Turpinia occidentalis, Geissanthus fragrans, among others. This community is also characterized by higher species richness (Fig. 7), due to the convergence of several species from the young secondary forest (transitional forest) and the advanced secondary forest. This community is found on steep slopes with over 30 % inclination usually facing W to NW, the soil texture can range from sandy clay to loamy sand. These stands are not completely abandoned as signs of cattle trampling were noticed in some relevés during fieldwork. Psychotria meridensis – Hedyosmum sp. Advanced secondary forest (Table 2/60 – 78)

Turpinia occidentalis Advanced secondary forest (Table 2/49 – 59) Structure and composition

Structure and composition

The canopy of this forest community is uniformly about 8 m in height and covers, on average, 60 % of the surface (Fig. 8h). In this layer are the remnant and emergent trees belonging to the species Calycolpus moritzianus, Fraxinus uhdei, and Myrsine coriacea intermingled with Miconia theaezans, Viburnum tinoides, Cordia cylindrostachya, Geissanthus fragrans, Montanoa quadrangularis, and the climbers Oligactis volubilis, Clematis populifolia, Stigmaphyllon bogotense and Baccharis decussata. Immediately under the canopy there is a subcanopy layer (4 – 7 m in height, 25 – 50 % cover, Fig. 8h) dominated by V. tinoides, M. theaezans, Palicourea leuconeura and C. moritzianus. The understory covers about 30 %, reaching 3 m in height (Fig. 8h); again the dominant species are M. theaezans, V. tinoides, P. leuconeura, accompanied by individuals of the differential species Turpinia occidentalis, Cestrum aff. tomentosum, Inga oerstediana, Tetrorchidium rubrivenium and Clusia multiflora. The ground layer retains some floristic elements of the grass-shrubland communities such as Pteridium aquilinum, Borreria laevis, Axonopus compressus, Rubus floribundus, Coccocypselum lanceolatum, but also more shade-tolerant species with higher constancy, including Rhamnus sphaerosperma, Psychotria aubletiana, Geissanthus fragrans; the sedge Uncinia hamata, the grasses Olyra latifolia, Ichnanthus tenuis, the herbaceous vine Valeriana bractescens and the fern Thelypteris balbisii. The palm Chamaedorea pinnatifrons can be found with low constancy.

The main canopy is between 8 and 10 m in height and covers 50 to 70 % (Fig. 8i,j), with some 14 m tall emergent Piptocoma niceforoi, Alchornea triplivervia, Hedyosmum sp., Tetrorchidium rubrivenium, Myrsine coriacea, Miconia theaezans, Hieronyma moritziana trees, and the lianas Manettia moritziana, Mikania banisteriae and Calea septuplinervia. In the subcanopy species of the Viburnum tinoides – Palicourea leuconeura group remain dominant together with individuals of Piper longispicum, Hedyosmum sp., the palms Wettinia praemorsa and Chamaedorea pinnatifrons, the lianas Mikania banisteriae, Smilax subpubescens, Manettia moritziana and less frequently the tree fern Cyathea caracasana (Photo 5). The understory is 1.5 to 3 m in height and covers about 20 % (Fig. 8i,j). The dominant species are Eugenia sp., Geissanthus fragrans, Hedyosmum sp., Piper longispicum, Prunus moritziana, Hieronyma moritziana, Miconia spinulosa, Psychotria meridensis and Psychotria amita. The ground vegetation is dominated by Psychotria aubletiana, P. meridensis, Piper prunifolium, the orchid Malaxis excavata and Cyclopogon sp., the Cyperaceae Rhynchospora tuerckheimii, the ferns Blechnum occidentale, Nephrolepis pendula, Serpocaulon fraxinifolium, the palm W. praemorsa, and juveniles of Myrcia fallax, H. moritziana, Hedyosmum sp., Myrsine coriacea. Ecology and distribution

These forests already have structural similarities to cloud forest with larger A. triplinervia and H. mori-

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Photo 5. Relevé 40 Psychotria meridensis – Hedyosmum sp. Advanced secondary forest.

together with the lianas Amphilophium pannosum and Securidaca sp. Additional characteristics include the presence of individual Hieronyma moritziana and Myrcia fallax trees appearing in the upper layers whereas the treelet Solanum sessile and juveniles of the tree species Billia rosea are common in the understory.

tziana trees and also a higher constancy of palms (W. praemorsa and C. pinnatifrons) and the bamboo Chusquea maculata. Although, there are not as many species as in the Turpinia occidentalis forest, there is an increase in the number of species compared with earlier successional stages (Fig. 7). Within the community, two subtypes can be differentiated mainly by the presence or absence of the bamboo species Chusquea maculata.


