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 3
The páramo vegetation of Ramal de Guaramacal, Trujillo,
Venezuela.
1. Zonal communities
Nidia L. Cuello A. and Antoine M. Cleef PHYTOCOENOLOGIA, 39 (3), 295–329. 20093.1 INTRODUCTION
Andean páramos play an essential role in the evolution and the ecology of the Andes (Vuilleumier & Monasterio 1986; Luteyn 1999; Hofstede et al. 2003; Hooghiemstra et al. 2006) and represent strategic ecosystems due to the environmental services they offer in the regional hydrological balance and agricultural production (Molinillo & Monasterio 1997, 2002; Monasterio & Molinillo 2003; Hofstede et al. 2003). Andean páramos are also, however, highly fragile ecosystems as a function of mounting demographic pressures, the expansion of agricultural and mining activities and of global warming, all of which represent major threats to the maintenance of environmental services and for the conservation of Andean biodiversity (Hofstede 2002; Van der Hammen 2002; Llambiet al. 2005).
Since the publication of the 'Flora de los Páramos de Venezuela' by Vareschi (1970), a substantial number amount of studies in but a few Venezuelan páramos has been published. The ecological studies by M. Monasterio and (own staff/foreign) collaborators (Monasterio 1980a; Sarmiento et al. 2003) were developed primarily in the central core of dry páramos in the state of Mérida. They remain ongoing in these páramos with highest altitude and most extension of the Cordillera of Mérida. At present, a great number of studies by researchers from the ICAE-ULA-Mérida, are available (see Sarmiento 2006 CD-ROM). These studies are mostly concerned with ecophysiology and functional processes in both natural and agro-ecosystems of the páramo and as such, remain unique in that there are not similar groups of this magnitude and focus elsewhere in the tropical Andes and high mountains of Central America and Mexico.
Despite a great environmental variability throughout a number of páramo areas and their associated vegetation communities along of the Cordillera de Mérida (Monasterio & Reyes 1980; Monasterio 1980b; Luteyn, 1999), little is currently known about páramo vegetation communities and their flora in other sectors of the Venezuelan Andes beyond the borders of Mérida state. To date, local floristic listings have appeared that include páramo areas such as those from Táchira and Trujillo states (Bono 1996; Dorr et al. 2000), there is a list of flowering plants of Venezuelan páramos (Briceño & Morillo 2002, 2006) and phytogeographical analyses of the páramo flora (Ricardi et al. 1997, 2000). Studies of classification and characterization of the vegetation communities in páramos of the Venezuelan Andes are limited to the descriptions of different sectors of Sierra Nevada de Mérida (Vareschi 1953, 1956; Baruch 1984; Berg 1998; Berg & Suchi 2000; Yánez 1998) and, as outlined above, to a general descriptive account for the whole region (Monasterio 1980b), floristic lists with comments on vegetation communities of páramos of Táchira state (Bono 1996) and a brief description of a selected area of Páramo Cendé in Trujillo state (Niño et al. 1997). In comparison, a much larger body of literature on plant diversity and vegetation exists for Colombian páramos (Cuatrecasas 1934, 1958; Cleef 1981; Sturm & Rangel 1985; Van der Hammen et al. 1983, 1984, 2003, 2005, 2008; Rangel 2000a, among others). Luteyn (1999) and Rangel (2000a) provide a summary of the flora and vegetation studies conducted throughout the last century in Colombian páramos.
Previous studies divided the north Andean páramo vegetation into several zones related to altitude (for a complete review we refer to Luteyn 1999). The Cuatrecasas (1934, 1958) altitudinal classification of superpáramo, páramo and subpáramo has since been widely adopted (Cleef1981;Acosta-Solís1984;Ramsay 1992; Jørgensen & Ulloa 1994; Hooghiemstra et al. 2006). For Venezuelan páramos, Monasterio (1980b) recognises two altitudinal zones called „pisos altitudinales‟: a High Andean zone or „Piso Altiandino‟ (4000-4800 m) and the Upper Andean zone or „Piso Andino Superior‟ (2800-4000 m) with a total of seven vegetation formation types and thirty four vegetation communities or “associations”. There are three vegetation types from the „Piso Altiandino‟, called 1) the High Andean Desert Páramo or „Páramo Desértico Altiandino‟, 2) the High Andean Periglacial Desert or „Desierto Periglacial Altiandino‟ and 3) the High Andean Forest of Polylepis sericea. Many authors agreed that the „Piso Altiandino‟ and the Superpáramo represent equivalent vegetation zones (Berg 1998; Luteyn 1999; Berg & Suchi 2000). In the „Piso Andino‟ zone, the four vegetation types recognized are 4) the Andean Páramo or „Páramo Andino‟, which includes heterogeneous páramo vegetation associations dominated either by rosettes or shrubs; 5) the Andean Grass Páramo or „Pajonal Paramero Andino‟, including páramo vegetation associations with high cover of tussock grasses; 6) the Andean Pasture Páramo or „Pastizal Paramero Andino‟, which is represented by vegetation associations with high cover of other non-tussock grasses; and 7) the Andean Páramo Forest or „Bosque Paramero Andino‟ (Monasterio 1980b). The wet páramo of Guaramacal found on the high summits of Ramal de Guaramacal (Fig. 1), has previously been reported as an important center of diversification of the genus Ruilopezia of the Espeletiinae (Cuatrecasas 1986). Moreover, due to its relative isolation, Ramal de Guaramacal is also an area with an endemic flora (Steyermark 1979; Ortega et al. 1987; Dorr et al. 2000). An important number of new and endemic species have been described from the forests and páramos of Guaramacal (Morillo 1988; Axelius & D' Arcy 1993; Carnevali & Ramírez 1998; Aymard et al. 1999; Benítez & Sawyer 1999; Taylor 2002; Stančik 2004; Stergios & Dorr 2003; Niño et al. 2005; Cuello & Aymard 2008). Endemic species of the Guaramacal subpáramo - páramo flora include:
Elaphoglossum appressum Mickel, Epidendrum guaramacalense Hágsater, Festuca guaramacalana Stančik, Ilex guaramacalensis Cuello & Aymard, Libanothamnus griffinii (Ruiz-Terán & López-Fig.) Cuatrec., Miconia aymardii
Wurdack, M. elvirae Wurdack, Rhynchospora guaramacalensis Strong and
Ruilopezia lopez-palacii (Ruiz-Terán & López-Fig.) Cuatrec., among others.
The zonal vegetation of the Páramo of Guaramacal is generally characterized by a mosaic of subpáramo formations (shrub páramo, bunchgrass páramo, most common bamboo páramo), intermingled with patches of dwarf forests. The páramo vegetation is distributed between 2800 and 3130 m. Due to its low altitude, the Páramo of Guaramacal has been catalogued by some authors as a subpáramo (Cuatrecasas 1986; Luteyn 1999). For the purpose of this paper, subdivison of subpáramo and grasspáramo, each in a lower and higher subzone, we refer to Cleef (1980, 1981).
Zonal and azonal vegetation is defined sensu Walter (1979). Zonal vegetation corresponds to the present vegetation as a function of the actual regional macroclimate. Zonal vegetation occurs on zonal soils and represents the majority of vegetation within the study area. Azonal vegetation is dependent on the special substrate conditions, such as where stress by water or dryness is experienced. Azonal vegetation communities in concave terrain is represented by peat bogs, mires or aquatic vegetation in the Guaramacal bamboo páramo, were treated separately (Cuello & Cleef 2009c).
The primary goal of the present study is to identify, define and characterize the zonal vegetation of Páramo de Guaramacal, and to establish a syntaxonomic scheme based on analysis of physiognomy, floristic composition, ecological relations and the altitudinal distribution of the different vegetation communities also in comparison to bamboo páramos elsewhere.
This work was carried out within the wider framework of a project aiming to study the diversity of flora and vegetation of the Guaramacal National Park (Cuello 1999, 2000, 2002, 2004; Dorr et al. 2000). Classification of forest vegetation and azonal páramo communities in Ramal de Guaramacal are described separately in Chapter 2 and 4 (Cuello & Cleef2009a, c).
3.2 STUDY AREA
Zonal páramo communities of the summit of Ramal de Guaramacal have been studied between 2800-3100 m, in the surroundings of 'Las Antenas' area (9o 14‟
1.02” N; 70o 11‟ 6.47” W) and Páramo El Pumar (9o 12‟ 45.6” N; 70o 12‟ 5.55”
W), 2.5 km Southwest of 'Las Antenas'. Ramal de Guaramacal is an outlier of the Venezuelan Andes, located South from the town of Boconó, Trujillo state, approximately 120 km Northeast of Mérida, in the centre of the Sierra Nevada de Mérida (Fig. 3.1).
The climatic characteristics of high humidity with permanent fog favour the development of great ground cover of Sphagnum spp. characteristic of the zonal shrub páramo vegetation associations and border of forests. This condition is very common all over the páramo areas of Ramal de Guaramacal and is not considered here as an azonality. First climatic records from a Davis Pro 2 climate station installed near the summit of Guaramacal (3100 m) by the first author since December 2006 to December 2007 (monthly precipitation in mm and monthly temperature in Celsius), registered a total amount of yearly rainfall of at least 2995.4 mm (some data were lost during some days in the most rainy months of june and july 2007). Relative humidity is extraordinary high, with a mean humidity of 96.88% throughout the year. The lowest mean relative humidity was observed in the month of February with a value of 92.35%. Mean temperature is 8.6oC, the
lowest temperatures of 1.3oC are recorded in December and January and the
highest temperature of 18.6oC in March. Detailed data of the Davis Pro 2 climate
station are intended to be published in a forthcoming paper on the upper forest line (Cuello et al. in prep.). For a more complete description of the study area the reader is referred to Chapter 2 and Cuello (1999).
