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

Cuello Alvarado, N.L.

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Citation for published version (APA):Cuello Alvarado, N. L. (2010). Flora, vegetation and ecology in the Venezuelan Andes: a case study of Ramal deGuaramacal. 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. 2009

3.1 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;

Llambi et 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.

The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities

67

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 (Cleef 1981; Acosta-Solís 1984; 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).

Flora, vegetation and ecology in the Venezuelan Andes

68

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 & Cleef 2009a, 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 (9

o 12‟ 45.6” N; 70

o 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).


69

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


70

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).


71

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. RUILOPEZIO LOPEZ-PALACII – CHUSQUEETALIA ANGUSTIFOLIAE Cuello & 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


72

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.


73

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 1Gaultheria 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 . 4Ageratina theifolia . . . . . . . . . . 2 1 . 1 . . . . 1 . . . 2 . . . . . . . . . . 3 . . . .Galium hypocarpium . . . . . . . . . . . . . . . . . . . . . 1 . 2 . . . . . . . . . . . . . .Polypodium funckii . . . . . . . . . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . 1Eriosorus 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 2Hypericum 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 . 3Neurolepis 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 juniperinumHypericum juniperinum . . . . . . . . . . 2 . . . . . . . . 2 . . 1 . . . . . . . . . . . 4 1 1 .Orthrosanthus acorifolius . . . . . . . . 1 . . . . . . . . . . . . . 2 . . . . . . . . . . . . . 1 .Calamagrostis sp. A . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paepalanthus pilosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4. Puyo aristeguietae - Ruilopezietum lopez-palaciiPuya 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. Disterigmo acuminatum - Arcytophylletum nitidum1. Ruilopezio - Neurolepidetum glomeratae


74

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


75

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. Disterigmo acuminatum - Arcytophylletum nitidum1. 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 4Chusquea 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 5Lycopodium clavatum subsp. contiguum 1 . . 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 3Ruilopezia lopez-palacii . 2 . 2 . 4 3 . 3 . . . 4 . . 1 . 1 2 . . 2 1 4 4 4 4 . . 5 4 . 1 4 4 2 2 3Geranium stoloniferum . . . . . . . . . . . . . 1 1 . 2 3 . 5 4 . 4 . 2 . . . 1 2 . . . 2 4 1 3 2Pernettya 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 1Rhynchospora 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 1Jamesonia 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 . . 2Sphagnum sparsum . . . . . 5 6 . . . . . . . . . . . . . . . . . 6 . . . . 3 . . 6 5 5 4 . 6Hieracium 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 00 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 10 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 10 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



76

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


. . . . . . . . . . . . . . . . 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 . 51 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 51 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 10 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 10 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 10 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 10 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 convex2 concave

Slope shape:


77

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


78

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.


79

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.


80

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.


81

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


82

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.


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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


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,


84

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


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


85

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.


86

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

Sphagnum sparsum and a diversity of lichens and bryophytes are present growing


87

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.


88

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.


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Cortaderio hapalotrichae – Hypericetum juniperinum

3.2. subassociation disterigmetosum acuminatum Cuello & Cleef 2009

Typus: Rel. No. 56 (Cuello L23b). Table 3.1, Figure 3.6

Subassociation of Disterigma acuminatum / Subasociación de Disterigma acuminatum.

Physiognomy and composition: The physiognomy and species composition is in

agreement with that of the association.

Syntaxonomy: This subassociation is represented in 12 line surveys with 36

vascular species. Diagnostic species are Arcytophyllum nitidum, in the shrub layer,

as well as Rhynchospora guaramacalensis and Disterigma acuminatum, in the

underbrush.

Ecology and distribution: The subassociation disterigmetosum acuminatum is

found at altitudes from 2820 to 3060 m, on the convex and steep slopes (5 to

almost 50 degrees) of hilltops, edges and other wind exposed areas. The soils are

mostly shallow, 13-41 (86) cm, in depth; consisting of sandy, dark coloured,

textures with small fragments of quartz, having pH from 3.6 to 4.2 in the upper

layers.

