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UvA-DARE is a service provided by the library of the University of Amsterdam (https://dare.uva.nl)

Flora, vegetation and ecology in the Venezuelan Andes: a case study of Ramal

de Guaramacal

Cuello Alvarado, N.L.

Publication date

2010

Link to publication

Citation for published version (APA):

Cuello Alvarado, N. L. (2010). Flora, vegetation and ecology in the Venezuelan Andes: a case

study of Ramal de Guaramacal. Universiteit van Amsterdam, Institute for Biodiversity and

Ecosystem Dynamics (IBED).

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Chapter 2

The forest vegetation of Ramal de Guaramacal in the

Venezuelan Andes

Nidia L. CuelloA. and Antoine M. Cleef PHYTOCOENOLOGIA 39(1):109-156.2009

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2.1 INTRODUCTION

Montane forests of the northern Andes are fragile ecosystems of significant biological and ecological diversity with a complex biogeographical history, and playing a major role in the regional hydrological balance (Gentry 1995; Cavalier & Goldstein 1989; Cavelier et al. 1996; Holder 2006; Kappelle & Brown 2001). Despite increased attention and conservation interest of the northern Andean forest ecosystem over the past two decades (Henderson et al. 1991; Churchill et al. 1995, Gentry 1995; Luteyn & Churchill 2000; Kappelle & Brown 2001; Van der Hammen et al. 1984), studies on montane forests of the Venezuelan Andes remain limited in area, subject and time. The majority of studies have been carried out in the montane forests of the state of Mérida. The silvicultural studies of La Mucuy and La Carbonera (Lamprecht 1954; Veillon 1965, 1985), vegetation ecology (Vareschi 1953, 1956; Yánez 1998); floristic analysis (Kelly et al. 1994, 2004); and several studies focusing on different aspects of ecophysiology, population ecology and hydroecology of cloud forests (Brun 1979; ICAE 2005), aspects of diversity, structure and biogeography on a succesional and mature forest stands close to the town of Mérida (Schneider et al. 2000; Schneider 2001) are particularly noteworthy. Few Andean montane forest areas outside of Mérida State have been studied (Bono 1996; Ortega et al. 1987; Cuello 1996, 1999, 2002 and Dorr et al. 2000). Beyond the Andes, other montane forest areas previously studied are Coastal Cordillera (Huber 1986; Howorth & Pendry 2006); Cerro El Avila (Vareschi 1955; Steyermark & Huber 1978; Meier 2004) and Cerro Copey in Margarita Island (Sugden 1985).

On tropical mountains, the altitudinal limit of forest formations varies with latitude (Troll 1959, 1968) or in response to local or regional peculiarities of topography or climate (Grubb & Whitmore 1966; Monasterio & Reyes 1980; Van der Hammen & Cleef 1986; Van der Hammen 1995; Lauer et al. 2001; Richter 2003). In the tropical Andes, the distribution of vegetation types and their qualitative and quantitative composition are thought to be determined largely by gradients of temperature, rainfall, and relative humidity (Van der Hammen & Cleef 1986; Van der Hammen 1995), and horizontal precipitation and mist deposition (Bendix et al. 2006; Richter & Moreira-Muñoz 2005). Gradients of temperature have pronounced effects on the pattern of vegetation zonation, especially at the limits of the upper forest line (Troll 1973; Rundel 1994).

In the northern Andes, altitudinal vegetation zonation has been distinguished as lowland tropical forests from 0-1000 m, lower montane (LMRF) or subandean forests from 1000-2300 m, upper montane (UMRF) or Andean forests from 2300-3500 (3200-3600) m and high Andean forests from 2300-3500 (3000-2300-3500)-3900 m. Open paramo vegetation is found over 3200-3900 m up to the nival zone (>4800-5000 m) (Cuatrecasas 1934, 1958; Van der Hammen 1974; Cleef et al. 1984; Van der Hammen & Hooghiemstra 2001).

The existence of altitudinal zonation with discrete vegetation belts in the northern Andes (Cuatrecasas 1958; Van der Hammen 1974) versus a continuous change in species composition in tropical mountains, have been subject of discussion. Some quantitative studies in other tropical mountain areas support the existence of

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discrete vegetation belts (Kitayama 1992; Hemp 2006), while other studies have been less conclusive about zonation (Nakashizucha et al. 1992; Kappelle et al. 1995; Lieberman et al. 1996; Vásquez & Givnish 1998). Differences in their scope, sampling methods, and analytical techniques may account for the discrepancy in their conclusions (Cuello 1996, 2000; Hemp 2006).

In the Venezuelan Andes, six ecological altitudinal zones have been recognized (Sarmiento et al. 1971; Monasterio 1980; Monasterio & Reyes 1980; Ataroff & Sarmiento 2004): (1) a Basal zone from piedemont to 1000 m, (2) a Subandean zone between 1000 and 2000 m, (3) a Lower Andean zone between 2000 and 3000 m, (4) an Upper Andean zone between 3000 and 4000 m, (5) a High Andean zone between 4000 and 4800 m and (6) a Snow zone above 4800 m. The distribution of vegetation types along altitudinal zones in the Venezuelan Andes differs between humid and dry slopes (Sarmiento et al. 1971; Monasterio 1980; Monasterio & Reyes 1980; Ataroff & Sarmiento 2004).

As elsewhere, the Venezuelan Andes are suffering increased human intervention. Many areas of montane forests continue to be converted into areas of agricultural or other land uses, while basic biodiversity studies remain scarce (Ataroff 2001). Fortunately, due to their steep slopes and inaccessibility there are still large parts of the Venezuelan Andes, with or without legal protection, where natural vegetation remains relatively undisturbed. One of such areas is Ramal de Guaramacal, the larger part of which is enclosed within a national park (Cruz Carrillo or Guaramacal National Park), not presently subject to a strong human intervention.

The main goal of the present study is to identify, define and characterize the montane rain forest vegetation of Ramal de Guaramacal and to establish a syntaxonomic scheme or classification, based on analysis of the physiognomy, floristic composition, ecological relations and spatial distribution of the different vegetation communities. This work was conducted within the framework of a larger project aiming to study the floristic and vegetation diversity of the Guaramacal National Park (Cuello 1999, 2000, 2002, 2004). The classification of the paramo vegetation of the Guaramacal summit area is described separately (Cuello & Cleef 2009 b, c).

2.2 STUDY AREA

Geography

Ramal de Guaramacal is a mountain range extending approximately 30 kilometers northeast towards the eastern end of the Venezuelan Andes between 9° 0521‟ N and 70° 0020‟ W. (Fig. 1). Parts of the Boconó Municipality in the State of Trujillo and Sucre Municipality of the Portuguesa state are included. This formation, in its larger extension, reaches altitudes over 2000 m. The Guaramacal range includes summits of 3130 m in Páramo de Guaramacal; 2970 m in Páramo El Pumar; 2800 m in Páramo Agua Fría and 2600 m in Páramo Los Rosarios.

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Much of the surface area of the Ramal de Guaramacal is protected by the Gral. Cruz Carrillo Nacional Park (or Guaramacal National Park), which includes, from the lowermost level of 1600 m, an approximate surface area of 21,466 hectares. Climate

The climate of the Venzuelan Andean cordillera, as in the whole country is largely determined by the Intertropical Convergence Zone (ITCZ). The great altitudinal interval with respect to the Llanos, in combination with full exposition to trade winds, causes high precipitation of at least 3000 mm/year from low altitude up to approximately 2400 m on the southeastern slopes. From this position upwards and northwards, precipitation decreases to around 2000 mm on the northern slopes (Reaud-Thomas 1989). The climate is characterized by a dry season from November to March and a rainy season from April to October. Maximum precipitation occurs during June and July (Cuello & Barbera 1999). Temperatures remain low throughout the year, averaging around 18 to 20 oC between 1000 to 1500 m, and 9 to 12 oC in the zone above 2500 m. According to Grubb (1977) and Sarmiento (1986) the decrease of temperature with altitude (lapse rate) is around 0.6 oC/100 m. Above 2000 m the nights are cold and seasonal frosts may occur at altitudes over 2,500 m (Reaud-Thomas 1989; Urriola 1999).