Chusquea maculata Subtype (Table 2/60 – 74)

This subtype is commonly found on S to SW exposed slopes with inclinations between 20 and 40 %, generally growing on loamy to sandy clay loam soils. This type is more abundant at higher elevations, i. e., around 2200 m a.s.l. According to the aerial photos, this forest type has developed after more than 40 years of succession. Some bamboo species of the genus


This subtype is characterized by high abundance of the bamboo Chusquea maculata in all vertical layers

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Fig. 6. NMDS scatter plot of the species composition of 78 relevés in a gradient of successional development after the abandonment of pastures in the LMCF. The dark gray (continue line) vectors represent the environmental variables: Age: time after pasture abandonment; distofo: the distance from relevés to the forest border in 1952; plotofar: distance from relevés to former farmhouses. Black (dash line) vectors represent the cover (Canopy) and height (Height) of the tree layer. The final stress for the 2 axes solution: 11.85, variance represent by the axes: axis1 = 73, 3%, axis2 = 15.7%.

Soil texture is generally loamy to sandy clay loam. This is the oldest forest community found within the successional sequence. The dominant species still belong to the Viburnum tinoides – Palicourea leuconeura species group, but species like P. demissa, Ocotea sp.1, Miconia meridensis, M. bernardii, Psychotria meridensis and P. amita indicate a successional trend towards the recovery of the cloud forest flora.

Chusquea are common elements of secondary montane forests (Kappelle et al. 1994, Bussmann 2001, Martínez 2007), the presence of this species is related to some degree to disturbance; however, C. maculata is not a pioneer of open stands. The high abundance of C. maculata in the upper layers is due to its ability to use trees as support to climb and dominate the subcanopy and even the uppermost forest canopy. Palicourea demissa Subtype (Table 2/74 – 78)

Secondary successional development The NMDS ordination displays the successional pathway of the abandoned fields in the LMCF of the Mucujún watershed (Fig. 6). It supports the classification described above. The best solution was reached with 2 dimensions representing about the 89% of the variance. The ordination was rotated to show the first axis as the main gradient, in this case it represents about 73 % of the variance. This axis shows the floristic and structural development (height and cover) of the forest in a time gradient after land abandonment. The main gradient along the first axis extends from the left side of ordination diagram beginning with


This subtype is characterized by Palicourea demissa which is a treelet common in the understory and subcanopy. Further diagnostic species are the shrub Psychotria blepharophora, and juveniles of a Lauraceae tree of the genus Ocotea, as well as the climber Mikania banisteriae in the canopy. Ecology and distribution

These forests can be found on SE to W exposed slopes with around 40 % inclination and over 2200 m a.s.l.

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the younger successional stages, on the left side, followed by shrubland and transitional secondary forest at the plot center and the older secondary forest on the right side. This axis also shows a relationship between the successional stage and the spatial distance to old growth forest (distofo, Fig. 6). The ASF were generally located closer to the forest border in 1952 than the earlier successional communities. The second axis represents the land use intensity. This is expressed by the distribution of the secondary forest communities along a gradient of distance from former farms. In the lower part of the diagram are the TSF of Peperomia galioides and the ASF of Turpinia occidentalis. These are relevés from stands situated closer to former farmsteads which are still influenced by higher land use intensity.

123

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Solanum (7), Piper (7), Desmodium (6), Paspalum (5), Psychotria (5) and Rhynchospora (5). Along the successional gradient (axis 1 Fig. 6), the total number of species increases. In the GS group 196 species were recorded, 146 genera in 53 families, and in the ASF group 246 species belonging to 75 families and 177 genera (Table 4). When comparing the communities and subtypes there is a general pattern of increasing species richness with advancing successional development (Fig. 7). The Asteraceae remained as the most species rich family in all successional communities. However, the composition of other families and genera varied with the successional development (Table 4). The families Poaceae and Cyperaceae were codominant in the earliest stages of succession and were then progressively replaced by woody families such as Melastomataceae and Rubiaceae; however, Poaceae remained the fourth most diverse family in the advanced secondary forest. At the genus level, Miconia, Paspalum and Baccharis were the most species rich genera in the GS commu-

Diversity A total of 368 morphospecies of vascular plants were recorded in the 78 relevés of the seral communities of the LMCF in the Mucujún watershed. This includes 219 species of dicotyledons belonging to 144 genera and 57 families, 69 species of monocotyledons in 46 genera and 13 families, 19 species in 13 genera and 11 families of pteridophytes and one gymnosperm species of the genus Pinus. Out of the total 368 morphospecies, 259 were identified to the species level, 50 to genus, 29 to family and 31 remained indeterminate, fourteen of them pteridophytes. The most species-rich plant families were Asteraceae, Poaceae, Melastomataceae, Rubiaceae, Cyperaceae and Fabaceae (Table 3); these families include mainly pioneer and long-lived pioneer species. Only seven families were represented by more than 10 species. The most species rich genera were Miconia (15),

Table 4. Most diverse plant families and genera of the main group of communities in the successional pathway of the LMCF.