Figure 3.6. Physiognomy of the vegetation of the association Cortaderio hapalotrichae -

Hypericetum juniperinum; subass. disterigmetosum acuminatum (L23b, 3030

m) An: Arcytophyllum nitidum; Ch: Cortaderia hapalotricha; Cha Chusquea

angustifolia; Da: Disterigma acuminatum; Gm: Geranium stoloniferum; Ho:

Hesperomeles obtusifolia; Hj: Hypericum juniperinum; Ji: Jamesonia

imbricata; Lc: Lycopodium clavatum subsp. contiguum; Ng: Nertera

granadensis; Pp: Pernettya prostrata; Rj: Ruilopezia jabonensis; Rm:

Rhynchospora macrochaeta; Vc: Vaccinium corymbodendron.


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4. Puyo aristeguietae – Ruilopezietum lopez-palacii Cuello & Cleef 2009

Typus: Rel. No. 65 (Cuello L10b). Table 3.1, Figure 3.7, Photo 3.3

Puya aristeguietae - Ruilopezia lopez-palacii grass páramo / Pajonal del páramo con

rosetas de Puya aristeguietae y Ruilopezia lopez-palacii

Physiognomy: Páramo vegetation with great abundances of ground and stem

rosettes with dominance of tussock grasses and some bamboos. The layer of big

Puya aristeguietae and Ruilopezia lopez-palacii rosettes, (terminal inflorescences

up to 1.5-2.5 m) attains 30-40% of cover. A layer of tall tussock grasses reaches up

to 90-125 cm with a cover of 35-45%. Small rosettes and other tussocky monocots

attain 45 cm. Additionally, a few low shrubs of 55-60 cm tall, 5-10% cover are

present. The ground layer (4-10 cm) consists of prostrate herbs and some

bryophytes. The presence of a few outcrops of rock (1.3 m) covered by abundant

lichens and bryophytes is common.

Photo 3.3. Páramo vegetation of the association of Puyo aristeguietae - Ruilopezietum

lopez-palacii, at ~2850 m in Páramo de Guaramacal, Ramal de Guaramacal,

Andes, Venezuela.

Composition and syntaxonomy: This association is defined on the basis of 17

line-intersect surveys with 45 vascular species. Diagnostic species are Ruilopezia

lopez-palacii, Puya aristeguietae and Rhynchospora guaramacalensis. The

dominant grasses in this association are Cortaderia hapalotricha (20 - 90 cm), and

the bamboo Chusquea angustifolia (50-125 cm), followed by others, such as:

Chusquea tessellata, Festuca guaramacalana, and Rhynchospora

guaramacalensis. Calamagrostis bogotensis and C. planifolia are common

species, but conspicuous only when fertile at the beginning of the rainy season.

Among the herbs Castilleja fissifolia, Daucus montanus, Hypericum cardonae,

Hieracium avilae and Jamesonia imbricata are common. Also present are prostrate

herbs like Geranium stoloniferum and Lycopodium clavatum subsp. contiguum as

well as small cushions of Oreobolus venezuelensis and Xyris subulata var.

acutifolia. Among the bryophytes Breutelia rythidioides, small cushions of

Campylopus subjugorum, and Herbertus pensilis as well as Campylopus richardii

growing over rocks are distinguished. Isolated and dispersed individuals of shrubs

or dwarf trees 1-1.5 (2.5) m, like Bejaria aestuans, Disterigma alaternoides,


91

Hypericum juniperinum, H. paramitanum, Ugni myricoides, and Vaccinium

corymbodendron are occasionally present.