Geology

Guaramacal mountain range is aligned to the South to the Boconó fault (Schubert 1980) and to the North of the Calderas fault. It constitutes a homogeneous block separated from the rest of the mountain range (and limiting in the North and West) by the interandean valley of Boconó River. To the South (Llanos slope) steep slopes descend towards the Calderas fault, from there, the smaller mountain formations decrease in altitude towards the piedemont at 400 m (Urriola 1999). This mountain chain is the product of orogenic processes which built the equatorial Andes. Some of the most important geological formations of the Venezuelan Andean Cordillera are displayed, from the oldest bedrock of the Precambric and the Paleozoic, to more recent deposits of the Neogene (Tertiary and Quaternary period). The most predominant geologic formations in Ramal de Guaramacal are: Sabaneta (Paleozoic), Palmarito (Paleozoic) y El Santuario or Gobernador (Tertiary), and, in the North slope at the end of the Boconó Fault, the Sierra Nevada formation (Grupo Iglesias, Upper Precambric) (Ministerio de Minas e Hidrocarburos, s/f; Urriola 1999) is present.

In the Páramo de Guaramacal area (Sector A), the Sabaneta formation predominates. This old formation consists of a sequence of gray to brown sandstones intercalated with limolites and red to red-violet sandstones. Around Páramo El Pumar (Sector C), the Santuario, or Gobernador, formation (Tertiary) arises. This formation consists of 80% friables to hardened gray quartzeous sandstones becoming brownish under weathered conditions, being locally conglomerated in thick layers with intercalations of light colored limolites, and laminations of dark lutites, being occasionally calcareous and making up to 20% of the formation (Urriola 1999).

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Soils

The soils of Guaramacal are determined by a set of climatic, geologic, and topographic conditions as well as vegetation characteristics for this mountainous landscape (Marvez & Schargel 1999). Periglacial phenomena are also an important landscape feature at the highest altitudes with páramos and high Andean forests. This is also true for the Guaramacal range. Glacial lakes, different types of morraines, roches mouttonnées, etc. account for the past presence of a glacial snow and ice cap in the summit zone during past glacial times.

The high precipitation of the region favors intense lixiviation and acidification of the soils; thus, acid soils of low pH (<5) predominate. The soils under forests and páramos generally display high organic matter content due to low temperatures and acid conditions which markedly reduce the activity of soil microorganisms. The steep slopes and the elevated water content of soils on weathered rocks, favor mass movements and landslides at different spatiotemporal intervals; also determining substantial variability in soil depth and coarse fragment content (diameter > 2 mm). The predominance of sandstone of the Sabaneta and El Santuario formations accounts for a high sand content of the soils. Nevertheless, the presence of fine grained sedimentary lutites causes clayey soils in some localities. The soils of the study area are predominantly of the Entisols, Inceptisols and Ultisols order (Marvez & Schargel 1999). Morphological and chemical characteristics of some of the soil profiles representative of Ramal de Guaramacal (from Marvez & Schargel 1999) are presented in Appendix 3.

Vegetation

The vegetation of Guaramacal Park area is predominantely represented by montane rain forests with height and density decreasing with altitude (Cuello 2002). Subpáramo and of páramo vegetation is locally evident from 2700 m; however, forest vegetation patches mixed with open páramo can be found to elevations of up to 2900-3000 m in the summit zone.

The montane forests of Ramal de Guaramacal are in the Tropical Lower Montane Very Wet Forest and Tropical Montane Rain Forest zones of the Holdridge climatic life zone system (Ewel et al. 1976). According to Huber & Alarcon (1988) the forests of Guaramacal are classified as „Bosques ombrófilos submontanos/montanos siempreverdes‟. According to the bioclimatic classification proposed by Costa et al. (2007) for the Andes of Merida, both forest and páramo zones of Guaramacal correspond to the Mesotropical belt.

Previous geobotanical explorations and flora research

Botanical exploration of Ramal de Guaramacal started in the 1960‟s when a rural road between Mosquey (Boconó - North slope) and the village of Guaramacal (South slope) was constructed, to provide access to a complex of communication antennas installed in the summit of Páramo de Guaramacal (Ortega et al. 1987, Dorr et al. 2000). The first floristic exploration was conducted by G.C.K. Dunsterville in 1963, who exploited this new track to collect orchids while workers still were felling trees for the new road (Dunsterville & Garay 1965, Dorr

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1999). Julian Steyermark visited Guaramacal at a later date, accompanied by Marvin Rabe in 1966; in 1970, with Basset Maguire and Celia K. Maguire; and finally alone in 1971 (Dorr 1999).

During the 1970‟s several botanists visited the Guaramacal range; the late Luis Enrique Ruiz-Terán, being among them in 1973. Dana Griffin III collected bryophytes in 1975; José Cuatrecasas in 1978 for Espeletiinae (Asteraceae), and James L. Luteyn in 1978 mainly for Ericaceae and other species of the páramo. Manuel López-Figueiras and Mason Hale also collected lichens in Guaramacal in 1979 (Dorr 1999).

During the 1980‟s, staff of Herbario Universitario (PORT) of the Universidad Nacional Experimental de los Llanos „Ezequiel Zamora‟ (UNELLEZ) in Guanare, headed by Francisco Ortega, Basil Stergios and Gerardo Aymard, made the first effort to catalogue the Guaramacal flora (Ortega et al. 1987, Rivero & Ortega 1989). In many of their visits they were accompanied by botanists from the United States of America, among them were Ronald Liesner and Henk van der Werff from the Missouri Botanical Garden; Alan R. Smith of the University of California, Berkeley; and Laurence J. Dorr, previously of the New York Botanical Garden (Dorr 1999).

In 1995, the first author initiated a floristic research project studying vegetation across an elevational gradient using plots for the vegetation analysis (Cuello 1996), consequently initiating explorations in other, more remote, areas of Ramal de Guaramacal. Simultaneously, Basil Stergios (UNELLEZ) and Laurence Dorr (Smithsonian Institution) initiated a project to document the Flora of Guaramacal. This project initially served as a supporting framework for the present study of the Guaramacal vegetation. The combined efforts of botanical exploration and floristic survey, together with contributions from an integrated multidisciplinary team of researchers from UNELLEZ, generated publication of a reference book dealing with several aspects of the nature of the Guaramacal National Park (Cuello 1999). In addition to the first results detailing forest composition and diversity (Cuello 1996, 2000, 2002, 2004), a first „Catalogue of the vascular plants of Guaramacal National Park‟ (Dorret al. 2000) has also been published. This catalogue accounts for a total 147 families, 517 genera and 1227 species of vascular plants.

During the last twenty years, as a result of continued integrated survey and botanical exploration, ca. 40 species new to science have been discovered in the Guaramacal range (Morillo 1988; Axelius & D‟ Arcy 1993; Badillo 1994, Carnevali & Ramírez 1998; Aymard et al. 1999; Taylor 2002; Stergios & Dorr 2003; Stancik 2004; Niño et al. 2005; Cuello & Aymard 2008, for the species list see also Stergios 1999 and Dorr et al. 2000).

The combined intensive vegetation surveys by the first author and botanical explorations by B. Stergios, L. Dorr and M. Niño (UNELLEZ) over the past ten years, have resulted in the addition of several new records to the Dorr et al. (2000) catalogue.

Collections of non-vascular plants have been neglected in Guaramacal to date. Despite the collections made during the 70‟s, a published list of bryophytes and lichens from Guaramacal does not yet exist. Only a few of the most prominent and

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conspicuous of bryophyte and lichen species present have been recently collected by first author, mainly from páramo vegetation. Many of the collections are undetermined at PORT, with only a present account of ca. 55 bryophyte species and 20 species of lichens.