1. Grassland- shrubland Families: 53 Genera: 146 Species: 196

Families Asteraceae Poaceae Cyperaceae Melastomataceae Fabaceae Piperaceae

N Sp 34 29 12 12 10 4

Solanaceae 2. Transitional Forest Families: 66 Genera: 142 Species: 178

Genera Miconia Paspalum Baccharis Chromolaena Rhynchospora Desmodium

N Sp 5 5 4 4 4 4

Miconia Baccharis Piper Palicourea Psychotria Rhynchospora

6 4 3 3 3 3

1

Asteraceae Melastomataceae Poaceae Rubiaceae Piperaceae Solanaceae

21 15 13 10 5 3

3. Advanced secondary forest Families: 75 Genera: 177 Species: 246

Asteraceae 28 Miconia 9 Rubiaceae 19 Piper 7 Melastomataceae 17 Psychotria 5 Poaceae 15 Solanum 5 Piperaceae 9 Baccharis 4 Solanacea 9 Palicourea 4 Note that genera do not correspond to the families in the left column. The values (N sp)refer to number of species

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nities; Miconia, Baccharis and Piper in the TSF; and in the ASF are Miconia, Piper and Psychotria (Table 4).

(sd < 3.3), and the maximum height reaches about 14 meters. The BA increases to more than 20 m2.ha-1 in about 40 years developing from a grassland to a forest; this pattern is also observed with the other structural variables. However, the recovery rate appears to be fast during the first 20 years and then decreases. The final BA amassed by the ASF, where about 63 % of the stems were over 10 cm, is 22.66 m2.ha-1.

Growth form spectrum We found a clear tendency towards diversification of growth forms with greater successional development. The number of grass-like (sedges and grasses) and forb species decreased drastically while trees, treelets, shrubs and climbers (lianas-vines) became the most abundant species of the advanced sere (Fig. 7). The GS communities showed the highest differentiation. In the community 1.1 (1.1.1 and 1.1.2) the forb and grass-like species represented over 60 % of the growth form spectra, while in the communities 1.2 and 1.3 shrub and grass-like species were the most abundant (Fig. 7). Outstanding were the number of lianas and shrubs in the advanced forest, both represented more than 40 % of the species, there were almost as many species of lianas as tree species. Only one bamboo species (Chusquea maculata) and two palms (Wettinia praemorsa and Chamaedorea pinnatifrons) were found, both of them in the ASF.

Discussion Successional communities composition and structure The type and intensity of land use practices determined the composition and structure of plant communities and also their persistence in the landscape (Purata 1986, Sarmiento 1997, Myster 2004). The common anthropogenic disturbances in Andean mountains are extensive grazing, small scale cultivation and use of fire for clearing weeds (Hueck 1961, Sarmiento et al. 1971, Sarmiento 2002). These disturbances result in patchy landscapes and forest stands with different successional communities many of them stable, but when land use ceases, succession advances. The progressive abandonment of pastures in the Mucujún watershed has allowed the establishment of plant communities that describe a successional development gradient. In this chronosequence, there is a clear trend towards the recovery of biodiversity and structure going from grasslands to advanced secondary forest. The total number of families, genera and species increases with succession. Also increasing is the diversity of growth forms, the cover of shrubs and trees and with it different strata are formed.

Structure The structural development of the successional communities after land use abandonment was more pronounced than was the floristic development. The earlier successional stages are mainly composed of a dense ground vegetation layer that, with time, develop into a well structured forest with a clear canopy of two to three or more vertical layers (Fig. 8). The increasing number of woody individuals, basal area (BA), mean DBH and height indicate the recovery of forest structure (Table 6). The DBH ranges between 5 and 25 cm, the average is around 9.0 cm

Table 5. Distribution of growth forms in the main successional groups of the LMCF.

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Fig. 7. Growth form spectra and species richness of the successional pathway of LMCF. The numbers on the X-axis are the successional communities (GS: 1.1-1.4, TSF: 2.1 and ASF 2.2). The secondary Y-axis represents the mean number of species.

tziana, Clusia multiflora, Alchornea triplinervia, Inga oesterdiana, Guettarda crispiflora, Weinmannia spp. (Hetsch & Hoheisel 1976, Cleef et al. 1984, Kelly et al. 1994, Meier 1998, Bussmann 2001, Schneider 2001, Howorth & Pendry 2004, Ramos & Plonczak 2007, Cortés 2008, Rangel-Ch et al. 2009, Schwarzkopf et al. 2011). The species of the Viburnum tinoides – Palicourea leuconeura group are also reported for a wide range of ecosystems, for instance Myrsine coriacea, Viburnum tinoides and Miconia theaezans, are commonly reported as shrubland and secondary forest species in montane ecosystems (Kappelle et al. 1994, 1996, Marin-Corba & Betancur 1997, Cortés 2003, Howorth & Pendry 2004, Lozano et al. 2007, Gutiérrez & Gaviria 2009) or even as oldgrowth forest species, albeit with lower constancy (Cleef et al. 1984, Kelly et al. 1994, Bussmann 2001, Schneider 2001, Cuello & Cleef 2009). The similarities with other cloud forests could be considered as an indicator of forest recovery towards becoming a mature forest. However, it is not possible to directly compare these studies because they were located at different elevations and had different sampling strategies (Table 7). Missing elements of the current flora are species of the family Podocarpaceae which are characteristic of Andean forest. The tree species Podocarpus oleifolius and Retrophyllum rospigliosii are common species of mature cloud forest (Bockor 1979, Cleef et al. 1984, Ulloa Ulloa & Jørgensen 1993, Kelly et al. 1994, Ramos & Plonczak 2007, Cuello & Cleef 2009) but seem to be rare in secondary forests (Schneider 2001, Ramos & Plonczak 2007). Other missing elements in the earlier sucesional stages are the fast-growing pioneer tree species (e. g.