Figure 3.1. Location of study area in the Venezuelan Andes.
3.3 METHODS Field Sampling:
Fieldwork on the zonal páramo vegetation of the Guaramacal range was conducted over a short altitudinal gradient between 2800 and 3100 m. Observations, general collections and quantitative sampling using line-intercept methods (Barbour et al. 1987), were conducted here. Lines of 10 m were laid down at ca. 10 m altitudinal
intervals on patches of vegetation with an apparently homogenous structure and composition; however, on occasion, it happened that the line also crossed other vegetation type(s). To avoid this, each line was divided into two sections of 5 m, a perpendicular 5 m line was then situated close to the first 5 m of the line to complete the 10 m. In few cases, some of those 5 m line segments on mixed vegetation were later excluded for the analysis. The horizontal measurement of interception of every plant species (vascular plants and cryptogams) touching the line was performed. The measurement of height and location of the plant with respect to the line was also registered, and together with measurements of relief variation each 25 cm, were used for drawing of vegetation and land form profiles. For the delineation of relief a cord extended horizontally along the length of the line (tape measure) leveled with a bubble level, was used as a reference. Soil sampling with an auger from 15 cm depth were conducted at the centre of each 5 m line interval. Soil pH and conductivity were later determined in the laboratory. A total of fifty observations sites and a hundred 5 m line sections were surveyed. At each observation site, information on topography, exposition, slope, geographic position (UTM coordinates), altitude and floristic composition were recorded. Botanical vouchers of all recorded species, including those with doubt as to their identification, equally found beyond the lines of interception as within were collected. Photographs, where possible, were also taken. The collected botanical material was processed, identified and deposited at Herbario Universitario PORT of UNELLEZ. For vascular plants, the nomenclature follows that of Dorr et al. (2000). Duplicates of mosses and lichens were sent to Dr. D. Griffin III (FLAS) and Dr. H.J.M. Sipman (B), respectively, for their identification. Additional duplicates were also deposited in MER, VEN and US. The collection number referred to is that of the first author.
Processing and data analysis:
Data for each survey were stored and processed using Microsoft Excel. For each species in each line section of zonal vegetation surveyed, the sum of the intersection and a percentage value of cover and relative cover were calculated. Percentage cover for each species is equal to the total sum of intersection for the species, multiplied by 100, then divided by the length of the line. Relative cover for each species is equal to the total sum of intersection for the species in the line, multiplied by 100, then divided by the total sum of intersections of all species. The number of individuals, relative abundance and the frequency of a species, based on the number of appearances of the species throughout 1 m sections of the line, were also computed.
A data matrix containing the percentage of relative cover of 91 vascular species recorded for ninety one 5 m-line surveys was processed with TWINSPAN (Hill 1979) using program PC-Ord 4 (McCune&Mefford 1999). Vegetation data were then interpreted in terms of syntaxonomical classification, based on cover and floristic affinities, following the Zürich-Montpellier approach (Braun-Blanquet 1979) and the International Code of Phytosociological Nomenclature (Weber et al. 2000).
The diverse subunits, recognized in a progressive way by the TWINSPAN procedure, were hierarchized in associations, and higher (alliances, order) and lower syntaxa (subassociations and variants).
In order to explore relationships between the species composition of vegetation types and some of the environmental variables measured in this study (altitude, slope angle, soil and humus depth), an ordination analysis, using canonical correspondence analysis (CCA), also available in the PC-Ord package, was performed.
3.4 RESULTS
Zonal subpáramo plant communities
Interpretation of the TWINSPAN table allowed recognition of 5 vegetation communities at association level, grouped into two alliances and one order (Table 3.1). The zonal subpáramo plant communities recognized in Ramal de Guaramacal are summarized as follows:
A.RUILOPEZIOLOPEZ-PALACII–CHUSQUEETALIAANGUSTIFOLIAECuello & Cleef 2009
I.HYPERICO PARAMITANUM–HESPEROMELETION OBTUSIFOLIAE Cuello & Cleef
2009
1. Ruilopezio paltonioides –Neurolepidetum glomeratae Cuello & Cleef 2009 1.1. variant of Disterigma alaternoides
1.2 variant of Ugni myricoides
2. Disterigmo acuminatum –Arcytophylletum nitidum Cuello & Cleef 2009 2.1. pentacalietosum cachacoensis Cuello & Cleef 2009
2. 2. subassociation typicum Cuello & Cleef 2009
II.HYPERICO CARDONAE–XYRIDION ACUTIFOLIAE Cuello & Cleef 2009
3. Cortaderio hapalotrichae –Hypericetum juniperinum Cuello & Cleef 2009 3.1. subassociation typicum Cuello & Cleef 2009
3.2. disterigmetosum acuminatum Cuello & Cleef 2009
4. Puyo aristeguietae –Ruilopezietum lopez-palacii Cuello & Cleef 2009 5. Rhynchosporo gollmeri –Ruilopezietum jabonensis Cuello & Cleef 2009
Lower Subpáramo
The zonal vegetation of the Guaramacal subpáramo corresponds to very dense shrub formations, growing on concave or wind protected slopes, forming the transition to high Andean forest (Subalpine rain forest or SARF). The subpáramo vegetation is represented by the new alliance Hyperico paramitanum - Hesperomeletion obtusifoliae, composed of two new associations Ruilopezio paltonioides - Neurolepidetum glomeratae and Disterigmo acuminatum - Arcytophylletum nitidum. Several species of small trees (typical) of the high-Andean forest are common, especially from the Ruilopezio paltonioides -Cybianthion marginati (Cuello & Cleef 2009a). They are growing in combination with high densities of tussock grasses dominated by Cortaderia hapalotricha, and the bamboo Chusquea angustifolia together with shrubs (up to 2 m) and proper woody páramo species, such as Hypericum juniperinum, Arcytophyllum nitidum,
Chaetolepis lindeniana, among other species of Hypericum, Asteraceae and
Ericaceae.
Upper Subpáramo
The zonal upper subpáramo vegetation corresponds to open vegetation pertaining to the new Hyperico cardonae - Xyridion acutifoliae alliance. This upper sub-páramo vegetation extends in greater proportion on low inclined convex slopes, and is represented by grasspáramo of the Puyo aristeguietae-Ruilopezietum lopez-palacii; bordered by or combined, with the vegetation of the new association Cortaderio hapalotrichae - Hypericetum juniperinum. There, the grasses
Corta-deria hapalotricha and Chusquea angustifolia also predominate, with variable
densities of rosettes of Ruilopezia lopez-palacii and Puya aristeguietae, prostrate herbs and a variable density of woody individuals among which the single-stemmed leptophyllous dwarfshrub (1.5 m) Hypericum juniperinum stands out. Towards the highest altitude (2900-3100 m), the open páramo vegetation of the (new) association Rhynchosporo gollmerii - Ruilopezietum jabonensis, located on concave slopes or in small depressions, is present. In this, the small (prostrate and erect) shrubs are absent (or very rare) and the 'frailejón' that dominates is the ground rosette Ruilopezia jabonensis. Cushion Cyperaceae, like Rhynchospora
gollmerii, and prostrate herbs occur more commonly. Another vegetation type
present in Páramo de Guaramacal is the bamboo-páramo ('chuscales') of the Carici bonplandii–Chusqueetum angustifoliae association (Chapter 4, Cuello & Cleef, 2009c), characterized almost exclusively by Chusquea angustifolia. The 'chuscales' of this association are located on humid, slightly sloping, ground of valleys or adjacent to lakes. They are considered azonal vegetation since they are periodically influenced by flood. As one move away from the chuscales, the density of individuals of Hypericum juniperinum increases, the number of clumps of Chusquea angustifolia bamboos decrease, and other grasses, rosettes and small shrubs appear conforming the vegetation of the corresponding association which is either Cortaderio hapalotrichae - Hypericetum juniperinum or that of Puyo aristeguietae - Ruilopezietum lopez-palacii.
Table 3.1. Phytosociological table of zonal páramo vegetation of Ramal de Guaramacal, Andes, Venezuela.