2.3 METHODS

Field surveys

The study was carried out during 1995, 1996, 1999, 2003, 2005 and 2006 (see Appendix 2.). Montane forest community composition of Ramal de Guaramacal, Venezuelan Andes, was studied along the altitudinal gradient on both sides of the range with different slope expositions. The study area was divided into three sectors: (A) Guaramacal, (B) Agua Fría and (C) El Santuario (Fig. 2.1). Sector A corresponds to both slopes (N and S) in the central and higher part of the mountain range. This area is crossed North-South by a road that leads up to the telecommunication repeater antennas at the top of the mountain (Páramo de Guaramacal) and then descends to the village of Guaramacal on the South slope. Sector B comprises both slopes of the northeastern end of Ramal de Guaramacal with. This sector includes samplings sites named after nearby villages and rivers. Such sites are “La Peña” and “Río Frío” on the South slope, Trujillo state; “La Divisoria” and “El Alto”, at the border of Trujillo and Portuguesa states on the South slope; and “Laguna Negra” in Trujillo state and “El Mogote”, at the border of Trujillo and Portuguesa states on the North slope. Sector C corresponds to the north-western end of the Park, and includes the site known as “Qda. Honda – El Santuario”.

Within the park access remains limited to the only existing road in the Guaramacal sector, whereas the other sites (La Divisoria, El Alto, El Mogote, Agua Fría, Río Frío, Laguna Megra and El Santuario) could only be reached on foot using new or existing pathways in the forest. The old paths crossing the mountain are also locally known as “caminos reales”; having previously served as commercial connections between towns located on the South side of Ramal de Guaramacal and the city of Boconó and its surroundings.

On the South slope of Agua Fría sector it was also possible to survey forests below the Park boundaries (1350 - 1550 m). The discovery of these natural forests at low altitude provided the opportunity to document information on this region of the country. No other forest inventories are known from slopes at these altitudes. On the North slope of the same sector, the lowest plot was at 1650 m, near the locality of El Mogote, also located outside the Park boundary. For a more complete description of the study area the reader is referred to Cuello (1999).

The forest survey contained a total of 44 samples (total area 3.705 hectares) located at different altitudes between 1300 and 3000 m, distributed throughout the different Park sectors (Fig. 2.1, Appendix 2). The samples include 35 plots of 1000 m2 (20 m x 50 m, divided in subunits of 10 m x 10 m) each. The forests located at higher elevations, due to access difficulties, low stature and low diversity of species, were surveyed in smaller plots. These include one plot of 500 m2, one of

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400 m2, one of 300 m2, three of 200 m2, two of 100 m2 and one of 50 m2 (see Table 2.2(b), Appendix 2).

Figure. 2.1. Geographic position of Ramal de Guaramacal in the Andes of Venezuela, with the outline of the National Park. Park sectors: A-Guaramacal, B-Agua Fría, C-Qda. Honda-El Santuario.

A B

C

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The altitudinal intervals between sampling sites were variable between 30 to 150 meters. The plots were selected taking into account accessibility, topography and vegetation physiognomy. Thus, plots were laid out in relatively accessible sites of less steep slope where possible and under a rather continuous canopy. In the highest and steepest part of the South slope, in the “Guaramacal” sector, the plots were laid out in sites where forest had been affected by nearby landslides. This was inevitable reflecting natural conditions, since the South slope is steeper than the North slope (Fig. 2.11). On the South slope of Agua Fría sector, the maximum altitude surveyed was 2125 m. Traveling by foot to this altitude was extremely difficult as the mountain relief in this sector is comprised of contiguous small ranges with low summits (1900-2000 m). From 1300 m (altitude of base camp), these lower summits had to be crossed (with lower valleys between at 1700 m) in order to reach the forest limits at around 2450 m. Unfortunately, the opportunity to arrive at this location remained unrealised at this time.

Within each plot, all rooted individuals – trees, shrubs, lianas, tall and thick-stem or climbing terrestrial herbs and hemiepiphytes– ≥ 2.5 cm dbh (diameter at breast height, taken at 1.3 m from the base of the trunk, or lower for shrubs and thick-stemmed herbs) were recorded, labeled with numbered aluminum tags and their dbh and height recorded. The 2.5 cm dbh minimum size was chosen to include most of the small woody understory species, as well as lianas and hemiepiphytes, and to make samples comparable with the studies of Gentry (1982, 1992, 1995). Epiphytes, non-vascular plants, small herbs and other growth forms with stems < 2.5 cm were not surveyed.

Height of trees was estimated using a 2 m clipper pole as reference. Multistemmed species were counted as single individuals, but entire stem diameters were recorded for calculation of basal area. The same criterion was applied to multiple aerial roots of hemiepiphytes, such as Clusia.

Individuals were assigned to morphospecies; a voucher sample of each morphospecies collected from each plot. For ambiguous species multiple vouchers were collected. Morphospecies were later matched for all plots. As voucher samples from plots were mostly sterile, general collections of fertile specimens outside the plots were also made.

Some individuals (mainly lianas or very tall trees) could not be vouchered. In these cases, only registry of growth form, dbh and height were taken. For each site, collections and observations of other species not included in the surveys, herbs and epiphytes for instance, were made. In total over 2000 botanical numbers were collected under the number of N. Cuello (et al.) from nr. 915 to 2900 and A. Licata from nr. 150 to 690 (see Appendix 1).

Data processing and analysis

Identification and processing of botanical specimens was made at Herbarium PORT of the Universidad de los Llanos (UNELLEZ) in Guanare, Venezuela. Other herbaria, such as MO and US, were also consulted. Some specimens were sent to specialists at other institutions to confirm identification. All specimens

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collected have been deposited at PORT, some duplicates have been sent to VEN, MER, MERF, MO and US.

All the information and field data were stored and handled using Microsoft Excel. The total listing of the inventoried species together with their respective collection numbers appears in Appendix 1.

For the physiognomic characterization of the forests structural profiles of 20 m x 10 m (Fig. 2.2-2.10) were elaborated in the direct neighbourhood of some of the surveyed plots. The sites selected for the elaboration of profiles are georeferenced in Appendix 2.

A data matrix of the relative abundance of 360 species and 44 plots was processed with TWINSPAN (Hill 1979) using the program PC-Ord 4 (McCune & Mefford 1999). The resulting TWINSPAN was interpreted in terms of syntaxonomical classification of the vegetation, on the basis of floristic affinities, according to the Zürich-Montpellier approach (Braun-Blanquet 1979, Westhoff & van der Maarel 1973).

For forest descriptions we followed Cuatrecasas‟s (1934) classification of subandean, Andean and high Andean forest. However, for discussion and comparison with other montane forests, we also referred to the equatorial montane rain forest zonation of LMRF, UMRF and SARF by Grubb (1977) used also elsewhere in the equatorial tropics.

2.4 RESULTS

Flora diversity

A total of 388 morphospecies with dbh ≥ 2.5 cm, corresponding to 189 genera and 78 families of vascular plants, were recorded from the 45 forest plots in Ramal of Guaramacal. These include: 4 families, 6 genera and 13 species of pteridophytes; 5 families, 13 genera and 19 species of monocots; 68 families, 170 genera and 355 species of dicots; and 1 gymnosperm species Podocarpus oleifolius var. macrostachyus. From the total of 388 morphospecies, 309 were identified to species level, 55 to genus, 9 to family and 15 were not identified. An additional 177 species of vascular epiphytes, herbs and small shrubs were annotated and collected for forest description, but not documented in the plot surveys. All the species registered and collected from the plots are listed in Appendix 1. Table 2.1 presents the most speciose families and genera based on the plot data from this study. Six families were represented by 20 or more species, while 3 genera were found with 10 or more congeners.

Forest structure

Table 2.2 (a and b) summarizes the structural parameters of the different plots by sector, slope exposure and elevation. A total of 14,895 individuals with dbh ≥ 2.5 cm were recorded in a total of 3,705 ha of accumulated forest plot. The number of individuals per 0.1-ha plot varied from 154 to 602, with an average density of 372.

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On the North slope, density in 0.1 ha plots increased with altitude up to 2480 m, decreasing towards 2890 m. However, in forest plots (SARF) over 2900 m altitude a high density of low diameter individuals occurs with an average density of 110.02 individuals in 100 m2 plots (about to 1100 individuals extrapolated to 0.1 ha). The canopy height was variable within plots, but generally decreased with altitude. Taller emergent trees were found at every altitude. Maximum and mean diameter was variable among plots, the individual with the greatest diameter (127.3 cm) found at 2480 m.