The total species richness increases about 25 % when going from grassland to advanced forest; however, the rate of species accumulation is not always constant, it appears to be faster at the earlier successional stages. The increase in species diversification with successional development has been observed in other successional studies of neotropical ecosystems (e. g. Kappelle et al. 1994, Aide et al. 1995, Pascarella et al. 2000, Guariguata & Ostertag 2001). In terms of floristic composition, the family Asteraceae remains the most species rich family in the seral stages studied. However, the relative importance of Asteraceae decreases with time, while Rubiaceae, Melastomataceae and Piperaceae become more diverse. The same family composition was observed by Schneider (2001), in upper montane secondary forests and also in the Mucujún watershed (excluding the Orchidaceae epiphytes, which were not considered in the present study). The diversity pattern partially coincides with that described by Gentry (1988, 1995) in most Andean montane forests at mid elevation (1500 – 2500 m a.s.l.), where the Lauraceae, Rubiaceae and Melastomataceae are the most important families. However, species and individuals of the Lauraceae family are still underrepresented in these successional forests, Lauraceae species seem to prefer old-growth forest. The genera composition also follows the general patterns of the Andean flora where Miconia, Piper, Psychotria were the most diverse genera (Gentry 1988, 1995). In relation to species composition there are some similarities with other floristic inventories of secondary and mature Andean montane forests. Some of the same species described as frequent or dominant in other mature cloud forests include Hieronyma mori-

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Cecropia spp. Trema spp. etc.) which are common in the secondary succession of tropical areas (Guariguata & Ostertag 2001). The absence of these pioneer tree species could be a common pattern in montane forest succession after anthropogenic disturbances as their absence was also reported for other montane ranges (Ewel 1980, Aide et al. 1995) Similar vegetation patterns may occur in other montane ranges with similar environmental conditions and in particular those with similar land use patterns. For instance, the grasslands establishing in recently abandoned fields are characterized by high constancy and abundance of the introduced grass species Melinis minutiflora, described as Melinetum by Vareschi (1969) in the valley around Caracas in the Coastal cordillera of Venezuela. This type of anthropogenic savanna is frequently found in dry open places that are often burned (Vareschi 1969, Barger et al. 2003), this community shares floristic and structural similarities to the grassland communities described here. The Hedyosmum pseudoandromeda and the Protium tovarense community groups described by Meier (1998) occurring in the cloud forest zone of the Avila National park (1700 – 2200 m a.s.l., Coastal cordillera) have floristic affinities to the secondary forest described here. Particularly, the Myrcia fallax community (Meier 1998), a secondary forest growing on sites with anthropogenic disturbances, has structural similarities with the TSF Escallonia paniculata – Clethra fagifolia and ASF of Psychotria meridensis – Hedyosmum sp. in that they share about 20 % of the species. Also for the Talamanca Cordillera (2200 – 3000 m a.s.l.) in Costa Rica where land use characteristics seem to be comparable, seral communities with very similar structure and growth form composition have been described (Kappelle et al. 1994). Because of biogeographical differences, the species composition differs from the communities described here. However, there are similarities in the successional development, growth forms and the structure of different successional stages. In this study there are also several genera (e. g. Rubus, Monochaetum, Viburnum, Miconia, Weinmannia, Chusquea, Ageratina, Escallonia) with similar growth forms related to equivalent vegetation types, and species common to both studies are Myrsine coriacea, Macleania rupestris, Blechnum occidentale and Pteridium aquilinum. Worth mentioning is the fact that similarities to old growth forest communities are not evident. Classifications of old growth cloud forest in the Andes cordillera in Venezuela (Hetsch & Hoheisel 1976, Cuello & Cleef 2009) show lower floristic and structural affinities with the secondary forest described here; in general the species in common with these mature forests are less than 18 % (Table 7). This indicates that passive forest restoration needs at least several decades more of succession. The structural development of the forest during succession follows the same general patterns with

Fig. 8. Structure diagrams of successional communities of the LMCF: a) 1.1.1 Oxalis latifolia–subtype; b) 1.1.2 Psidium guineense subtype; c) 1.2 Ischaemum latifolium savanna; d) 1.3 Rhynchospora rugosa pasture; e) 1.4 Baccharis nitida shrubland; f) 2.1.1 Peperomia galioides TSF; g) 2.1.2 Escallonia paniculataClethra fagifolia TSF; h) 2.2.1 Turpinia occidentalis ASF; i) 2.2.2.1 Chusquea maculata subtype ASF ; j) 2.2.2.2 Palicourea demissa subtype ASF.