Ecology and distribution: The open páramo vegetation of the association of Puya

aristeguiate and Ruilopezia lopez-palacii extends over large surfaces of Páramo de

Guaramacal between 2800-3040 m. It is present both on convex and concave

slopes varying between 5-18 degrees. The soils are comparatively deep, 30-120

cm, with sandy, sand-loam to silt-loam textures of brown-grayish color and pH

from 3.6 to 4.1 in the upper layers.

Figure 3.7. Physiognomy of the vegetation of the association of Puyo aristeguietae -

Ruilopezietum lopez-palacii (L10b, 2840 m). Bs: Blechnum schomburgkii; Ch:

Cortaderia hapalotricha; Cha: Chusquea angustifolia; Da: Disterigma

acuminatum; Dal: D. alaternoides Ji: Jamesonia imbricata; Lc: Lycopodium

clavatum subsp. contiguum; Md: Myrsine dependens; Ov: Oreobolus

venezuelensis; Pa: Puya aristeguietae; Pp: Pernettya prostrata; Rle:

Rhynchospora lechleri; Rm: Rhynchospora macrochaeta; Rsp: Rhynchospora

sp.; Rl: Ruilopezia lopez-palacii; Um: Ugni myricoides.

5. Rhynchosporo gollmerii – Ruilopezietum jabonensis Cuello & Cleef 2009

Typus: Rel. No. 82 (Cuello L14b). Table 3.1, Figure 3.8, Photo 3.4

Ruilopezia jabonensis - Rhynchospora gollmeri grass páramo / Pajonal de páramo con

Ruilopezia jabonensis y Rhynchospora gollmeri

Physiognomy: Low bunchgrass páramo with a high density of small ground

rosettes, cushion grasses and the presence of a few bamboos. Shrubs are absent

and the dominating silvery-leaved rosette species is Ruilopezia jabonensis. The

upper layer is composed of dispersed and low Chusquea angustifolia bamboo

clumps and bunches of Cortaderia hapalotricha of around 40-50 cm in height with

5 to 20% cover. The layer of rosettes reaches about 20-30 cm in height, covering

approximately 65%. There is a layer of small tussock and cushion grasses of up to


92

10 cm in stature and 30-40% cover. An open and discontinuous ground layer (2-3

cm) consists of mosses and small prostrate herbs (1%). The presence of rocks

outcrops (1%), bare ground and senescent material (3%) after fire is common.

Composition and syntaxonomy: The association is represented by 11 line-

intersect surveys with 22 vascular species.

The diagnostic species are: Ruilopezia jabonensis, Rhynchospora gollmeri,

Isidrogalvia robustior and Gentianella nevadensis. Other species with lower

density and cover are small herbs, like: Carex bonplandii, Geranium stoloniferum,

Hypericum cardonae, Lycopodium clavatum subsp. contiguum and Pernettya

prostrata, the grasses Calamagrostis planifolia and Polypogon elongatus together

with the terrestrial orchid Pterichis multiflora. In the ground layer the mosses

Campylopus richardii, Rhacocarpus purpurascens and Sematophyllum swartzii,

the lichens Cladia aggregata, species of Cladonia, as well as Rimelia reticulata

growing over the rocks, are present.

Photo 3. 4. Páramo vegetation of the association Rhynchosporo gollmerii - Ruilopezietum

jabonensis at ~2950 m in Páramo El Pumar, Ramal de Guaramacal, Andes,

Venezuela.

Ecology and distribution: The vegetation of Rhynchosporo gollmerii -

Ruilopezietum jabonensis is always located at altitudes superior to 2900 m.

Generally, it forms small patches, on concave slopes, or in small depressions, on

gently slooping ground (11-28 degrees). The vegetation of this association is in

downslope contact with that of the association of Puyo aristeguietae–

Ruilopezietum lopez-palacii var. Chusquea tessellata and upslope with the

association of Cortaderio hapalotrichae - Hypericetum juniperinum. It also borders


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the azonal vegetation association of Carici bonplandii - Chusqueetum angustifoliae

(Cuello & Cleef, 2009c.). The soils are of variable depth, 18-115 cm (average 49.6

cm), and are of sandy, sand-loam to sand-silt-loam texture, of gray and light colour

and of pH from 3.8 to 4.2 in the upper layers.