Table 2.1. Most diverse woody plant families with dbh ≥ 2.5 cm (>10 species) and genera (>5 species) in the plots of the montane forests of Ramal de Guaramacal, Andes, Venezuela.

Families Species Genera Species

LAURACEAE 34 Ocotea 21 RUBIACEAE 30 Miconia 19 MELASTOMATACEAE 27 Eugenia 13 MYRTACEAE 24 Piper 8 ASTERACEAE 20 Persea 7 MYRSINACEAE 12 Palicourea 6 ERICACEAE 12 Psychotria 6 EUPHORBIACEAE 10 Cybianthus 5

Table 2.2(a) Summary of structural parameters for each forest 0.1 ha plot by slope and Sector of Ramal de Guaramacal, Venezuela. Species richness, number of individuals, basal area, mean and maximum height, canopy height, mean and maximum diameter (in cm).

Slope

Sector Alt. m Plot Nr. Num spp Num Ind Basal Area Tree height Diameter Max Med Canopy Max Med

North Guaramacal 1850 5 36 182 4.57 4 7.3 10-20 64.0 9.2 1960 1 41 358 2.66 20 8.0 10-15 38.2 7.2 2070 19 46 446 6.64 24 7.6 10-18 114.6 7.9 2100 2 35 401 3.31 18 6.5 10-15 62.1 7.1 2170 18 41 316 5.00 26 9.4 10-20 90.0 9.3 2300 3 50 377 8.57 21 7.1 8-15 111.4 10.5 2350 20 60 479 5.19 22 8.1 8-15 60.0 7.9 2400 4 59 547 6.91 19 6.5 6-15 108.2 8.3 2480 17 36 602 4.59 23 7.0 6-14 127.3 6.3 2580 16 33 413 5.67 19 8.6 6-14 69.0 9.4 2750 39 41 458 5.2 17 9.8 7-14 42.7 8.6 2870 37 18 231 4.35 16 8.2 6-12 44 10.1 2890 35 27 423 3.65 12 6.7 5-10 40 8 Agua Fría 1830 14 53 390 4.94 24 7.3 10-20 64.0 9.2 1900 25 60 320 7.08 28 10.8 8-24 116.5 7.8 2100 26 44 492 5.15 26 9.4 9-23 61.0 6.9 2260 27 61 438 5.43 24 9.1 7-18 105 8.2 North-west (Qda. Honda) 1880 21 43 227 4.65 32 11.6 14-29 83 9.9 2100 22 55 324 5.35 30 10.4 13-26 70 8.5 2250 23 35 257 4.03 18 8.6 7-14 50 9.5

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Slope

Sector Alt. m Plot Nr. Num

spp Num Ind Basal

Area

Tree height Diameter Max Med Canopy Max Med

Guaramacal 1950 7 36 500 4.62 22 8.9 9-18 76 6.32

2100 9 29 301 2.82 20 7.7 9-13 76 7.84

2300 8 31 306 2.98 21 7.9 9-16 41.4 8.3

2470 6 31 378 2.98 13 6.5 8-11 50 7.7

2580 24 34 309 3.78 18 8.6 8-14 56 8.0

South Agua Fría 1330 28 40 154 3.76 28 10.4 10-24 69 11.7

1450 29 45 191 3.63 27 8.8 10-24 82 8.6 1550 13 47 265 5.44 24 11.7 13-22 123 8.68 1600 10 52 328 4.95 25 7.3 12-20 105 7.93 1770 31 45 376 4.37 28 11.5 11-24 34 9.1 1800 11 42 482 5.96 26 7.2 12-20 90 6.73 1875 30 38 342 3.73 26 11.5 11-22 61 8.9 1880 15 43 472 4.01 25 7.2 15-22 66 6.82 1950 12 42 506 5.96 18 6.7 9-13 85 7.28 2125 32 42 420 5.37 24 9.9 9-18 56 6.5 Average 371.74 4.78 8.29 Total 13011 167.3

Table 2.2(b) Summary of structural parameters for forest plots (< 0.1 ha) in Ramal de Guaramacal, Andes, Venezuela.

Slope

Sector Alt. m Plot Nr. Plot area Num spp Total Num Ind/plot Average Num Ind/100 m2 Basal Area

Tree height Diameter Max. Med. Canopy Max. Med. 2474 33 300 29 175 58.3 0.97 12 6.1 6-10 27 6.8 North Guaramacal 2810 38 200 21 407 203.5 0.91 8.3 4.5 3-6 13 4.7 2830 PL3 100 10 41 41.0 0.19 4.3 2.8 2-4 16 6.8 2870 36 500 21 172 34.4 1.71 12 5.8 5-10 41.5 7.7 3050 34 200 18 263 131.5 0.81 6.5 4.1 3-5 21 5.3 3050 44 100 12 131 131 0.26 6 3.5 3-4 17 4.1 2950 41 400 22 272 68.0 1.77 10 6.2 5-8 25 6.9 South Guaramacal 2950 40 200 19 277 138.5 1.1 6 3.6 3-5 27.5 5.6 3060 43 50 14 92 184 0.25 5 3.1 3-5 12.8 5.2 Forest classification

The interpretation of the TWINSPAN table, based on affinities of floristic composition and relative species abundance, allowed recognition of seven vegetation communities at association level, grouped in three alliances and one major group equivalent to order level (Table 2.3). Three subandean forest (LMRF) communities and four Andean - high Andean forest (UMRF-SARF) communities are distinguished. The classification and description of the forest vegetation communities of Ramal de Guaramacal are presented below.

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Table 2.3. Phytosociological table of montane forests of Ramal de Guaramacal, Venezuela. 1: Simiro erythroxylonis - Quararibeet

um magnificae; 1.1: typicum; 1.2: bunchosietosum armeniaceae. 2: Conchocarpo larensis -

retum moritzianae. 3: Croizatio brevipetiolatae - Wettinietum praemorsae; 3.1: hedyosmetosum cuatrecazanum; 3.2: var.

Protium tovarense

. 4: Schefflero ferrugineae - Cybianthetum laurifolii; 4.1: typicum; 4.2: miconietosum

suaveolentis. 5: Geissantho andini - Miconietum jahnii; 5.1: Subcommunity of

Freziera

serrata

. 6: Libanothamnetum griffinii. 7: Gaultherio anastomosantis - Hesperomeletum obtusifoliae.

Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23 27 16 33 6 9 24 8 39 35 36 37 38 43 44 PL3 34 40 41 Area 1x10 (m 2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 30 100 100 100 100 100 100 40 100 20 5 10 10 20 20 40 No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35 61 33 29 31 29 34 31 41 27 21 18 17 9 10 10 18 19 22 E 1 1 111 2 11 1 1 2 2 2 11 1 2 2 2 11 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 33 2 3 2 2 l 434 8 8 1 8 6 6 9 1 3 1 9 9 901 1 8 8 74 43 2 2 54 4 1 53 7 8 8 8 8 00 8 09 9 e 535 5 8 0 05 0 0 0 0 7 6 5 570 2 8 77 5 8 55 6 8 77 0 8 0 5 9 7 7 165 3 5 5 5 v 0 0 000 0 0 0 0 0 0 0 00 0 0 0 0 5 0 5 0 0 0 0 000 40 0 0 00 00 0 0 0 0 0 0 0 0 Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO N N N S S S S N N N N N S N N N S S Park sector B B B A CC B B B B A A A A A B A BBB B B A A A CB A B A A A A A A A A A A A A A A A Order Alliance Association 34 5 6 7 Subassociation 1.1 4.1 5.1 Variant 3.2