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Table 6. Structural characteristics of woody plants in the successional pathway of the LMCF.

'HQVLW\ ,QGKD D

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E

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7UDQVLWLRQDO6HFRQGDU\ )RUHVWQ $GYDQFHGVHFRQGDU\ IRUHVWQ

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Table 7. Structural comparison of woody components in secondary and mature forests in different locations.

Location and source

Mucujún watershed Andes Venezuela (2300–2600 m asl) plot size 0.1 ha (Schneider 2001)

Forest types MF (n=1) SF (n=1)

Species richness (shared) 2562 (20 %) 383 (38%)

Density

Basal area (m2 ha-1)

(ind.ha-1)

Total

Min/max

Heigh (m)

DBG H (cm) Mean

Max.

Mean

Max.

1500c 750b

55.2a

-

-

-

27

38

3240c 1290b

34.0a

-

-

-

13

17

20.1

30

La Carbonera Andes Venezuela (22002500 m asl) plot size 0.25 ha (Hetsch & Hoheisel 1976)

MF (n= 17)

602 (18 %)

905b

35.7b

15/54.6

22.3

Guaramacal Andes Venezuela (1850– 2580 m asl) plot size 0.1 ha, n=35 (Cuello & Cleef 2009)

LMRF

2662 (10 %)

2552a 650b

47.3a

26.6/70.8

8.2

123.0

9.4

32

UMRF

1692

3430a 727b

49.0a

29.8/85.7

8.5

127.3

8.95

23

10 yr (n=1)

91

1561c

12.1c

-

-

-

6.0

9

20 yr (n=2)

11.51

1546c

30.2c

26.2/34.2

-

-

7.0

14.5

35 yr (n=2)

181

2002c

48.3c

41.2/55.4

-

-

9.8

18.5

MF (n=3)

26.31

2442c

56.8c

48.9/62.1

-

-

11.3

27

8-20 yr (n=5)

211 (10%)

1248d 280b

9.8d

Coastal Cordillera Venezuela (1800– 1930 m asl) plot size 0.04 ha (Howorth & Pendry 2004)

Talamanca Cordillera, Costa Rica (2900–3000 m asl) plot size 0.1 ha (Kappelle et al. 1996)

10

17.4d 25-32 20.31 1603d 15.5 (10%) yr 628b (n=4) 1197d 60.7d MF 19.71 36.0 (5 %) (n=3) 634b a: dhb FPEGKEFPFGEKFP d: dhb FP PHDQWUHHVSHFLHVQXPEHUWRWDOVSHFLHVQXPEHUWUHHVSHFLHVLQWKHVHFRQGDU\IRUHVWDQGSORWVDWPDVO0) 2Od growth forest, /05)ORZHUPRQWDQHUDLQIRUHVW805)XSSHUmontane rain forest

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an increase of cover, mean stem diameter, basal area and height (e. g. Aide et al. 1995, Guariguata & Ostertag 2001, Howorth & Pendry 2004). These observations are comparable to those made in low land tropical forests (Guariguata & Ostertag 2001), where there is a sharp increase in the basal area and consequently woody species cover in the first decade of succession after which they slow down. However, the recovery of structural features in LMCF in the Mucujun watershed seems to be slower compared to other successional sequences (Kappelle et al. 1996, Howorth & Pendry 2004). The stem density is similar to the values reported for other secondary and mature forests (Table 6 and Table 7). However, compared to mature cloud forest the stem diameter is small. The stems are under 25 cm DBH and relatively homogeneous in diameter (CV under 44 %) while the diameter size distribution tends to be irregular with a high coefficient of variation in old secondary and in mature forests (Guariguata & Ostertag 2001). The basal area (BA) and mean tree height are relatively low compared to other secondary forests. The basal area observed in this study is comparable to forests in the Coastal cordillera with approximately 20 years of development, a 25 – 32 year old forest in Talamanca, and to low density stands in Guaramacal forest (Table 7). The low BA values can be explained by the absence of mid to large sized trees which account for a good portion of the total BA. Along with the structural development there is also a diversification of growth forms. The number of species of trees, treelets, shrubs and lianas-vine species increases with successional development, while forb and grass species decrease (Fig. 7). The same successional pattern was observed in Talamanca (Kappelle et al. 1994). The high number of lianas and vine species in the ASF is comparable to the lowland tropical forest. About 20 – 25 % of the total flora is shared with lowland forests (Gentry & Dodson 1987). Also the density of lianas in montane forests seems to be comparable to that observed in lowland forests (Homeier et al. 2010). The high number and cover of liana species found in the older secondary forest could be an indication of disturbances and slow recovery because lianas seem to be well adapted to disturbances in tropical ecosystems (Letcher & Chazdon 2009).