Releve number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
Releve (field number) 47a 47b 48b 32b 48a 39a 39b 11a 32a 3a 12a 12b 2a 19a 19b 2b 46a 46b 3b 45a 45b 29b 37a 29a 18b 34a 34b 43a 43b 18a 31a 31b 49b 7a 17a 17b 37b 7b
A 3 3 3 2 3 2 2 2 2 2 2 2 2 3 3 2 3 3 2 3 3 2 2 2 3 2 2 3 3 3 2 2 3 3 3 3 2 3
L 0 0 0 8 0 8 8 8 8 8 9 9 9 0 0 9 0 0 8 0 0 9 9 9 0 8 8 0 0 0 9 9 0 0 0 0 9 0
T 3 3 0 6 0 6 6 6 6 8 5 5 8 4 4 8 8 8 3 6 6 5 2 5 4 5 5 0 0 4 6 6 3 4 2 2 2 4
(m) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 5 5 0 0 0 0 0 0 0 0 0 0 0 0 5 0
Slope exposition NW NW N SE N NW NW NW SE SE S S W NW NW W SE SE NE SW SW NE S NE N NW NW S S N NE NE NW SE N N S SE
Slope angle (degrees) 45 45 30 19 30 30 30 18 17 20 25 30 18 13 37 22 22 22 20 35 35 36 10 29 12 24 24 25 25 12 21 37 23 28 18 25 10 29
Slope shape 2 2 1 1 1 2 2 2 1 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 1 2
Soils depth (cm) 30 50 50 46 95 106 45 38 60 >90 25 34 40 53 >80 45 40 13 >55 10 10 41 4 33 67 60 56 75 62 20 35 35 >110 53 56 17 30 25
pH 4 4.0 4.0 3.7 4.0 3.6 3.9 3.7 3.9 3.9 3.7 3.8 4.0 4.0 3.3 3.5 3.7 3.9 4.0 4.1 4.1 4* 3.5* 3.5* 3.70 3.7 3.7 4.5 4.5 3.7 3.7 3.8 4.2 4.2 3.8 3.7 3.4 3.7
Soils texture Fa aF FAa La Fa F La FL a Fla aL FL FaL FaL aL a aF Fa FLa aF aF a a a aF La La Fa Fa aF a a A FLA aL a aL FaL
No. vascular species 17 8 15 17 10 12 14 12 17 19 16 17 18 18 16 22 17 20 19 11 13 18 17 21 17 14 18 18 14 16 17 15 11 15 16 17 13 13 % outcrops and/or bare soil <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 6 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1
% Cov. Shrubs & dwarf trees >60 cm 35 30 80 50 70 10 15 15 25 60 45 50 25 70 55 85 100 60 50 15 15 70 65 85 45 70 40 45 30 5 35 20 35 25 30 35 30 20
% Cov. Small shrubs < 60 cm 5 5 15 30 10 5 10 5 10 20 20 40 39 20 30 20 45 25 45 50 45 5 10 20 20 20 10 45 35 5 30 25 15 15 20 30 15 10
% Cov. Grasses & rosettes > 10 cm 100 75 80 45 100 85 60 90 65 30 65 25 65 15 20 30 10 15 30 10 20 40 20 45 35 60 85 65 80 90 35 45 45 65 25 35 40 50
% Cov. Ground < 10 cm (including Cryptogams) 20 15 5 5 10 35 40 25 10 25 45 10 25 50 45 25 30 40 10 60 25 5 15 15 45 15 35 35 5 15 30 40 50 35 60 35 10 45
Order Alliance Association Subasociacion Variant Ruilopezia paltonioides . . 4 3 4 2 . 4 3 3 2 . . . 3 . . 4 . 1 . . 3 . 1 . . . . Disterigma alaternoides 1 . 3 2 2 . . . 5 . 3 . . . . Nertera granadensis 1 . 1 . . . 1 . . 1 . . . . Pentacalia greenmaniana . . 2 1 . . . . Sphyrospermum buxifolium . . 2 . . . . Disterigma acuminatum 1 . . . . 1 3 . . 4 4 5 3 4 4 2 5 4 4 5 5 2 2 3 4 3 4 3 4 2 4 4 2 4 4 4 4 1 Gaultheria hapalotricha 1 . . . 1 1 1 2 2 1 1 . . . . 1 . 1 1 . 1 1 . . 1 . . . 2 2 . . Arcytophyllum nitidum 1 . . . 4 2 . 4 2 . . . 4 . . 1 1 2 3 2 2 1 4 4 . 4 Ageratina theifolia . . . 2 1 . 1 . . . . 1 . . . 2 . . . 3 . . . . Galium hypocarpium . . . 1 . 2 . . . . Polypodium funckii . . . 1 . . . 1 Eriosorus flexuosus . . . 1 . . . 1 . . . . Hymenophyllum myriocarpum . . . 1 . 1 . . . . Pentacalia cachacoensis . . . 1 2 . . 3 2 2 . 1 2 3 3 . . . 3 . Vaccinium corymbodendron . 4 3 . . . 1 1 1 4 . 2 2 2 . . 1 . . . 2 . . . . Melpomene moniliformis . . . 1 1 1 . . 1 1 1 3 . . . . 1 . . . . Gaultheria anastomosans . . . 2 2 2 2 . . 1 . . . 1 . . . 2 . . . . Themistoclesia dependens . . . 2 . . . . 1 3 . . . 1 . . . . Hesperomeles sp. . . . 2 . . . . Ugni myricoides . . . . 1 1 3 . . 2 . . . 1 . . 2 . . . . 3 . 1 . 3 1 . 2 2 . . . 2 . . Rubus acanthophyllos . . . 1 . 2 2 . 1 . 1 1 . . . . Ilex guaramacalensis . . . 1 1 . . . . 1 . . . . Valeriana quirorana . . . 1 . 5 . . . . . Blechnum schomburgkii 1 3 3 3 4 . 1 2 2 2 . 1 . 3 4 4 4 5 2 . 2 4 5 4 4 5 4 4 4 . 4 4 . 3 . . 2 2 Hypericum paramitanum 1 . 3 4 2 1 2 3 3 2 2 2 3 2 1 2 2 2 4 4 4 1 . 1 2 . . 4 3 2 1 2 3 3 1 3 . 3 Neurolepis glomerata 5 5 5 . 5 5 5 3 . 2 1 2 1 2 4 2 . . . 1 . . 4 . 4 3 5 . . . . Cybianthus marginatus . . . . 1 . 1 . . 5 3 4 . 1 4 1 4 2 4 . . . . 4 . . . 1 . . 1 . . . . Hesperomeles obtusifolia 4 3 . 4 . . . . 1 . 3 5 4 4 4 2 2 2 . 2 2 . . . 2 . . 1 . . . 1 1 . . Sphagnum meridense 4 3 2 . . . 2 3 . . 6 . 6 6 . . . 2 . . . 4 . . . . Libanothamnus griffinii 1 . . . . 2 2 . . . 4 . 3 3 . 5 . 4 . . . . Elaphoglossum cf. lingua 1 . . . 1 . . . 1 . . 1 . . . 1 . . . . 1 1 . . Puya sp. . . . 2 1 . . . 4 2 . . . 3 . . . . 1 . . . . Miconia tinifolia . . . 1 . . . 1 . . . 2 . . . . Muehlenbeckia tamnifolia . . 1 . . . 2 . . . . Epidendrum frutex . . 1 . . . 1 . . . . Myrsine dependens . . . 2 . 2 . . . 1 . 1 2 . . . . 1 . . . . Diplostephium obtusum . . . 2 . . . . 3 . . . . 1 3 . . . 4 . . Rhynchospora sp. . . . 3 . . 2 . 2 . . . . 3.Hypericetum juniperinum Hypericum juniperinum . . . 2 . . . 2 . . 1 . . . 4 1 1 . Orthrosanthus acorifolius . . . 1 . . . 2 . . . 1 . Calamagrostis sp. A . . . . Paepalanthus pilosus . . . . 4. Puyo aristeguietae - Ruilopezietum lopez-palacii
Puya aristeguietae . . . 3 . . . 2 . . . . Chusquea tessellata . . . . Castilleja fissifolia . . . . Festuca guaramacalana . . . . Monnina sp. . . . . Bejaria aestuans . . . . Rhynchospora lechleri . . . . Oreobolus venezuelensis . . . . 2.2. typicum 2.1. pentacalietosum cachacoensis 2.2. typicum
1. Ruilopezio paltonioides - Neurolepidetum glomeratae
2. Disterigmo acuminatae - Arcytophylletum nitidum
RUILOPEZIO LOPEZ-PALACII -
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE
2.1. pentacalietosum cachacoensis
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE 2. Disterigmo acuminatum - Arcytophylletum nitidum 1. Ruilopezio - Neurolepidetum glomeratae