Subandean forests (LMRF) Assoc. 1. Simiro erythroxylonis - Quararibeetum magnificae Simira er

ythrox ylon 1 5 3 1 3 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Parathesis venezuelana 1 3 1 2 1 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Urera caracasana 2 1 1 4 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Quararibea ma gnifica . 4 3 . 4 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Cuatresia riparia . 1 3 . 3 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Picramnia sp. . 1 1 . 1 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ae giphila floribunda 1 . 1 2 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Diplazium hians 1 . .. 3 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . In ga edulis . 1 . 1 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Vasconcella microcarpa . 1 . 1 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ocotea cernua 1 . .. 1 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Huertea glandulosa 1 . . 1 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Subassoc. 1.1. typicum Aphelandra macroph ylla 3 1 2 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ps ychotria trichotoma 3 2 1 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Alchornea glandulosa 1 1 1 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Cecropia sararensis 3 . 1 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ocotea sp. C . 3 2 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . St ylo gy ne lon gifolia . 2 3 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Hippotis albiflora 2 . 2 .. . . 1 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Matisia sp. . 1 1 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Nectandra aff. membranacea 1 1 ... . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . 1.2 4.2 3.1 Meliosma tachirensis - Alchornea grandiflora

montane forest order group

Geonomo undatae -Posoq. coriaceae

Farameo killipii - Prunion moritzianae

12

Ruilopezio paltonioides - Cybianthion marginati

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Trichilia pallida . 1 1 .. . . . 1 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Zyg ia bisin gula 2 . ... . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ficus sp. 2 . ... . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Paullinia capreolata 2 . ... . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Tammsia anomala . 2 ... . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Trichilia hirta . 2 ... . . . . . . 1 .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Piper hispidum . . 2 .. . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . .

Subassoc. 1.2. bunchosietosum armeniaceae Acal

ypha macrostach ya . . . 4 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ps ychotria fortuita . . . 4 1 2. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Bunchosia armeniaca . . . 2 1 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Pleuroth yrium costanense . . 1 1 4 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ficus tonduzii . . 1 1 1 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Saurauia tomentosa . . . 3 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Diplazium celtidifolium . . .. 2 . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Piper s p. (Liana ) . . .. 2 2. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . H ydran

gea aff. peruviana

. . . 2 . . . . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Cestrum bi gibbosum . . .. 1 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Solanum nudum . . .. 1 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Coussarea moritziana . . ... 2 1 1 3 3 . . . 1 . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Conchocarpus larensis 4 . ... . 5 1 5 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Alsophila erinacea . . ... 1. 4 . 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Sloanea guianensis . . .. 1 1 1 1 1 1 . . . 3 . . 1 . . . . 1 . . . ... .. . . .. .. . . . . . . . . Miconia lonchoph ylla . . ... 1 1 1 1 3 . 1 .. . 1 . 2 . 1 . . . . 1 ... .. . . .. .. . . . . . . . . Meliosma pittieriana . . ... . 1 1 1 . . . 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . C yathea kalbre yeri . . ... 1. 2 1 . . . .. . . . . 1 . . 3 . . . ... .. . . .. .. . . . . . . . . Eschweilera perumbonata . . ... . 1 . 1 2 . . .. . 1 . . . . 1 1 . . . ... .. . . .. .. . . . . . . . . Chr ysoph yllum cf. cainito 1 . ... . 1 . 2 1 . . . 2 . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Sloanea rufa . . .. 1 1. . 1 1 . . . 1 . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Mouriri barinensis 1 . ... 2. . 1 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Asplundia va gans . . ... . . 3 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Pseudolmedia ri gida . . ... . . 1 1 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Dussia coriacea . . .. 1 1 1 . . . 1 . 1 . . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Eu genia grandiflora . . ... . 1 . 1 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Eu genia sp. 1 . . ... . 1 . . 1 . . .. . . . 1 . . . . . . . ... .. . . .. .. . . . . . . . . In ga aff. densiflora . . ... 1 1 . . . . . .. . . . . . . . 1 . . . ... .. . . .. .. . . . . . . . . Machaerium cf. floribundum . . ... . . 1 . 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Picramnia sp. A 1 . ... . . 1 . 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ocotea rubrinervis . . ... . 1 . 1 . . . .. . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . St ylo gy ne sp. A . . ... 1. . . 1 . . .. . . . . 1 . . . . . . ... .. . . .. .. . . . . . . . . Eu genia sp. 3 . . .. 1 3. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Chionanthus sp. . . ... 2. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Toco yena costanensis . . ... . . 2 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . .

Assoc. 2. Conchocarpo larensis - Coussaretum moritzianae

The forest vegetation of Ramal de Guaramacal

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Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23 27 16 33 6 9 24 8 39 35 36 37 38 43 44 PL3 34 40 41 Area 1x10 (m 2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 30 100 100 100 100 100 100 40 100 20 5 10 10 20 20 40 No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35 61 33 29 31 29 34 31 41 27 21 18 17 9 10 10 18 19 22 E 1 1 111 2 11 1 1 2 2 2 11 1 2 2 2 11 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 33 2 3 2 2 l 434 8 8 1 8 6 6 9 1 3 1 9 9 901 1 8 8 74 43 2 2 54 4 1 53 7 8 8 8 8 00 8 09 9 e 535 5 8 0 05 0 0 0 0 7 6 5 570 2 8 77 5 8 55 6 8 77 0 8 0 5 9 7 7 165 3 5 5 5 v 0 0 000 0 0 0 0 0 0 0 00 0 0 0 0 5 0 5 0 0 0 0 000 40 0 0 00 00 0 0 0 0 0 0 0 0 Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO N N N S S S S N N N N N S N N N S S Park sector B B B A CC B B B B A A A A A B A BBB B B A A A CB A B A A A A A A A A A A A A A A A Order Alliance Association 34 5 6 7 Subassociation 1.1 4.1 5.1 Variant 3.2 1.2 4.2 3.1 Meliosma tachirensis - Alchornea grandiflora

montane forest order group

Geonomo undatae -Posoq. coriaceae

Farameo killipii - Prunion moritzianae

12

Ruilopezio paltonioides - Cybianthion marginati

Petrea pubescens . . ... . . . 2 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Piper arboreum . . ... . 2 . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Eu genia sp. . . ... . . 2 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Salacia aff. cordata . . ... . . 1 . 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Clusia sp. 1 . . ... . . . . 1 . . .. . . . 1 . . . . . . . ... .. . . .. .. . . . . . . . . Cecropia telenitida . . . 2 . . . 1 . . . . .. 1 . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Eu genia sp. 2 . . ... 1. . . . . . .. . . . . . 1 . . . . . ... .. . . .. .. . . . . . . . . Prunus cf. skutchii . . ... . 1 . . . . . . 1 . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Simira lezamae . . ... . . . . 1 . . .. . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . .

Alliance 1. Geonomo undatae - Posoquerion coriaceae Guarea kunthiana

1 1 3 2 2 2. 1 1 1 . 1 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Pouteria baehniana 5 . .. 1 1 2 1 2 2 . . . 2 . . 1 . . 1 . . . . . ... .. . . .. .. . . . . . . . . Geonoma undata 1 1 4 . 1 . 1 4 1 . . . .. . . . . . . . 1 . . . ... .. . . .. .. . . . . . . . . Calatola venezuelana . . 1 . 3 5 1 2 . 1 1 . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Posoqueria coriacea 1 . 2 1 2 2. 2 1 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Rud gea nebulicola . . 1 . 2 . . 2 3 2 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Mata yba camptoneura 1 1 1 . 1 . 1 1 1 1 . . .. . . 1 . . . 1 2 . . . ... .. . . .. .. . . . . . . . . Mabea occidentalis 3 . ... . . 2 3 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Persea peruviana . . 1 . 1 1 1 1 . 1 1 . 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Eu genia moritziana . 1 1 2 . 2. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Marc gravia brownei . . 1 . 1 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Tapirira guianensis 1 . ... . . . 1 1 . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Gordonia fruticosa . . . 1 . . 1 . 1 . . 1 .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ps ychotria lon girostris . . . 1 . . . 1 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Rollinia mucosa . . 1 .. . . 1 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Tri gy naea duckei . 1 ... . . . 1 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . H ydran gea cf. preslii . 1 ... 1. . . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Ficus n ymphaeifolia . 1 ... . . 1 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Nectandra aff. purpurea . 1 ... . . . 1 . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . . 22

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Paullinia cf. latifolia . . 1 .. . . 1 . . . . .. . . . . . . . . . . . ... .. . . .. .. . . . . . . . .