ondary forest communities studied were more closely associated with land use intensity, represented by the distance of the relevés from former farmhouses (Fig. 6). Generally, the farther a stand was located from farmhouses, the lower the land use intensity. The gradient analysis showed two clear tendencies in the successional pathways represented by two series of species groups occurring in certain secondary communities. These series are: 1) the Calycolpus moritzianus – Clematis populifolia series, corresponding mainly with the communities of TSF of Peperomia galioides and the ASF of Turpinia occidentalis which developed in stands closer to farmhouses, represented by the relevés plotted in the lower half of the right side of the NMDS diagram (Fig. 6, Table 2) the series of Piptocoma niceforoi – Miconia spinulosa, composed of the TSF of Escallonia paniculata – Clethra fagifolia and the ASF of Psychotria meridensis – Hedyosmum sp., occupy the upper part of the NMDS diagram (Fig. 6, Table 2). The forest communities closest to farmhouses are most likely those disturbed by various land uses, and are frequently dominated by the tree species Calycolpus moritzianus (Table 2). This species is promoted by farmers who use it as shade tree, firewood, and for construction on their farms (personal observation). Individuals of C. moritzianus are frequently found in pastures at low densities but become more dominant with increasing time after stand abandonment especially in the shrubland and TSF stages, but their dominance decreases with ongoing successional development. It is common to find decaying individuals of C. moritzianus in older secondary forests suggesting that there was once pasture there with scattered trees and then abandoned. Another good indicator of the Calycolpus moritzianus – Clematis populifolia series is the tree species Fraxinus uhdei. It has a higher abundance and constancy in the Baccharis nitida and Peperomia galioides communities (Table 2). This species was introduced in the 1950s and 60s through reforestation programs (Vilanova et al. 2008); the plantations were mostly established in accessible locations close to farms. The series of Piptocoma niceforoi – Miconia spinulosa represents succession under less intense land use. It corresponds to the secondary communities developed farther away from farmhouses. These relevés have a relatively high presence of Piptocoma niceforoi, but it does not occur as abundantly as C. moritzianus in the other series (Table 2). The colonization by P. niceforoi happens early in the GS communities (1.2 and 1.3, Table 2), where it is found mainly as a juvenile under 3 m tall. Once established the trees reach the higher layers and become important elements of the canopy and sub-canopy of the TSF and ASF (2.2 and 2.4.1, Table 2). The community is easy to identify in the landscape by the yellowish foliage of P. niceforoi. The two series seem to converge in the older stages of Palicourea demissa ASF (subtype 2.4.2, Table 2), with

Succession The current Andean landscape is the result of hundreds of years of land use which has had a large influence on the composition and development of the present vegetation pattern. It has been shown that time is the best predictor of forest development after the land has been abandoned (Fig. 6). This has also been documented in other cultural landscapes (Aide et al. 1995, Chinea 2002). However, the differences observed in the species composition within the sec-

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the presence of species such as Alchornea triplinervia that reach the canopy in older successional stages. In general, the succession of the LMCF parallels the phases proposed by Guariguata & Ostertag (2001 p. 200). The GS communities (group 1) represent the “initial colonization” phase still dominated by grasses, ferns and other pioneers. Although there were no fast-growing woody pioneers such as Cecropia spp. Heliocarpus sp. or Trema sp.; the TSF communities match the “early forest development” by the presence of short-lived pioneers such as species in the genera Baccharis or Monochaetum. The woody species established in this phase remained dominant in the ASF, namely, the species belonging to the Viburnum tinoides – Palicourea leuconeura group.

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the extensive cattle grazing observed in the Mucujún watershed, most likely has promoted the establishment of woody pionners species. Abandoned land is usually burned less often by humans (Aide et al. 2011). Frequent fires hinder the establishment of many woody plants through the destruction of seed banks or killing of the juveniles (Aide & Cavelier 1994), for example those in the Ischaemum latifolium community. On the other hand infrequent fires could prevent the dominance of invasive grass species (Aide et al. 2011). Therefore, when fire use is reduced and burning is not done in the dry season, the colonization and establishment of woody plants improves as was observed in the case of shrubs and treelets in the Asteraceae, Melastomataceae and Rubiaceae (Vareschi 1969), and this same outcome was observed on most of the abandoned stands in the Mucujún watershed. Another important factor favoring forest recovery was the early coverage of abandoned lands by species of the Viburnum tinoides – Palicourea leuconeura group together with the trees C. moritzianus, Myrcia fallax and P. niceforoi. These tree species grow in patches, have multiple stems and umbrella-shaped crowns. This structure provides excellent coverage of the soil creating favourable partly shaded microhabitats. Additionally, many of these species (e. g. Myrcia fallax, Miconia spp. Palicourea spp.) have fleshy fruits that attract birds that disperse the seed. As shown by other studies, the establishment of shade-tolerant species on one time agricultural fields is higher under remnant or planted trees (e. g. Guevara et al. 1992, Otero-Arnaiz et al. 1999). However, the establisment of the LMCF species seems to be relatively slow, probably due to the long time that the land was under management and the degree to which the LMCF’s original tree population was reduced thereby eliminating seed sources. The introduced tree species Fraxinus uhdei could hinder the establishment of other important native woody species. Fraxinux uhdei is a common component of the secondary communities in all vertical layers, especially in the TSF (Table 2). This species is an invasive pioneer which competes with and often replaces native plants at different successional stages. It seems that cattle grazing hampers the growth of young F. uhdei as they were frequently browsed in the relevés. Based on the results we conclude that the restoration of LMCF in traditional landscapes could be achieved by reducing (it’s not necessary to eliminate) the grazing intensity during the first 2 decades after field abandonment and by reducing the fire frequency. The recovery could be accelerated by planting species of the Viburnum tinoides – Palicourea leuconeura group in pastures or in disturbed stands. Another possibility for enhancing the recovery would be to plant or seed species belonging to the advanced successional stages and old growth forest in earlier secondary forests, e.g. Alchornea triplinervia (A. grandiflora), Hieronyma moritziana, Hedyos-