39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 41a 41b 13a 15b 22a 22b 25b 40a 40b 8b 9a 9b 13b 1b 20a 20b 23a 23b 44a 44b 6a 6b 15a 25a 49a 10a 10b 1a 28a 28b 35a 35b 38b 42a 42b 8a 50a 50b 38a 11b 21a 21b 14a 14b 24b 5b 16a 16b 24a 4a 5a 4b 27a 3 3 3 2 3 3 3 3 3 2 2 2 3 2 3 3 3 3 3 3 3 3 2 3 3 2 2 2 2 2 2 2 2 3 3 2 3 3 2 2 2 2 2 2 3 2 2 2 3 2 2 2 3 0 0 0 9 0 0 0 0 0 8 9 9 0 8 0 0 0 0 0 0 0 0 9 0 0 8 8 8 8 8 8 8 8 0 0 8 0 0 8 8 8 8 9 9 0 9 9 9 0 9 9 9 0 2 2 1 8 5 5 6 2 2 8 1 1 1 2 5 5 3 3 4 4 4 4 8 6 4 4 4 2 6 6 7 7 7 2 2 8 4 4 7 0 8 8 6 6 5 9 6 6 5 6 9 6 5 5 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 0 0 0 5 0 0 0 0 S S W NE SE SE E S S NW S S W NE SE SE SW SW SE SE N N NE E NW N N NE E E NW NW E E E NW N N E NW N N NE NE E E E E E E E E SW 8 8 12 21 14 18 19 14 13 16 11 11 12 5 48 31 12 12 32 32 7 7 21 19 23 15 15 5 9 9 15 15 11 21 21 11 26 26 11 18 12 12 11 11 13 28 19 19 13 24 23 16 5 2 3 1 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 2 2 1 1 2 1 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 1 1 1 2 1 1 1 2 1 2 2 2 21 18 31 28 40 22 17 28 29 115 60 9 31 20 15 63 86 29 72 40 80 80 13 41 25 75 30 40 120 120 52 38 28 51 15 115 20 32 120 55 106 80 26 65 20 56 40 73 10 48 60 28 120 4.1 4.1 3.9 3.8 4.0 3.7 4.2 4.0 3.8 3.3 3.7 3.5 3.6 3.7 4.2 3.9 4.0 3.8 4.0 4.0 4.1 3.7 3.8 ## 4.3 3.9 3.9 4.5 4.7 4.9 3.7 3.9 3.6 3.7 3.7 3.7 4.1 4.4 3.6 4.1 4.1 4.1 4.0 3.9 3.9 4.2 3.9 3.8 4.0 4.0 4.1 4.2 3.9 aL aL a aF a aL aL aL aL La La aL a L LA aL aL AF Fa Fa La aL aL aF aF La FLA FL aL aL LF LF aL aL aL LF Aa a L FaL aL aL a a aL FaL aL aL aL L FLa a L
12 14 13 11 18 15 13 14 17 9 15 11 14 10 11 15 14 11 16 10 12 15 19 13 15 11 13 10 9 8 14 16 10 11 15 12 14 10 10 10 12 17 12 12 12 7 9 12 11 10 9 9 7 <1 10 5 <1 <1 <1 2 1 5 <1 1 2 5 20 5 5 1 2 <1 <1 1 15 <1 <1 <1 1 5 10 1 1 15 <1 <1 1 1 1 <1 <1 <1 <1 5 5 15 15 25 5 15 15 20 10 5 10 10 30 30 20 <1 10 5 15 5 10 10 30 10 20 20 50 15 40 45 20 50 20 20 30 3 30 10 0 5 0 0 0 10 0 0 0 25 0 2 0 30 0 0 0 0 0 0 0 0 0 0 0 0 0 5 5 10 5 5 20 3 5 10 5 5 15 5 1 5 5 15 35 40 20 15 15 5 5 10 5 5 1 <1 <1 15 5 5 10 10 5 5 2 1 5 1 5 2 2 <1 0 <1 <1 <1 3 0 4 0 50 40 50 70 50 30 45 70 45 45 60 60 25 70 25 80 55 45 75 30 55 10 40 60 75 90 70 50 75 85 75 65 90 75 70 60 100 100 90 60 65 65 60 50 60 80 70 70 70 60 85 80 75 10 15 40 25 50 50 20 40 15 20 20 10 40 15 35 15 10 5 25 25 30 20 25 10 50 5 5 10 5 5 20 10 15 5 20 5 45 1 15 40 10 15 20 30 15 15 20 10 10 10 10 5 10 . . 1 . . . 3 . . . 5 . . . . . . . 3 . . . . . 1 . . . 2 1 . . . . 1 . 1 . . . . . . . . . . . . . . . 2 1 3 2 3 4 3 3 3 2 1 . 1 1 . . . 3 . . 2 . . . . . . . 1 . . . 1 . . . . . . . 4 . 3 1 4 3 2 . . . 1 . . . 1 . . . . . . 1 . . . 2 . 1 . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . 1 1 . . . . 1 . 4 . . 1 1 . . . 1 2 . . . 4 . . . . . . . . 1 . . . . . . . 2 . . . . 3 2 . . . 1 . . . . . . . . 1 . . . . . . . 1 . . . 2 . . . 2 1 . . . . . . . . . . . . . . . 3 . . . 1 . . . . . . . 1 . 2 1 . 5 . . 1 . . . . 1 2 . . . 1 . 1 . . . . . . 1 . 2 4 . . . 1 . . . 2 . 1 . . 2 1 1 . . . 2 . 1 . . . . 3 . . . 3 . . . . 1 . . . 1 . . . 1 . . . 2 . . . . . . 1 . . . 2 . . . 3 . . 1 1 . 1 2 2 2 . 1 4 . . . . . . . 3 . . . . . . . 4 . . . 3 5 . 3 . . . 3 . . . . . . . 1 . . . . . . . . . . . . . . . . . . . 1 . . . . . . . 3 . . . . 2 . . . 1 . . . . 5 4 . . . 1 . . . 1 . . . 3 . . . . . . . 1 1 . . . . 3 . . . 1 . . . 2 . . . 3 . . . . 3 4 4 . 4 1 4 5 2 4 2 2 1 4 5 2 4 4 2 . 3 2 2 2 . . . 2 . . . 1 4 . 2 . . . 1 . 1 2 . . . . . 2 2 . 1 3 . 3 1 . 2 . . . 3 . . . 1 . . . . . . . 1 . 2 . 1 . . . . 2 . . . 1 . . . . . . . 3 . . . 1 4 2 3 3 3 2 3 . . 3 . . 4 . . 3 . . . . . . . 5 4 . 4 3 . . . . . . . 1 1 . . . 1 1 . . . . . . . 1 2 . . . . . . . 2 . . . . . . . 2 . . . . . . . 2 . . . . . . . 2 . . . 1 . . . 1 . . . 1 . 1 . . . 2 - CHUSQUEETALIA ANGUSTIFOLIAE
3.1. typicum 3.2. disterigmatosum acuminatum
3. Cortaderio hapalotrichae - Hypericetum juniperinum 4. Puyo aristeguietae - Ruilopezietum lopez-palacii 5. R. gollmeri - Ruilopezietum jabonensis HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
Releve number 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38
Releve (field number) 47a 47b 48b 32b 48a 39a 39b 11a 32a 3a 12a 12b 2a 19a 19b 2b 46a 46b 3b 45a 45b 29b 37a 29a 18b 34a 34b 43a 43b 18a 31a 31b 49b 7a 17a 17b 37b 7b
A 3 3 3 2 3 2 2 2 2 2 2 2 2 3 3 2 3 3 2 3 3 2 2 2 3 2 2 3 3 3 2 2 3 3 3 3 2 3
L 0 0 0 8 0 8 8 8 8 8 9 9 9 0 0 9 0 0 8 0 0 9 9 9 0 8 8 0 0 0 9 9 0 0 0 0 9 0
T 3 3 0 6 0 6 6 6 6 8 5 5 8 4 4 8 8 8 3 6 6 5 2 5 4 5 5 0 0 4 6 6 3 4 2 2 2 4
(m) 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 5 5 0 0 0 0 0 0 0 0 0 0 0 0 5 0
Slope exposition NW NW N SE N NW NW NW SE SE S S W NW NW W SE SE NE SW SW NE S NE N NW NW S S N NE NE NW SE N N S SE
Slope angle (degrees) 45 45 30 19 30 30 30 18 17 20 25 30 18 13 37 22 22 22 20 35 35 36 10 29 12 24 24 25 25 12 21 37 23 28 18 25 10 29
Slope shape 2 2 1 1 1 2 2 2 1 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 2 2 1 1 1 2
Soils depth (cm) 30 50 50 46 95 106 45 38 60 >90 25 34 40 53 >80 45 40 13 >55 10 10 41 4 33 67 60 56 75 62 20 35 35 >110 53 56 17 30 25
pH 4 4.0 4.0 3.7 4.0 3.6 3.9 3.7 3.9 3.9 3.7 3.8 4.0 4.0 3.3 3.5 3.7 3.9 4.0 4.1 4.1 4* 3.5* 3.5* 3.70 3.7 3.7 4.5 4.5 3.7 3.7 3.8 4.2 4.2 3.8 3.7 3.4 3.7
Soils texture Fa aF FAa La Fa F La FL a Fla aL FL FaL FaL aL a aF Fa FLa aF aF a a a aF La La Fa Fa aF a a A FLA aL a aL FaL
No. vascular species 17 8 15 17 10 12 14 12 17 19 16 17 18 18 16 22 17 20 19 11 13 18 17 21 17 14 18 18 14 16 17 15 11 15 16 17 13 13 % outcrops and/or bare soil <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 6 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1 <1
% Cov. Shrubs & dwarf trees >60 cm 35 30 80 50 70 10 15 15 25 60 45 50 25 70 55 85 100 60 50 15 15 70 65 85 45 70 40 45 30 5 35 20 35 25 30 35 30 20
% Cov. Small shrubs < 60 cm 5 5 15 30 10 5 10 5 10 20 20 40 39 20 30 20 45 25 45 50 45 5 10 20 20 20 10 45 35 5 30 25 15 15 20 30 15 10
% Cov. Grasses & rosettes > 10 cm 100 75 80 45 100 85 60 90 65 30 65 25 65 15 20 30 10 15 30 10 20 40 20 45 35 60 85 65 80 90 35 45 45 65 25 35 40 50
% Cov. Ground < 10 cm (including Cryptogams) 20 15 5 5 10 35 40 25 10 25 45 10 25 50 45 25 30 40 10 60 25 5 15 15 45 15 35 35 5 15 30 40 50 35 60 35 10 45
Order Alliance Association Subasociacion Variant 2.2. typicum RUILOPEZIO LOPEZ-PALACII - 2.1. pentacalietosum cachacoensis
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE 2. Disterigmo acuminatum - Arcytophylletum nitidum 1. Ruilopezio - Neurolepidetum glomeratae
Ruilopezia jabonensis . . . .
Rhynchospora gollmeri . . . .
Isidrogalvia robustior . . . .
Gentianella nevadensis . . . .
Xyris subulata var. acutifolia . . . 1 . . . .