Assoc. 3. Croizatio brevipetiolatae - Wettinietum praemorsae Croizatia brevipetiolata

. . ... . 5 . . 1 5 5 5 2 . 4 4 5 2 . . 1 . . . ... .. . . .. .. . . . . . . . . Wettinia praemorsa . . ... . 2 3 2 4 2 . 2 5 5 5 4 5 5 2 1 1 . . . . 3 . .. 3 . .. .. . . . . . . . . Meriania grandidens . . ... 1. . . . 1 2 1 . 1 1 2 1 1 . . . . . . ... .. . . .. .. . . . . . . . . Aniba cf. cinnamomiflora . . ... . 2 1 1 3 1 . . 3 . 1 2 . . 2 2 2 . . . ... .. . . .. .. . . . . . . . . C ybianthus cuspidatus . . ... . . . 3 . . 1 . 2 . 1 . . . 3 . . . . . ... .. . . .. .. . . . . . . . . Miconia cf. minutiflora . . ... . 1 . . . 1 . 1 1 . 1 . . . . 1 1 . . . ... .. . . .. .. . . . . . . . . Elaea gia ruiz-teranii . . . . . 1 . . . . 1 4 . 1 . 1 1 . 1 . . . . 1 . . 1 .. . . .. .. . . . . . . . .. Ocotea sp. A . . ... . . . . 1 . 1 .. 1 1 . . . 1 . . . . . ... .. . . .. .. . . . . . . . . Miconia lucida . . ... . . . . 1 . . . 2 . 1 1 1 2 1 1 . . . . . 1 . .. . . .. .. . . . . . . . . Hed yosmum cf. gentr yi . . ... . 1 1 1 1 . . .. . . 1 1 . 1 2 2 . . . ... .. . . .. .. . . . . . . . . Faramea guaramacalensis . . ... . 1 . . . . . .. 1 1 . 1 1 . . 1 . . . ... .. . . .. .. . . . . . . . . Ma ytenus sp. A . . ... . 1 . . . . . .. 1 1 . . 1 . . . . . . ... .. . . .. .. . . . . . . . . Ocotea aff. puberula . . ... . . . . . . . . 2 . . 2 . . . . . . . . ... .. . . .. .. . . . . . . . . M yrcia acuminata . . ... . 1 . . . . . .. . 2 . . 1 . 1 2 . . . ... .. . . .. .. . . . . . . . .

3.2. var. Protium tovarense Protium tovarense

. . ... . 2 1 1 2 . . .. 2 1 . . . 4 5 4 . . . ... .. . . .. .. . . . . . . . . Coccoloba cf. llewel ynii . . ... . . . . . . . .. . 1 . 2 1 2 1 2 . . . . 1 . .. . . .. .. . . . . . . . . Aiphanes ster giosii . . ... . . . . . . . .. . . . . 1 2 1 2 . . . ... .. . . .. .. . . . . . . . . Persea meridensis . . ... . . . . . . . . 1 . . 1 1 . 1 2 2 . . 1 ... 1 . . . .. .. . . . . . . . . Miconia sp. B . . ... . . . . . . . .. . . . . 2 . . . . . . ... .. . . .. .. . . . . . . . . M yrcia sp.1 . . ... . . . . . . . .. . . . . . 2 . 1 . . . ... .. . . .. .. . . . . . . . .

Subassoc. 3.1. hedyosmetosum cuatrecazanum Hedyosmum cuatrecazanum

. . ... . . . . . 2 2 3 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Palicourea demissa . . . 1 . . . . . . 2 3 2 . . . . . . . . . 3 . . ... .. . . .. .. . . . . . . . . Sapium st ylare . . ... . . . . . 2 1 2 . . . . . . . . . . . . ... .. 2 . .. .. . . . . . . . . Ae giphila ternifolia . . . 1 . . . . . 1 1 1 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Casearia tachirensis . . .. 2 3. . . . 1 1 3 . . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Palicourea puberulenta . . ... . . . . . 2 3 . 2 . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Meriania macroph ylla . . .. 2 1. . . . 2 . 2 . . . 1 . . . . . . . . ... .. . . .. .. . . . . . . . . Perrottetia quinduensis . . ... . . . . . 1 . 3 . . . . . . . . . . . . ... .. 1 . .. .. . . . . . . . . Guettarda crispiflora . . ... 1. . . . 1 . 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Turpinia occidentalis . . ... 1. . . . 1 . 1 . . . . . . . . . . . . ... .. . . .. .. . . . . . . . . Cestrum darc yanum . . ... . . . . . . 1 1 . . . . . . . . . . . . ... .. . . . 1 .. . . . . . . . . Miconia amilcariana . . . 1 . . . . . . 1 .. . .. . . . . . . .. . . . . . . .. 1 . . ... . . . . . .

Andean Forest (UMRF) Assoc. 4. Schefflero ferrugineae - Cybianthetum laurifolii Cybianthus laurifolius

. . . . . . . . . . . . . . . . . . . . . . 2 1 1 424 41 . 3 21 . . . . . . . . . . M yrsine coriacea . . . . . . . . . . . . . 1 . . . . . . . . 1 2 1 111 11 . 1 21 . . . . . . . . . . Schefflera ferru ginea . . . . . . . . . .. . . . . . . ... . . 1 2 1 1 1 1 2 . . 2 1 .. . . . . . . . .. Hed yosmum crenatum . . ... . . . . . . . .. . . . . . . . . 3 4 3 2 2 2 .. . 3 .. .. . . . . . . . . Miconia ulmarioides . . . . . . . . . .. 1 . . . . . ... . 2 3 2 3 . 2 . 2 1 . . 1 .. . . . . . . . .. Ilex laurina . . . . . . . . . . . . . .. . . . 1 . 1 . 1 1 . 1 1 2 1. 1 1 1. . . . ... . . . .

The forest vegetation of Ramal de Guaramacal

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Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23 27 16 33 6 9 24 8 39 35 36 37 38 43 44 PL3 34 40 41 Area 1x10 (m 2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 30 100 100 100 100 100 100 40 100 20 5 10 10 20 20 40 No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35 61 33 29 31 29 34 31 41 27 21 18 17 9 10 10 18 19 22 E 1 1 111 2 11 1 1 2 2 2 11 1 2 2 2 11 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 33 2 3 2 2 l 434 8 8 1 8 6 6 9 1 3 1 9 9 901 1 8 8 74 43 2 2 54 4 1 53 7 8 8 8 8 00 8 09 9 e 535 5 8 0 05 0 0 0 0 7 6 5 570 2 8 77 5 8 55 6 8 77 0 8 0 5 9 7 7 165 3 5 5 5 v 0 0 000 0 0 0 0 0 0 0 00 0 0 0 0 5 0 5 0 0 0 0 000 40 0 0 00 00 0 0 0 0 0 0 0 0 Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO N N N S S S S N N N N N S N N N S S Park sector B B B A CC B B B B A A A A A B A BBB B B A A A CB A B A A A A A A A A A A A A A A A Order Alliance Association 34 5 6 7 Subassociation 1.1 4.1 5.1 Variant 3.2 1.2 4.2 3.1 Meliosma tachirensis - Alchornea grandiflora