Forest restoration There are some indicators of forest recovery, including increasing species richness, diversity of growth forms, forest cover and woody species densities (Table 5, 6 and 7). However, the older successional forest (ASF) is still in an intermediate recovery stage, between early and late forest development according to the phases proposed by Guariguata & Ostertag (2001). About 50 years were necessary to achieve the current stage of development of the ASF stands. The species Billia rosea, Nectandra truxillensis, Prunus moritziana, Clusia multiflora, Wettinia praemorsa are known to be important components of the cloud forest (Sarmiento et al. 1971, Hetsch & Hoheisel 1976, Schneider 2001, Cuello & Cleef 2009). However, they only occur with very low constancy and are mostly present as juveniles in the understory and ground layer in the ASF stands. It will take several more decades for these species to become a structural part of the canopy composition and to develop the floristic and structural properties of mature forests. The slow recovery rates coincide with observations made in other mountain ranges with similar land use characteristics (Aide et al. 1995, Sarmiento et al. 2003). However, taking into account the long time that the study stands were under agricultural use (Gutiérrez in prep.), and the variability of land use intensity and practices, the recovery has been relatively successful, especially when compared to other locations where the cultural land use resulted in degraded savannas (e. g. Cavelier et al. 1998). There are some important factors that contributed to the establishment of secondary forest in the Mucujún watershed. Introduced African grass species (e. g. Melinis minutiflora) have a negative impact on the recovery of forest ecosystems by hindering the establishment of woody species (Aide & Cavelier 1994, Sarmiento 1997, Pivello et al. 1999, Hoffmann et al. 2004). However, it has been observed that low grazing intensity reduced the biomass of the palatable grass species allowing the establishment and growth of woody pioneers (Posada et al. 2000). Therefore,

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mum spp., Clusia multiflora (Clusia spp.), Prunus moritziana, Palicourea demissa, Miconia lonchophylla, and Myrcia fallax. These species seem to have the physiological plasticity, documented in A. triplinervia (García-Nuñez et al. 1995, Rôças et al. 1997), allowing them to adapt to site conditions early on and in later successional stages and could facilitate the recovery process. However, for most of the species in question we do not know anything about their autoecology. More studies will be needed to understand them and their role in restoration ecology of the Andean cloud forest. The phytosociological approach used to describe the seral communities in the LMCF belt in the Mucujún watershed, has allowed the development of a successional model in abandoned cattle pastures. The species replacement associated to the time after abandonment, site properties and land use intensity could be considered to undertake the ecological restoration on montane forest in areas with similar conditions or to accelerate the recovery of intermediate successional stages.