Hypericum cardonae . . . 1 . . . 1 . . . . Carex bonplandii . . . . Ruilopezia viridis . . . . Calamagrostis planifolia . . . . Cortaderia hapalotricha . . . 4 . 3 4 5 3 2 4 2 5 4 4 4 2 3 4 2 4 3 1 2 4 2 3 4 3 4 2 2 . . 3 4 4 4 Chusquea angustifolia 3 4 1 4 5 . . 3 4 2 5 4 . . . 1 . . 2 . 2 4 3 3 1 4 4 4 4 2 2 4 5 5 3 4 5 5
Lycopodium clavatum subsp. contiguum1 . . 2 . 4 4 4 3 4 . . 3 2 1 1 . 1 2 2 1 . 4 1 1 1 2 2 . 2 2 2 3 2 3 . 2 3
Ruilopezia lopez-palacii . 2 . 2 . 4 3 . 3 . . . 4 . . 1 . 1 2 . . 2 1 4 4 4 4 . . 5 4 . 1 4 4 2 2 3 Geranium stoloniferum . . . 1 1 . 2 3 . 5 4 . 4 . 2 . . . 1 2 . . . 2 4 1 3 2 Pernettya prostrata 1 1 1 2 2 1 1 1 1 1 . . 1 1 2 . 1 1 2 3 4 1 2 1 1 2 1 2 . 1 3 2 4 1 1 3 1 1 Rhynchospora guaramacalensis 2 1 2 1 . . . . 3 3 . . . 2 2 2 3 . . . 3 3 3 2 4 1 . . . . Rhynchospora macrochaeta . . . 3 4 . . . 1 . 1 . . . 2 3 3 3 1 2 3 1 Jamesonia imbricata 2 . . 1 . 2 1 2 2 . . . 4 . . . 1 . . . . 2 . . . 1 2 1 . 1 3 2 . . . . Chaetolepis lindeniana . . 1 3 . . . 2 . . 1 1 2 2 . 1 . 2 3 . . 2 2 2 2 2 2 2 2 1 . . . 3 . . 4 . Daucus montanus 1 . . . 1 . . . . 1 . . . 1 1 1 . . . . 1 1 . . . . 2 . 2 2 1 1 . . 2 Sphagnum sparsum . . . 5 6 . . . 6 . . . . 3 . . 6 5 5 4 . 6 Hieracium avilae . . . 1 . . . 1 . . . 1 . 1 . . . . Hymenophyllum trichomanoides . . . 1 . . . 1 . . Hypericum sp. . . . 1 . . . 1 . . . . 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 1 1 0 0 1 1 1 0 0 0 0 0 0 0 0 1 1 1 1 1 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 0 0 0 1 1 1 1 1 0 0 0 1 1 0 1 1 1 1 1 0 0 0 0 0 1 1 1 1
Cybianthus laurifolius? 3a(1) Huperzia amentacea 3b(1) Melpomene xiphopteroides 17b(1) Gaultheria erecta 34b(1) Hymenophyllum sp. 34a(1) Polypodium sp. 34b(1) Greigia sp. 44a(1) Melpomene flabelliformis 17a(1) Utricularia alpina 21b(1)
5. R. gollmeri - Ruilopezietum jabonensis
RUILOPEZIO LOPEZ-PALACII - CHUSQUEETALIA ANGUSTIFOLIAE HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 41a 41b 13a 15b 22a 22b 25b 40a 40b 8b 9a 9b 13b 1b 20a 20b 23a 23b 44a 44b 6a 6b 15a 25a 49a 10a 10b 1a 28a 28b 35a 35b 38b 42a 42b 8a 50a 50b 38a 11b 21a 21b 14a 14b 24b 5b 16a 16b 24a 4a 5a 4b 27a 3 3 3 2 3 3 3 3 3 2 2 2 3 2 3 3 3 3 3 3 3 3 2 3 3 2 2 2 2 2 2 2 2 3 3 2 3 3 2 2 2 2 2 2 3 2 2 2 3 2 2 2 3 0 0 0 9 0 0 0 0 0 8 9 9 0 8 0 0 0 0 0 0 0 0 9 0 0 8 8 8 8 8 8 8 8 0 0 8 0 0 8 8 8 8 9 9 0 9 9 9 0 9 9 9 0 2 2 1 8 5 5 6 2 2 8 1 1 1 2 5 5 3 3 4 4 4 4 8 6 4 4 4 2 6 6 7 7 7 2 2 8 4 4 7 0 8 8 6 6 5 9 6 6 5 6 9 6 5 5 5 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 0 0 0 5 0 0 0 0 S S W NE SE SE E S S NW S S W NE SE SE SW SW SE SE N N NE E NW N N NE E E NW NW E E E NW N N E NW N N NE NE E E E E E E E E SW 8 8 12 21 14 18 19 14 13 16 11 11 12 5 48 31 12 12 32 32 7 7 21 19 23 15 15 5 9 9 15 15 11 21 21 11 26 26 11 18 12 12 11 11 13 28 19 19 13 24 23 16 5 2 3 1 1 1 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 2 2 1 1 2 1 1 1 2 2 1 1 1 1 1 2 2 2 1 2 1 1 1 1 2 1 1 1 2 1 2 2 2 21 18 31 28 40 22 17 28 29 115 60 9 31 20 15 63 86 29 72 40 80 80 13 41 25 75 30 40 120 120 52 38 28 51 15 115 20 32 120 55 106 80 26 65 20 56 40 73 10 48 60 28 120 4.1 4.1 3.9 3.8 4.0 3.7 4.2 4.0 3.8 3.3 3.7 3.5 3.6 3.7 4.2 3.9 4.0 3.8 4.0 4.0 4.1 3.7 3.8 ## 4.3 3.9 3.9 4.5 4.7 4.9 3.7 3.9 3.6 3.7 3.7 3.7 4.1 4.4 3.6 4.1 4.1 4.1 4.0 3.9 3.9 4.2 3.9 3.8 4.0 4.0 4.1 4.2 3.9 aL aL a aF a aL aL aL aL La La aL a L LA aL aL AF Fa Fa La aL aL aF aF La FLA FL aL aL LF LF aL aL aL LF Aa a L FaL aL aL a a aL FaL aL aL aL L FLa a L
12 14 13 11 18 15 13 14 17 9 15 11 14 10 11 15 14 11 16 10 12 15 19 13 15 11 13 10 9 8 14 16 10 11 15 12 14 10 10 10 12 17 12 12 12 7 9 12 11 10 9 9 7 <1 10 5 <1 <1 <1 2 1 5 <1 1 2 5 20 5 5 1 2 <1 <1 1 15 <1 <1 <1 1 5 10 1 1 15 <1 <1 1 1 1 <1 <1 <1 <1 5 5 15 15 25 5 15 15 20 10 5 10 10 30 30 20 <1 10 5 15 5 10 10 30 10 20 20 50 15 40 45 20 50 20 20 30 3 30 10 0 5 0 0 0 10 0 0 0 25 0 2 0 30 0 0 0 0 0 0 0 0 0 0 0 0 0 5 5 10 5 5 20 3 5 10 5 5 15 5 1 5 5 15 35 40 20 15 15 5 5 10 5 5 1 <1 <1 15 5 5 10 10 5 5 2 1 5 1 5 2 2 <1 0 <1 <1 <1 3 0 4 0 50 40 50 70 50 30 45 70 45 45 60 60 25 70 25 80 55 45 75 30 55 10 40 60 75 90 70 50 75 85 75 65 90 75 70 60 100 100 90 60 65 65 60 50 60 80 70 70 70 60 85 80 75 10 15 40 25 50 50 20 40 15 20 20 10 40 15 35 15 10 5 25 25 30 20 25 10 50 5 5 10 5 5 20 10 15 5 20 5 45 1 15 40 10 15 20 30 15 15 20 10 10 10 10 5 10 - CHUSQUEETALIA ANGUSTIFOLIAE
3.1. typicum 3.2. disterigmatosum acuminatum
3. Cortaderio hapalotrichae - Hypericetum juniperinum 4. Puyo aristeguietae - Ruilopezietum lopez-palacii 5. R. gollmeri - Ruilopezietum jabonensis HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
. . . 2 . . . 3 . . . 5 5 5 5 5 5 5 4 5 5 2 . . . 1 . . . 2 2 1 1 3 1 1 . . . 5 . . . 1 . . . 1 . . . 2 2 . . . . . . . 1 . . . . 1 . . . . 5 4 3 3 1 4 1 2 2 3 . . 2 . 1 . . . . 2 . . 1 . . . . 1 1 4 . 1 . 2 3 4 . . 2 2 2 2 4 3 4 5 1 2 1 2 3 . 5 1 1 2 2 1 1 1 1 1 . 1 . 1 1 . . . 1 1 . . . 1 1 1 . . . . 1 1 1 . 1 1 1 . . . . 3 1 . . . 2 2 . . . 1 . . . 1 . . . . 4 1 . . . 3 . . . 2 . . . . . . . 1 . . 1 . . . . 3 5 4 4 5 4 5 5 4 1 3 5 4 5 3 3 4 4 3 4 5 5 3 4 2 5 4 5 4 4 5 5 4 4 4 3 4 2 4 5 5 4 5 5 5 4 5 4 5 5 4 5 1 . . 4 3 . 1 2 4 5 3 4 2 . 3 . 5 4 4 4 . 3 2 3 . 5 3 2 1 5 3 4 3 5 1 3 3 5 5 . . . . 3 3 3 1 4 5 4 4 3 5 5 1 3 3 3 3 3 3 3 3 3 2 3 2 4 2 3 2 2 2 4 3 3 2 1 3 4 4 3 4 4 2 3 2 3 1 4 3 . 3 5 4 3 2 3 1 2 4 2 2 2 3 . . . . . 5 2 . 4 2 1 5 3 4 . 3 . . . . 4 . . 1 4 5 3 5 5 5 4 5 3 5 5 5 5 5 3 5 5 . 5 5 4 . . . 4 . . . 5 3 5 5 4 . . 2 4 4 5 3 5 4 5 5 3 4 5 5 3 4 . . . 2 . . . . 3 . 2 1 . . 2 . . . 1 2 1 . . 2 . 3 3 1 . 2 2 3 2 1 2 2 3 4 3 2 4 1 1 2 1 3 4 3 3 1 2 1 3 3 2 1 1 . . 3 2 1 . 1 2 1 1 1 1 1 1 . 1 1 . . 1 . 1 . . . . . . 3 2 . . 2 4 1 . . . 4 3 4 4 1 4 4 . . . 3 . 4 3 . . . . 1 2 1 3 2 . 1 . . 1 3 . . . 2 . . . 3 3 3 3 1 . . . 1 2 2 . . . . 2 1 3 1 1 1 2 3 . 3 . . . . 1 . . 1 . . . 1 2 1 1 . 3 . . . 1 2 3 3 1 . . . 1 2 3 2 2 . 2 4 1 1 1 4 . . . 2 . 1 1 2 . . 3 . . 1 2 . 2 1 . 5 2 . . . 2 1 . . . 2 . 1 . . . 1 . . 2 . . . . . 1 . . . 2 3 . . . 1 . . . . 2 2 . . 1 . . . 4 1 . . . . . . . 5 3 . 3 . . . 2 . . 6 . . . 6 . . . . . . 1 . 1 . . . 1 . . . 1 1 . . . . 1 . . . 1 . . . 1 . 2 1 1 . 1 . . . . . . . 3 . . . 1 . . . . . . . 2 . . . . 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 1 1 0 0 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 1 1 0 0 0 0 1 1 1 1 1
Soil texture: F franco (loamy) a arena (sand) A arcilla (clay) L Limo (mud/silt) 1 convex 2 concave Slope shape:
RUILOPEZIO LOPEZ-PALACII – CHUSQUEETALIA ANGUSTIFOLIAE Cuello & Cleef
2009
Representative alliance: Hyperico paramitanum - Hesperomeletion obtusifoliae Provisional order of zonal humid lower páramo of Ruilopezia lopez-palacii and Chusquea
angustifolia / Orden provisional de páramo húmedo bajo zonal de Ruilopezia lopez-palacii
y Chusquea angustifolia
Physiognomy and composition: A vegetation mosaic of very humid subpáramo and páramo, with rosettes and bamboos growing among patches of ecotonic dwarf forest. A variety of growth forms is characteristic, including: acaulescent and stem rosettes, dwarf trees, small (upright and prostrate) shrubs; epiphytic, erect prostrate and trailing herbs, and grass tussocks and bamboos. Also noticeable are a variety of ferns and a dense cover of bryophytes and lichens. Locally appear patches of reddish Sphagnum mosses. Diagnostic species are: Chaetolepis lindeniana,
Chus-quea angustifolia, Cortaderia hapalotricha, Daucus montanus, Geranium stoloni-ferum, Hymenophyllum trichomanoides, Jamesonia imbricata, Lycopodium con-tiguum, Pernettya prostrata, Rhynchospora guaramacalensis, R. macrochaeta and Ruilopezia lopez-palacii.