montane forest order group

Geonomo undatae -Posoq. coriaceae

Farameo killipii - Prunion moritzianae

12

Ruilopezio paltonioides - Cybianthion marginati

Drim ys granadensis . . . . . . . . . .. . . . . . . ... . . 1 1 1 1 . 1 . . . 1 . 1 1 2 1 . . . . . .. Prestoea acuminata . . . . . . . . . .. . . . 3 . . ... . . . . . 1 1 . 2 1 1 . 1 .. . . . . . . . .. Palicourea apicata . . ... 1. . . 3 . . . . . . . 1 . 2 . 2 . 3 4 3 2 3 . 1 . . .. .. . . . . . . . . Weinmannia glabra . . . . . . . . . .. . . . . . . ... 1 2 1 2 1 . 2 3 3 . . .. .. . . . . . . . .. Rhipidocladum geminatum . . . . . . . . . .. . . . . . . ... . . . 4 . . . 3 . . . 2 . 2 . . . . . . . . .. Podocarpus oleifolius . .. . . . .. . . . . . . . ... 1 .. . 1 1 .. 1 2 2 . . . . 2 . . . . . . . . . . Ilex m yricoides . . . . . . . . . .. . . . . . . ... . . 1 . . . . 2 3 . . .. 1 . . . . . . . . .. Persea aff. mutisii . . . . . . . . . .. . . . . . . ... . . . 1 . . . 2 1 . . 1 . 1 . . . . . . . . .. Weinmannia fa garoides . . ... . . . . . . . .. . . . . . . . . . . . ... 2 . . 4 . 1 .. . . . 1 . . . . By rsonima sp. . . . . . . . . . . . .. . .. . . 1 . . . .. . 1 1 . 1 . .. . . . ... . . . . . . Ocotea jelski . . . . . . . . . .. . . . . . . ... . . 1 . . . . 1 1 . . .. .. . . . . . . . .. Ilex sp.2 . . ... . . . . . . . .. . . . . . . . . . . . 1 .. .. . 1 1 . .. . . . . . . . . Persea sp.1 . . ... . . . . . . . .. . . . . . . . . . . . 1 . 2 .. . . .. .. . . . . . . . . By rsonima karstenii . . ... . . . . . . . .. . . . . . . . . 1 . . ... 2 . . . .. .. . . . . . . . . Viburnum tinoides . . ... . . . . . . . .. . . . . . . . . . 1 . ... . 1 . . .. .. . . . . . . . . Arthrost ylidium venezuelae . . ... . . . . . . . .. . . . . . . . . 1 . . ... .. 1 . .. .. . . . . . . . . Palicourea jahnii . . ... . . . . . . . .. . . . . . . . . . . . ... 1 . . . .. .. . 1 . . . . . . Pentacalia vicelliptica . . ... . . . . . . . .. . . . . . . . . . . 1 ... .. . . .. 1 . . . . . . . . . Subassoc. 4.1. typicum Cal yptranthes cf. meridensis . . ... . . . 1 . . 1 .. . . . 2 2 . . . 3 2 1 3 1 2 .. . . .. .. . . . . . . . . Brunellia cf. inte grifolia . . ... . . . . . . . .. . . . . . 1 1 . 1 1 1 1 2 . .. . . .. .. . . . . . . . . Panopsis suaveolens . . ... . . . . . . . .. . . . . . . . . 1 1 1 ... .. . . .. .. . . . . . . . . M yrcia aff. guianensis . . ... . . . . . . . .. 1 . . . . . . . 1 1 . . 1 . .. . . .. .. . . . . . . . . Dioicodendron dioicum . . ... . . . . . . 1 .. . . . . . . . . 1 . 1 . 1 . .. . . .. .. . . . . . . . . Gaiadendron punctatum . . ... . . . . . . . .. . . . . . . . . 1 . 1 ... 1 . . . . 1 .. . . . . . . . . Ilex truxillensis var. bullatissima . . ... . . . . . . . .. . . . . . . . . 1 . . 1 1 . .. . . .. .. . . . . . . . . Meliosma venezuelensis . . ... . . . . . . . .. . . . . . . . . 1 . 1 ... .. . . 1 . .. . . . . . . . . Sy mplocos bo gotensis . . ... . . . . . . . .. . . . . . . . . 1 . 1 ... .. . . .. .. . . . . . . . . Ocotea sericea . . ... . . . . . . . .. . . . . . . . . . . 1 1 .. .. . . .. .. . . . . . . . . 24

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Ocotea sericea . . ... . . . . . . . .. . . . . . . . . . . 1 1 .. .. . . .. .. . . . . . . . .

Subassoc. 4.2. miconietosum suaveolentis Critoniopsis paradoxa

. . ... . . . . . . . .. 1 . . . 1 . . . 1 3 . ... . 3 4 2 4 . .. . . . . . . . . Hed yosmum sp. A . . ... . . . . . . . .. 2 . . . 3 . . . . . . ... . 1 2 2 4 . .. . . . . . . . . Miconia suaveolens . . ... . . . . . . . .. . . . . . . . . . . . ... . 3 1 3 .. .. . . . . . . . . H yeronima scabrida . . ... . . . . . . . .. . . . . . . . 1 . . . ... . 1 1 1 2 . .. . . . . . . . . Ae giphila moldenkeana . . ... . . . . . . . .. . . . . . . . . . . . ... . 2 1 . .. .. . . . . . . . . Chusquea purdieana . . ... . . . . . . . .. . . . . . . . . . . . ... . 3 . . .. .. . . . . . . . . Ocotea sp. B . . ... . . . . . . . .. . . . . . . . . . . . ... . 1 1 . .. .. . . . . . . . . Monnina meridensis . . ... . . . . . . . .. . . . . . . . . . . . ... . 2 . . .. .. . . . . . . . .

Alliance 2. Farameo killipii - Prunion moritzianae Faramea killipii

. . . . . . . . . . 1 2 . 1 1 2 1 2 1 . 1 . 4 3 2 1 4 4 1. . . 1. . . . ... . . . . Clethra fa gifolia . . . . . . . . . . . 1 . . 1 . . 11 . 1 . 1 2 1 1 . 11 5 2 22 21 . . 1 . . . . . . Anaectocal yx bracteosa . . ... . . . . . . . .. 1 1 . 1 1 1 1 1 3 3 2 3 3 2 1 1 1 1 .. .. . . . . . . . . C yathea pauciflora . .. . . . .. . . . . . . 1 .. 2 2 . 4 . 1 ... 2 1 2 . 3 .2 .. . . . . . . . .. Prunus moritziana . . ... . . . . . . 1 1 1 1 1 1 1 1 . 1 1 1 1 1 2 1 . . 1 . 1 .. .. . . . . . . . . Zanthox ylum melanostictum . . . . . . . . . .. . . 1 1 1 2 . 1 1 3 1 1 . 2 . 1 .. . 1 .. .. . . . . . . . .. C yathea caracasana 1 . ... . . . . . . . .. 2 . . . . 1 . . 3 3 . 2 1 . . 2 . . .. .. . . . . . . . . C ybianthus iteoides . . . . . . . . . .. . . 2 . 1 2 . 1 . 2 1 1 . . 2 1 .. . . .. .. . . . . . . . .. Eu genia cf. tamaensis . . . . . 2 . . . .. 3 . 1 . . . 1 . 1 . . 3 . 2 . 1 .. . . .. .. . . . . . . . .. Weinmannia aff. balbisiana . . ... . . . . . . . .. 1 1 . 1 1 1 . . . . 2 2 .. .. . 1 .. .. . . . . . . . . Rud gea ta yloriae . .. . . . .. . 1 . . . . 1 .. 2 2 .. .. ... 2 . 1 . 1 . 1 . . . . . . . . . . . Aiouea dubia . . . . . 1 . . . .. 1 . 1 . . 2 . 1 . . . 1 1 1 . 1 .. . . .. .. . . . . . . . .. Miconia mesmeana . .. . . . .. . . . 1 . . . ... 1 .. .. ... . 1 . . 4 . . 1 . . . . . . . . . . Miconia tovarensis . . ... . . . . . . . . 2 2 . . . . . . 1 . . . ... .. 3 . 2 . .. . . . . . . . . M yrcia cf. sanisidrensis . . . . . . . . . .. . . . 1 1 . 1 . 2 . . . . . 3 2 .. . . .. .. . . . . . . . .. Ocotea va ginans . . ... . . . . . . 2 1 . 1 . 1 . 1 . . . . . 2 ... .. . 1 .. .. . . . . . . . . Hieron yma moritziana . . ... . . 1 . . . 1 1 1 1 . 1 . . . . 1 1 . 1 ... .. . . 1 . .. . . . . . . . . Sy mbolanthus vasculosus . . . . . . . . . .. . . . . 1 . .. 1 1 . 1 . 1 2 . .. . . . 1 .. . . . . . . . .. cf. Elaeoluma nuda . . ... . . . . . 1 1 1 . . 1 . . . . 1 . 1 . 1 ... .. . . .. .. . . . . . . . . Ocotea leucox ylon . . . . . . . . . .. . . . . 1 . 1 . 1 . . 1 . 1 . 1 .. . . .. .. . . . . . . . .. Mikania banisteriae . .. . . . .. . . 1 . . . . . 2 1 . .. .. ... 1 . . . . . . . . . . . . . . . . . Citronella costaricensis . . ... . . . . . . 1 1 . . . . . . . . . 1 1 . ... .. . . .. .. . . . . . . . . Geonoma jussieuana . . ... . . . . . . . .. . 1 . . . 1 . . . . 1 ... .. . . 1 . .. . . . . . . . . Dio genesia tetrandra . . ... . . . . . . . .. . . . . . . 1 . . . 1 ... .. . 2 .. .. . . . . . . . . Ocotea cf. hexanthera . .. . . . .. . . . . . . . .. 2 . .. .. ... 2 . . . . . . . . . . . . . . . . . Saurauia yasicae . . ... . . . . . . 1 .. 1 . . . . . . . 1 . . ... .. . . .. .. . . . . . . . . Dicksonia sellowiana . . ... . . . . . . . .. . . . . . 1 . . . 1 . . 1 . .. . . .. .. . . . . . . . . Mikania ni gropunctulata . . ... . . . . . . . .. . . . 1 . . . . . . 1 ... .. . . .. .. . . . . . . . . Eu genia albida . . ... . . . . . . . .. . . 1 . . . . . . . . .. 1 .. . . .. .. . . . . . . . .