References Aide, T., Ruiz-Jaen, M. & Grau, H. (2011): What is the state of tropical montane cloud forest restoration? In: L. Bruijnzeel, F.N. Scatena & L.S. Hamilton (eds.), Tropical Montane Cloud Forests: Science for Conservation and Management, pp. 101 – 109. – Cambridge University Press, New York. Aide, T.M. & Cavelier, J. (1994): Barriers to lowland tropical forest restoration in teh Sierra Nevada de Santa Marta Colombia. – Restor. Ecol. 2: 219 – 229. Aide, T.M. & Grau, H.R. (2004): Ecology: Globalization, Migration, and Latin American Ecosystems. – Science 305 (5692): 1915 – 1916. Aide, T.M., Zimmerman, J.K., Herrera, L., Rosario, M. & Serrano, M. (1995): Forest recovery in abandoned tropical pastures in Puerto Rico. – Forest. Ecol. Manag. 77: 77 – 86. Aide, T.M., Zimmerman, J.K., Rosario, M. & Marcano, H. (1996): Forest recovery in abandoned cattle pastures along an elevational gradient in northeastern Puerto Rico. – Biotropica 28: 537 – 548. Aldana, A.A. & Bosque, J.B. (2008): Cambios ocurridos en la cobertura/uso de la tierra del Parque Nacional Sierra de la Culata. Mérida-Venezuela. período 1988 – 2003. – Geofocus 8: 139 – 168. Ataroff, M. (2001): Venezuela. In: M. Kappelle & A.D. Brown (eds.), Bosques Nublados del Neotrópico, pp. 397 – 442. – Instituto Nacional de Biodiversidad (INBio), Costa Rica. Ataroff, M. & Rada, F. (2000): Deforestation impacts on water dynamics in a Venezuelan Andean cloud forest. – Ambio 29: 440 – 444. Ataroff, M. & Sarmiento, L. (2003): Diversidad en los Andes de Venezuela. I mapa de unidades ecológicas del estado Mérida. CD-ROM. – Universidad de Los Andes, Mérida, Venezuela. Ataroff, M. & Sarmiento, L. (2004): Las unidades ecológicas de los Andes de Venezuela. In: E. La Marca & P. Soriano (eds.), Reptiles de los Andes de Venezuela, pp. 9 – 26. – Biogeos, Mérida. Barger, N.N., D’Antonio, C.M., Ghneim, T. & Cuevas, E. (2003): Constraints to colonization and growth of the African grass, Melinis minutiflora, in a Venezuelan savanna. – Plant. Ecol. 167 (1): 31 – 43. Bockor, I. (1979): Analyse von Baumartenzusammensetzung und Bestandesstrukturen eines andinen Wolkenwaldes in Westvenezuela als Grundlage zur Waldtypengliederung. – Dr. Dissertation, Universität Göttingen, FRG. 138 pp. Bussmann, R.W. (2001): The montane forest of reserva biológica San Francisco (Zamora-Chinchipe, Ecuador) vegetation zonation and natural regeneration. – Die Erde 132: 9 – 25. Cavelier, J., Aide, T.M., Santos, C., Eusse, A.M. & Dupuy, J.M. (1998): The Savannization of moist forests in the Sierra Nevada de Santa Marta, Colombia. – J. Biogeogr. 25: 901 – 912. Chinea, J.D. (2002): Tropical forest succession on abandoned farms in the Humacao Municipality of eastern Puerto Rico. – Forest. Ecol. Manag. 167: 195 – 207. Cleef, A.M., Rangel, O., Van der Hammen, T. & Jaramillo, R.M. (1984): La vegetación de las selvas del transecto Buritaca. In: T. van der Hammen & P. Ruiz (eds.), La Sierra Nevada de Santa Marta (Colombia) – Transecto Buritaca – La Cumbre, pp. 267 – 406. – J. Cramer, Berlin, Stuttgart. Cortés, S.P. (2003): Estructura de la vegetación arbórea y arbustiva en el costado oriental de la Serranía de Chía (Cundinamarca, Colombia). – Caldasia 25: 119 – 137. Cortés, S.P. (2008): La vegetación boscosa y arbustiva de la cuenca alta del río Bogotá. In: T. van der Hammen, A.P. Preciado & P.P. E (eds.), Studies on tropical Andean ecosystems 7, p. . – J. Cramer, Berlin, Stuttgart.

Resumen: Se estudió la recuperación del la selva nublada montano baja en pastizales abandonados ubicados entre 1800 y 2400 m s.n.m., en la cuenca del Río Mucujún. Para tal fin, se utilizó la estrategia de muestreo de espacio por tiempo, ubicando 83 inventarios en un gradiente sucesional de unos 50 años, desde pastizales recientemente abandonados hasta bosques secundarios avanzados. Se registraron 368 plantas vasculares pertenecientes 205 géneros y 82 familias. Un total de ocho comunidades sucesionales fueron determinadas mediante una clasificación phytosociológica de los inventarios. Los bosques sucesionales se caracterizaron por la frecuencia y abundancia de las especies pertenecientes al grupo Viburnum tinoides – Palicourea leuconeura. Estas especies logran establecerse en estadios tempranos de sucesión y forman una parte importante de la estructura del bosque secundario. El tiempo fue la variable que mejor determina la recuperación del bosque, mientras que la intensidad de uso determinó dos caminos sucesionales diferenciados por su composición florística. La recuperación del bosque es evidente sobre todo en su estructura y diversidad, sin embargo la recuperación de la diversidad florística original está aún en una fase intermedia. Acknowledgement: The study was possible with the support from the Fundación Gran Mariscal de Ayacucho (FUNDAYACUCHO) and the Deutscher Akademischer Austauschdienst (DAAD), the Verband der Freunde der Universität Freiburg i. Br., the International Ph.D. Program “Forestry in Transition” of the Faculty of Forest and Environmental Sciences of the University of Freiburg, and the Universidad de los Andes. The authors gratefully acknowledge the personal of the Jardín Botánico de la Facultad de Ciencias of the Universidad de los Andes and Darwin Gutierrez for the support during the field sampling and species identification. We thank Antoine Cleef, Bernhard Thiel and an anonymous reviewer for improving the final version of the manuscript.

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