Syntaxonomy: This provisional order is defined on the basis of 91 line-intersect surveys with 85 vascular species. This order groups both the alliances of humid shrub subpáramos of Hyperico paramitanum -Hesperomeletion obtusifoliae and Hyperico cardonae - Xyridion acutifoliae of shrub páramos and grassy lower subpáramos.
Ecology and distribution: The order unifies all communities of zonal vegetation (excluding dwarf forests) present in the summit region of Ramal de Guaramacal between 2800 and 3130 m.
HYPERICO PARAMITANUM – HESPEROMELETION OBTUSIFOLIAE Cuello & Cleef 2009
Typus: Ruilopezio paltonioides–Neurolepidetum glomeratae.
Shrubpáramo of the Hypericum paramitanumi and Hesperomeles obtusifolia alliance / Subpáramo de arbustales de la alianza de Hypericum paramitanum y Hesperomeles
obtusifolia
Physiognomy and composition: This alliance groups vegetation communities with a high proportion of shrubs and dwarf tree species. The shrubpáramo displays variable densities of Ruilopezia paltonioides and R. lopez-palacii stem rossettes, within a matrix of Cortaderia hapalotricha tussock grasses and Chusquea
angustifolia bamboos. These shrub formations can reach heights of 1.5-2 m,
occasionally reaching upwards of 3 m in wind protected areas. In the understorey, very common low shrubs of Hypericum paramitanum and prostrate shrubs of
Disterigma acuminatum are present. A variable density of the tall and wide-leaved
bamboo Neurolepis glomerata and an abundant turf cover of Sphagnum and other bryophytes are distinctive
Dwarf tree species of high Andean forest (or subalpine rain forest, SARF) are common, such as: Cybianthus laurifolius, C. marginatus, Gaultheria erecta,
Hes-peromeles obtusifolia, Ilex guaramacalensis, Libanothamnus griffinii, Miconia tinifolia, Myrsine dependens, and Vaccinium corymbodendron. Also present are
typical open páramo dwarf treelets, such as: Ageratina theifolia, Hypericum
juniperinum and Hesperomeles sp.
Between the shrubs, and distinctive in the sequence of abundance, are: Hypericum
paramitanum, Chaetolepis lindeniana, Arcytophyllum nitidum, Ugni myricoides, Disterigma alaternoides, Pentacalia cachacoensis, Valeriana quirorana, Gaul-theria anastomosans, Diplostephium obtusum, Pentacalia greenmaniana, Hype-ricum juniperinum x cardonae. Small ericaceous prostrate shrubs including: Disterigma acuminatum, Pernettya prostrata, Gaultheria hapalotricha, Themisto-clesia dependens and Sphyrospermum buxifolium are also present.
Apart of the prominent bamboos Chusquea angustifolia and Neurolepis glomerata are also important tussocks of Cortaderia hapalotricha, Rhynchospora
guara-macalensis and R. macrochaeta.
Other species include herbs like Daucus montanus, Epidendrum frutex, Hypericum
cardonae, Geranium stoloniferum, Nertera granadensis and ferns and clubmosses
such as: Elaphoglossum cf. lingua, Eriosorus flexuosus, Huperzia amentacea,
Jamesonia imbricata, Lycopodium clavatum subsp. contiguum, Polypodium funckii, Hymenophyllum myriocarpum, H. trichomanoides, Melpomene flabella-formis, M. moniliformis and M. xiphopteroides. The trailings Rubus acantho-phyllos and Muehlenbeckia tamnifolia are also present.
Syntaxonomy: Thirty-eight line-intersect surveys are recognized as belonging to this alliance with a total of 65 vascular species accounting for species richness. Diagnostic species for the alliance are: Blechnum schomburgkii, Cybianthus
mar-ginatus, Hesperomeles obtusifolia, Hypericum paramitanum, Libanothamnus griffinii and Neurolepis glomerata.
This new provisional alliance contains two associations: Ruilopezio paltonioides - Neurolepidetum glomeratae and Disterigmo acuminatum - Arcytophylletum nitidum.
Ecology and distribution: This alliance groups zonal vegetation characteristic of humid shrub subpáramo in the páramo-forest ecotone. Vegetation of this type is situated mainly on predominantly convex slopes between 2830 and 3080 m, with slopes of between 5 to 48 degrees. The soils are, in general, comparatively deep, with a layer of organic matter, sand-muddy textures and acidic (average pH 3.8) in the superficial layers. The associations of this alliance shares many species in co-mmon with those of dwarf forests alliance of Ruilopezio paltoniodes–Cibianthion marginatus, and may be contiguous in the field, however, differences in ecology (soil depth, light exposition, humidity level in underbrush) and the presence of proper open páramo diagnostic species in the shrubpáramo associations help to difference between alliances.
1. Ruilopezio paltonioides –Neurolepidetum glomeratae Cuello & Cleef 2009 Typus: Rel. No. 3 (Cuello L48b). Table 3.1. Figure 3.2. Photo 3.1
Humid shrub páramo of Ruilopezia paltonioides and Neurolepis glomerata / Pajonal-arbustal de subpáramo húmedo de Ruilopezia paltonioides y Neurolepis glomerata Physiognomy and composition: Shrub community with a high density of tall tussock grasses and wide-leaved bamboos (1-1.5 m) and between 35-50% cover, growing among a layer of dwarf trees and dispersed shrubs (Fig. 3.2). Tall conspicuous espeletioid stem rossettes reaching 2 (3) m with 15 to 25% cover are also present.
The upper layer is composed of discrete Chaetolepis lindeniana, Hesperomeles
obtusifolia, Hypericum paramitanum and Ugni myricoides shrubs, together with
tall (2-3 m) Ruilopezia paltonioides stem rosettes and lower ones of Ruilopezia
lopez-palacii and Blechnum schomburgkii. In the tall grass layer, additional to the
dominance of Neurolepis glomerata (20-40% cover), Chusquea angustifolia and
Cortaderia hapalotricha are also present. Further, there is also a low herb layer
containing prostrate shrubs Disterigma acuminatum and Pernettya prostrata, the sedges Rhynchospora guaramacalensis and R. macrochaeta, small herbs like
Daucus montanus, and the ferns Jamesonia imbricata and Lycopodium clavatum
subsp. contiguum, growing over a turf of Sphagnum sparsum and S. meridense among other bryophytes.
Syntaxonomy: This association is defined on the basis of 10 line-intersect surveys, with a total of 41 vascular species. Ruilopezia paltonioides and Neurolepis
glomerata are diagnostic. Other diagnostic species in this association include: Disterigma alaternoides, Pentacalia greenmaniana and Sphyrospermum buxifolium.
Two provisional variants are distinguished for this association: a variant of
Disterigma alaternoides and a variant of Ugni myricoides.
Ecology and distribution: Transitional ecotonic shrubby vegetation of the humid sub-páramo located close to the upper forest line, consisting of (subalpine rain forests or SARF sensu Grubb, 1977) of Libanothamnus griffinii, and Gaultheria
anastomosans and Hesperomeles obtusifolia dwarf forests (Cuello&Cleef, 2009). This association has been observed between 2860 to 3000 m on concave or convex slopes with NW-SE exposition and slope angles between 18 and 30 degrees. This community can also be found near rock outcrops or along fractured rocks crossed by small streams. The soils are 38-106 cm deep, loamy to loam-sandy loam in texture, with gray to brown yellowish colours and of pH, 3.6 to 3.9 in the upper layer.