The forest vegetation of Ramal de Guaramacal

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Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23 27 16 33 6 9 24 8 39 35 36 37 38 43 44 PL3 34 40 41 Area 1x10 (m 2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 30 100 100 100 100 100 100 40 100 20 5 10 10 20 20 40 No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35 61 33 29 31 29 34 31 41 27 21 18 17 9 10 10 18 19 22 E 1 1 111 2 11 1 1 2 2 2 11 1 2 2 2 11 1 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 2 33 2 3 2 2 l 434 8 8 1 8 6 6 9 1 3 1 9 9 901 1 8 8 74 43 2 2 54 4 1 53 7 8 8 8 8 00 8 09 9 e 535 5 8 0 05 0 0 0 0 7 6 5 570 2 8 77 5 8 55 6 8 77 0 8 0 5 9 7 7 165 3 5 5 5 v 0 0 000 0 0 0 0 0 0 0 00 0 0 0 0 5 0 5 0 0 0 0 000 40 0 0 00 00 0 0 0 0 0 0 0 0 Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO N N N S S S S N N N N N S N N N S S Park sector B B B A CC B B B B A A A A A B A BBB B B A A A CB A B A A A A A A A A A A A A A A A Order Alliance Association 34 5 6 7 Subassociation 1.1 4.1 5.1 Variant 3.2 1.2 4.2 3.1 Meliosma tachirensis - Alchornea grandiflora

montane forest order group

Geonomo undatae -Posoq. coriaceae

Farameo killipii - Prunion moritzianae

12

Ruilopezio paltonioides - Cybianthion marginati

Montane forest order group of

Meliosma tachirensis and Alchornea grandiflora Clusia trochiformis . 1 . . . 1 . . . . . 1 . .1 1 . . 2 . 2 . 2 2 3 323 42 4 3 3 . . . . . . . . . . . Alchornea grandiflora . 1 . 2 1 2 1 2 1 1 1 1 1 3 1 1 2 1 . 1 3 2 1 . 3 1 1 . .. 2 . .. .. . . . . . . . . Piper lon gispicum . . . 1 1 3 1 3 1 2 3 3 3 2 . 1 2 1 2 1 . 1 1 . 1 ... .. . . .. .. . . . . . . . . Clusia alata . . . 1 . . 1 . 1 . . . 1 . . 1 1 . . 1 . 1 . 2 . 211 23 2 4 31 . . . . . . . . . . Miconia theaezans . .. 1 . . 1 1 2 1 2 1 2 . 2 ... 1 .. . 1 ... 1 1 . 1 3 1 3 . . . . . . . . . . . Rua gea pubescens . 2 2 .. . . 1 . . 2 3 1 . 2 1 1 2 3 1 . 1 1 . . ... .. . . 2 . .. . . . . . . . . Geissanthus fra grans . . 2 2 . 1 1 . 1 1 2 1 1 1 2 1 1 .. 1 . 1 . . 1 1 . .. . 1 .. .. . . . . . . . .. Besleria pendula . . 1 .. . 1 1 1 1 . . . 3 . . 2 1 2 3 2 2 . 1 . ... .. . . .. .. . . . . . . . . C yathea fulva . . ... . . . . 2 . 2 1 . . 3 2 1 . 1 2 3 . . 1 . 1 . .. . 1 . 1 1 . . . . . . . . . Beilschmiedia tovarensis . . ... 1 1 . . 1 1 2 1 1 1 2 1 1 . 1 1 1 . . 2 . 1 . .. . . .. .. . . . . . . . . Billia rosea 1 . .. 1 1 1 . . 1 . 1 . 1 1 1 1 1 1 1 1 1 1 . 1 1 1 . .. . . .. .. . . . . . . . . Hieron yma cf. oblon ga . . ... 1 1 . 1 1 . 1 1 1 2 1 2 1 2 . . . 1 . 1 ... .. 1 . .. .. . . . . . . . . Psammisia hookeriana . . 1 .. . . . . 1 1 . .. . 1 . 1 . 1 1 . 1 . 2 2 .. .. . 1 .. .. . . . . . . . 1 Blakea schlimii . . 1 .. 1 1 1 . 2 . . 1 2 1 1 1 1 . 1 1 . . . . ... .. . . .. .. . . . . . . . . Meliosma tachirensis . . ... 2. . . 1 1 1 1 . . . . 1 . . 1 2 1 . 1 ... . 1 . . 1 . .. . . . . . . . . Sphaeradenia laucheana . . 1 . 1 . . . 1 . . .. . .. . . . 5 1 . .. . 1 2 . 3 . .. . . . ... . . . . . . Tetrorchidium rubrivenium 1 . . 1 1 . . 2 . . 1 1 1 1 . . 1 . 1 . . . . . 1 ... .. 1 . .. .. . . . . . . . . Palicourea an gustifolia . . ... 1. 1 . 1 . . . 3 . . 3 . . . . 1 . . . ... . 1 . . .. .. . . . . . . . . Dendropanax arboreus 1 . ... . . 1 2 1 . . . 1 . . 1 . . 1 . 2 . . . ... .. . . .. .. . . . . . . . . Ocotea karsteniana . . . 1 . . . . . .. . . 1 1 1 1 ... . . 1 1 1 . 1 .. . . .. .. . . . . . . . .. Eu genia cf. oerstediana 1 . ... . . 1 1 2 . . .. . . . . . . 2 . . . . ... .. 2 . .. .. . . . . . . . . Miconia cf. dolichopoda 1 1 1 1 . . . . . 1 . . 1 . . . . . . . . . .. 1. 1 . . . .. . . . ... . . . . . . Trichilia septentrionalis . . .. 1 1. . . . 1 1 1 . . . . . . . . . . 1 . ... .. . 1 .. .. . . . . . . . . Ocotea floribunda . . ... . . . 1 . 1 2 .. . . . 1 . . . . 2 . 1 ... .. . . .. .. . . . . . . . . Weinmannia sorbifolia . . . 1 . . . . . .. . . . . . . ... 2 2 . . . ... .. . . .. .. . . . . . . . . Ocotea macropoda . . ... . . 1 . 1 . . .. . . . . . 1 . . 1 . 1 1 . 1 .. . . .. .. . . . . . . . . Elaea gia karstenii . . ... . 1 . . . . . .. . . . 2 . . . . . . 2 ... .. . . .. .. . . . . . . . . Tabebuia gua yacan 1 . ... . . 1 1 . . . .. . . . . . . . . . . 1 ... .. . . .. .. . . . . . . . . 26

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