1.1. variant of Disterigma alaternoides
Physiognomy and composition: Dense shrubby-grass vegetation dominated by Neurolepis glomerata bamboo clumps (1-1.5 m, 35-40% cover), a layer of discrete
shrubs and dwarf trees (2-3 m, 20-25% cover) and small prostrate shrubs in the interior. Species composition is as described for the association.
Diagnostic species are Disterigma alaternoides, Sphyrospermum buxifolium,
Pentacalia greenmaniana and Vaccinium corymbodendron. This variant is
distinguished from the variant of Ugni myricoides by the low presence of
Cortaderia hapalotricha and a greater presence of Chusquea angustifolia.
Ecology and distribution: This variant corresponds to the vegetation of the association of Ruilopezia paltonioides and Neurolepis glomerata located at altitudes of around 3000 m, generally transitional and adjacent to dwarf forests of
Libanothamnus griffinii.
Photo 3.1. Closer view of a shrub páramo vegetation of the Ruilopezio paltonioides -Neurolepidetum glomeratae on the border of a patch of dwarf forest at ~2890 m in Páramo de Guaramacal, Ramal de Guaramacal, Andes, Venezuela. Notice the dominance of the tall stem rosette Ruilopezia paltonioides.
Figure 3.2. Physiognomy of the vegetation of the association Ruilopezio paltonioides - Neurolepidetum glomeratae var. Disterigma alaternoides (L48b 3000 m). Bs:
Blechnum schomburgkii; Cha: Chusquea angustifolia; Chl: Chaetolepis lindeniana; Da: Disterigma alaternoides; Ef: Epidendrum frutex; Hp: Hype-ricum paramitanum; Mp: Muehlenbeckia tamnifolia; Ng: Neurolepis glome-rata; Ngr: Nertera granadensis; Pg: Pentacalia greenmanniana; Pp: Pernettya prostrata; Rgu: Rhynchospora guaramacalensis; Rp: Ruilopezia paltonioides; Sb: Sphyrospermum buxifolium; Vc: Vaccinium corymbo-dendron.
1.2 variant of Ugni myricoides
Physiognomy and composition: Dense shrubby-grass vegetation of high Neurolepis glomerata clumps (15-20%), dispersed shrubs (15-20%) and a high Flora, vegetation and ecology in the Venezuelan Andes
cover of low tussocks (25-30%) with a dominance of Cortaderia hapalotricha. See for species composition the association.
The diagnostic species in this variant are Ugni myricoides and Disterigma
acuminatum. The presence of Cortaderia hapalotricha is also significant and a
greater presence and cover of Lycopodium clavatum subsp. contiguum, Ruilopezia
lopez-palacii and Jamesonia imbricata distinguish this variant.
Ecology and distribution: This variant corresponds to the vegetation of the association of Ruilopezia paltonioides and Neurolepis glomerata located at altitudes of 2800-2900 m. Stands are generally adjacent to both dwarf forests of
Libanothamnus griffinii or those of Gaultheria anastomosans and Hesperomeles obtusifolia (Cuello & Cleef 2009a), in addition to their presence along small streams.
2. Disterigmo acuminatum –Arcytophylletum nitidum Cuello & Cleef 2009 Typus: Rel. No. 31 (Cuello L31a). Table 3.1. Figure 3.3
Humid Disterigma acuminatum and Arcytophyllum nitidum shrub páramo / Arbustal de páramo húmedo de Disterigma acuminatum y Arcytophyllum nitidum
Physiognomy and composition: Dense shrubby vegetation, with a variable frequency of tall stem rosettes and tussock grasses. The aspect is a layer of shrubs and dwarf trees around 1-1.5 (3) m tall, with 20-40% cover, and a layer of tall tussock grasses that reach up to 1.5-2 m with 20 to 25% cover. In the dwarf shrub layer are ericaceous prostrate shrubs (30-50 cm and 15-18% cover), other grasses (15-45 cm and 2-6% cover) and a ground layer consisting of cushions species of
Sphagnum and other bryophytes (60-80% cover).
Among the shrub and dwarf tree (dt) species with substantial cover are
Arcytophyllum nitidum, Chaetolepis lindeniana, Cybianthus marginatus (dt), Disterigma alaternoides, Hesperomeles obtusifolia (dt), Hypericum paramitanum, Libanothamnus griffinii (dt), Pentacalia cachacoensis (dt), Ugni myricoides and Vaccinium corymbodendron (dt).
Among the bamboo and tussock grasses are Chusquea angustifolia and Cortaderia
hapalotricha in the shrub layer; Rhynchospora guaramacalensis, R. macrochaeta, Orthrosanthus acorifolius and Xyris subulata var. acutifolia are present in the herb
layer. The stem rosettes of Blechnum schomburgkii, Ruilopezia lopez-palacii and
Ruilopezia paltonioides are conspicuous. Common small shrubs include Disterigma acuminatum, Gaultheria hapalotricha, Hypericum cardonae, Pernettya prostrata and Themistoclesia dependens, and scandents or climbers like Muehlenbeckia tamnifolia and Rubus acanthophyllos. Further, the tall erect
terrestrial orchid Epidendrum frutex, small or prostrate herbs like Daucus
montanus, Galium hypocarpium, Geranium stoloniferum, and a diversity of ferns
and club mosses, such as Elaphoglossum cf. lingua, Eriosorus flexuosus, Huperzia
amentacea, Hymenophyllum myriocarpum, H. trichomanoides, Jamesonia imbricata, Lycopodium clavatum subsp. contiguum, Melpomene moniliformis, M.
flabelliformis, M. xiphopteroides and Polypodium funckii, are also present, among
others.
Syntaxonomy: This is a highly diverse association represented by 28 line-intersect surveys with 61 species of vascular plants.
Diagnostic species are Arcytophyllum nitidum, Ageratina theifolia, Disterigma
acuminatum and Gaultheria hapalotricha. Two subassociations are distinguished,
pentacalietosum cachacoensis and the typicum one.
Ecology and distribution: This subpáramo bamboo shrub is generally found surrounding areas of dwarf forests (SARF), at edges of slopes or hill tops, and in contact with communities of Ruilopezia paltonioides and Neurolepis glomerata. It represents humid shrub páramo, transitional between forest and páramo.
Figure 3.3. Physiognomy of the vegetation of the association Disterigmo acuminatum - Arcytophylletum nitidum subass. Typicum (L31a 2960 m). An: Arcytophyllum
nitidum; Bs: Blechnum schomburgkii; Cha: Chusquea angustifolia; Cm: Cybianthus marginatus; Da: Disterigma acuminatum; Dm: Daucus montanus; Gh: Gaultheria hapalotricha; Hp: Hypericum paramitanum; Ji: Jamesonia imbricata; Lc: Lycopodium clavatum subsp. contiguum; Ng: Neurolepis glomerata; Pp: Pernettya prostrata; Ra: Rubus acanthophyllos; Rl: Ruilopezia lopez-palacii; Rm: Rhynchospora macrochaeta; Um: Ugni myricoides; V: Valeriana quirorana.
Disterigmo acuminatum – Arcytophylletum nitidum
2.1. subassociation pentacalietosum cachacoensis Cuello & Cleef 2009 Typus: Rel. No. 17 (Cuello L46a). Table 3.1. Figure 3.4
Pentacalia cachacoensis subassociation / Subasociación de Pentacalia cachacoensis
Physiognomy: Dense shrubby vegetation in a matrix of tussock grasses of Cortaderia hapalotricha and bamboos of Chusquea angustifolia and Neurolepis glomerata; shrubs, dwarf trees (1-1.5 (3) m) and prostrate shrubs are present at
high density. There is a carpet of species of Sphagnum, together with other mosses,
as well as the presence of liverworts, such as Scapania portoricensis and species of
Plagiochila.
Composition and syntaxonomy: This subassociation is represented in 13 line-intersect surveys containing 50 vascular species. Diagnostic species are Pentacalia
cachacoensis and Vaccinium corymbodendron, together with Ageratina theifolia, Cybianthus marginatus, Gaultheria anastomosans, Hesperomeles obtusifolia, Themistoclesia dependens and the fern Melpomene moniliformis. The ground layer
of this vegetation unit is dominated by Sphagnum meridense and S. sparsum and among them Breutelia rithidoides and Cladonia furcata can also be found. Other epiphytes on small trunks are species of Riccardia (2955), Frullania (3038, 3039) and Plagiochila (2957).
Some facies may be distinguished for this subassociation: a facies of Vaccinium
corymbodendron, characterized also with a prominent presence of Melpomene monniliformis and Gaultheria anastomosans and another facies with a greater
presence of Libanothamnus griffinii.
Ecology and distribution: The shrub páramo of the subassociation of Pentacalia cachacoensis is located at altitudes between 2920-3080 m, and occuring on the
edges of convex or concave slopes of 10-37 degrees. The soils attain a depth of 4-80 cm, with mixed textures predominantly sandy (sand-muddy to sand-silty or silt-sand-loam), with pH 3.3-4.1 and dark colors in the superficial layers, varying in colour until reddish and grayish with a high clay content at increased depth. Disterigmo acuminatum –Arcytophylletum nitidum
2. 2. subassociation typicum Cuello & Cleef 2009 Typus: Rel. No. 31 (Cuello L31a). Table 3.1. Figure 3.3
Subassociation of Arcytophyllum nitidum / Subasociación de Arcytophyllum nitidum Physiognomy: Shrubs and dwarfed trees dominate (up to 2 m, 20-40% cover); with a presence of tall stem rosettes of up to 3.5 m.
Composition and syntaxonomy: The subassociation is represented in 15 line-intersect surveys with a total of 50 vascular species. Diagnostic species are the same as the association as well as Ugni myricoides and Rubus acanthophyllos.
Rhynchospora guaramacalensis also being a further diagnostic species. In the
vegetation of this subassociation a ground layer of high bryophyte cover is common and comprised mainly Sphagnum sparsum and S. meridense. Other common species are Breutelia squarrosa, Campylopus flexuosus, C. nivalis,
Scapania portoricensis, Herbertus sp. (2980), Plagiochila tabinensis and other
species of Plagiochila and Frullania. Epiphytic bryophytes are also present on the smaller trunks. Some lichens, such as Cladia aggregata and Cladonia squamosa, can be found in the ground layer or over rocks. Peltigera neopolydactyla is found also on the dry leaves of Blechnum schomburgkii.
Some variants may also be distinguished for this subassociation, one variant characterized with a dominance of Rhynchospora guaramacalensis and a greater
presence of Ruilopezia paltonioides; the other variant dominated by Rhynchospora
macrochaeta.
Ecology and distribution: The shrubs of the subassociation typicum are located at altitudes of 2850-3040 m, at the base of convex slopes, with slopes between 10-37 degrees. Soils are 17-75 cm deep and consist of sandy, loam-sandy to silt-sandy textures, with dark brown grayish colours and pH of 3.4-4.5 in the upper layers.
Figure 3.4. Physiognomy of the vegetation of the association Disterigmo acuminatum - Arcytophylletum nitidum subass. pentacalietosum cachacoensis (L46a 3080 m). An: Arcytophyllum nitidum; Bs: Blechnum schomburgkii; Ch: Cortaderia
hapalotricha; Cm: Cybianthus marginatus; Da: Disterigma acuminatum; Dm: Daucus montanus; El: Elaphoglossum lingua; Ga: Gaultheria anastomosans; Gh: G. hapalotricha; Gm: Geranium stoloniferum; Ho: Hesperomeles obtusifolia; Hp: Hypericum paramitanum; Lg: Libanothamnus griffinii; Mm: Melpomene moniliformis; Pc: Pentacalia cachacoensis; Pp: Pernettya prostrata; Vc: Vaccinium corymbodendron.
HYPERICO CARDONAE –XYRIDION ACUTIFOLIAE Cuello & Cleef 2009
Typus: Cortaderio hapalotrichae -Hypericetum juniperinum
Hypericum cardonae - Xyris subulata var. acutifolia alliance / Alianza de Hypericum cardonae y Xyris subulata var. acutifolia
Physiognomy: This alliance includes zonal open grass páramo, with a high proportion of rosettes, whitin a variable density matrix of tussock grasses and bamboos. The presence of a few species of shrubs and dwarf trees varies from total absence to extreme densities.
Composition and syntaxonomy: This alliance is defined on the basis of 53 line-intersect surveys represented by 64 vascular species. Diagnostic species are: Xyris
subulata var. acutifolia and Hypericum cardonae. Although less frequent, Ruilopezia viridis is also a diagnostic occurance. The dwarf tree species Hypericum juniperinum is present in this alliance, present at very variable densities
among the different associations. The most important species, in sequence of cover, are: Ruilopezia lopez-palacii, Cortaderia hapalotricha, Chusquea
angustifolia, Geranium stoloniferum, Lycopodium clavatum subsp. contiguum, Hypericum juniperinum, Xyris subulata var. acutifolia, Pernettya prostrata, Rhynchospora guaramacalensis, Jamesonia imbricata, Puya aristeguietae, Libanothamnus griffinii, Rhynchospora macrochaeta, Disterigma acuminatum,
and Chusquea tessellata, among others.
This alliance contains three associations, Puyo aristeguietae-Ruilopezietum lopez-palacii; Cortaderio hapalotrichae -Hypericetum juniperinum; and Rhynchosporo gollmerii-Ruilopezietum jabonensis.
Ecology and distribution: The vegetation of the associations of the alliance of Hypericum cardonae and Xyris subulata var. acutifolia can be found between 2820
and 3060 m, located over ample extensions or forming small patches, on convex or concave slopes between 5 and almost 50 degrees.
3. Cortaderio hapalotrichae –Hypericetum juniperinum Cuello & Cleef 2009 Typus: Rel. No. 45 (Cuello L25b). Table 3.1. Figure 3.5. Photo 3.2
Cortaderia hapalotricha - Hypericum juniperinum shrub-grass páramo / Páramo de
arbustal-pajonal de Cortaderia hapalotricha e Hypericum juniperinum
Physiognomy and composition: Páramo vegetation with low density and diversity of shrubs and dwarf trees in the upper layer. Leptophyllous dwarf treelets of Hypericum juniperinum, 0.8-1.5 (2) m, 20-25% cover, with slender twigs and canopies oriented in the wind direction are noticeable. A dense grass layer is present at 10-60 cm in height, dominated by tussock grasses and small shrubs with some rosettes. The ground layer is dominated by Geranium stoloniferum and a variable cover of mosses and lichens. Rocky outcrops and areas of bare ground are common. In the upper layer, the dominance of Hypericum juniperinum is particularly noteworthy, together with a few other species of small trees like
Hesperomeles obtusifolia, Arcytophyllum nitidum and Chaetolepis lindeniana. In
the medium layer common Hypericum paramitanum grows among Chusquea
angustifolia bamboos, Cortaderia hapalotricha and Rhynchospora guaramacalensis tussock grasses. There are also the prostrate shrubs Disterigma acuminatum and Pernettya prostrata. Among the ground rosettes Puya aristeguietae and Ruilopezia lopez-palacii are more frequent and abundant, Ruilopezia jabonensis and R. viridis are occasionally present. In the herbaceous
layer Orthrosanthus acorifolius, Hypericum cardonae, Jamesonia imbricata,
Daucus montanus, Hieracium avilae and Lycopodium clavatum subsp. contiguum
are present, among others. In narrow valleys and humid areas, dense carpets of
over rocks and bases of trunks, such as Cladia aggregata, Cladonia squamosa, C.
andesita, C. pyxidata, C. arcuata, Jamesoniella rubricaulis, Herbertus juniperoides, Breutelia squarrosa, Plagiochila spp. (2961), Campylopus insignis
and, Riccardia spp. (2965). In these conditions, individuals of Hypericum
juniperinum and Chusquea angustifolia are found to reach their greatest heights of
up to 2-2.5 m.
Syntaxonomy: This association is defined on the basis of 25 line-intersect surveys containing 50 vascular species. Diagnostic species are Cortaderia hapalotricha,
Geranium stoloniferum and Hypericum juniperinum. Orthrosanthus acorifolius is
also diagnostic. Two subassociations are recognised, the subassociation typicum and that of disterigmetosum acuminatum.
Ecology and distribution: The association Cortaderio hapalotrichae -Hypericetum juniperinum is widely distributed between 2820 to 3060 m covering the entire upper ridge of Páramo of Guaramacal and Páramo El Pumar. The vegetation of this associaction extends over convex slopes with inclinations of 5 up to almost 50 degrees on hilltops or slope ridges exposed to wind. Patches of this vegetation additionally located on slope bases, concave sloping ground, or at the bottom of small valleys with slopes of 7-23 degrees.
The soils are variable in depth, 9-115 cm, with predominantly sandy textures, (sandy-loam, sand silt, silt-sandy, loam-sandy), pH 3.3-4.2 and dark grayish brown colors in the upper layers.
Figure 3.5. Physiognomy of the vegetation association Cortaderio hapalotrichae - Hypericetum juniperinum (L9a, 2910 m). Páramo El Pumar. At: Ageratina
theifolia; Ch: Cortaderia hapalotricha; Cha: Chusquea angustifolia; Chl: Chaetolepis lindeniana; Ga: Gaultheria anastomosans; Gm: Geranium stoloniferum; Hc: Hypericum cardonae; Hj: Hypericum juniperinum; Lc: Lycopodium clavatum subsp. contiguum; Oa: Orthrosanthus acorifolius; Pp: Pernettya prostrata; Rl: Ruilopezia lopez-palacii; Rm: Rhynchospora macrochaeta; Vc: Vaccinium corymbodendron.
Photo 3.2. Landscape of Páramo El Pumar in the surrounding areas of Laguna El Pumar, 2880–2950 m, Ramal de Guaramacal, Andes, Venezuela.
Cortaderio hapalotrichae –Hypericetum juniperinum 3.1. subassociation typicum Cuello & Cleef 2009 Typus: Rel. No. 45 (Cuello L25b). Table 3.1
Composition: This subassociation is represented in 12 line-intersect surveys with a total of 37 vascular species. The diagnostic species are the same as for the association. Orthrosanthus acorifolius, Xyris subulata var. acutifolia and
Hypericum cardonae are also diagnostic in the herb layer. The presence of Calamagrostis sp. A, Paepalanthus pilosus and Carex bonplandii, as well as some
cryptogams like Breutelia rhythidioides, Frullania sp. (2976), Cladia aggregata and Cladonia isabellina are distinctive. Diagnostic also is the absence of
Arcytophyllum nitidum.
Ecology and distribution: Vegetation belonging to this subassociation was observed at altitudes of 2890-3050 m, at the tops of hills and on convex slopes of low inclination (8-21 degrees), generally with S, SE, NE exposition. The soils are shallow, 9-30 cm in depth, on outcrops of bedrock, with sandy textures, dark grayish brown colours and pH in the range 3.3-4.2 in the upper layers. In this subasociation, shrub communities (1.5 up to 2 m), located in wind-protected areas at the base of the slopes, or along and to the base of small valleys with gently slooping ground (8-16 degrees), are also included. Soils are sandy-loam in texture, dark brown grayish or gray dark in colour and pH from 3.8 to 4.1 in the upper layers. Soil depth is 60 to 115 cm.