Figure 3.8 Physiognomy of the vegetation of the association Rhynchosporo gollmerii -

Ruilopezietum jabonensis (L14b, 2960 m). Ch: Cortaderia hapalotricha; Cha:

Chusquea angustifolia; Gm: Geranium stoloniferum; Ha: Hieracium avilae;

Hc: Hypericum cardonae; Ir: Isidrogalvia robustior; Lc: Lycopodium

clavatum subsp. contiguum; Rm: Rhynchospora macrochaeta; Rg:

Rhynchospora gollmeri; Rj: Ruilopezia jabonensis; Rl: Ruilopezia lopez-

palacii; Xs: Xyris subulata.

Flora diversity and composition

A total of 91 vascular plants, 33 species of bryophytes and 20 species of lichens

have thus far been documented from fifty 10 m-line intercept transects in zonal

páramo vegetation in Páramo de Guaramacal, Ramal de Guaramacal. The vascular

plants include 49 species belonging to 36 genera and 18 families of dicots; 24

species, 15 genera and 8 families of monocots and 18 species, 12 genera and 9

families of ferns. All plant species recorded in the studies of páramo vegetation

from Ramal de Guaramacal are listed in Appendix 4. It is expected that ongoing

sampling will yield further other bryophyte and lichen species.

Table 3.2. Most diverse plant families and genera in zonal paramo of Ramal de Guaramacal,

Venezuela.

FAMILY

#

GENERA # SPP GENERA # SPP

ASTERACEAE 6 13 Hypericum 4

ERICACEAE 7 10 Rhynchospora 4

POACEAE 4 7 (+3 indets) Ruilopezia 4

CYPERACEAE 3 6 Melpomene 3 (+2 indets)

CLUSIACEAE 1 4 Gaultheria 3

MELASTOMATACEAE 3 3 Hymenophyllum 3

MYRSINACEAE 2 3 Pentacalia 3

ROSACEAE 2 3

RUBIACEAE 3 3

GRAMMITIDACEAE 2 3 (+2 indets)

BROMELIACEAE 2 3


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Table 3.2 presents the most speciose families and genera for the páramo vegetation

of Ramal de Guaramacal based on the line-intersect data of this study. Asteraceae

and Ericaceae are the most speciose families followed by Poaceae and Cyperaceae.

The most diverse genera are Ruilopezia in the Asteraceae, Rhynchospora in the

Cyperaceae and Hypericum in the Clusiaceae. The flora diversity and most diverse

families for each vegetation type are presented in Table 3.3. Diversity decreases

from the most diverse shrubpáramo association of Disterigmo - Arcytophylletum

to the open grasspáramo of Rhynchosporo gollmerii - Ruilopezietum jabonensis.

Table 3.3. Flora diversity and most diverse families for each páramo vegetation association

found in Ramal de Guaramacal, Venezuela.

Life forms and growth forms

Species number for each life and growth forms for each vegetation association

registered from the line-intersect data from the páramo vegetation of Ramal de

Guaramacal are presented in Table 3.4 (a and b, respectively).

Generally the most representative life form in terms of both number of species and

cover in the study area are the phanerophytes, especially of the micro-

phanerophytic type, followed by hemicryptophytes of caespitose habit.

The growth forms with the highest species richness are upright shrubs, represented

mainly by members of the Clusiaceae, Ericaceae, Rubiaceae and Asteraceae

families, followed by tussock plants of the Poaceae, Cyperaceae, Xyridaceae and

Iridaceae families. The shrubpáramo association of Disterigmo - Arcytophylletum

shows the greatest diversity of growth forms and species.

Ordination analysis

The standard canonical coefficients as well as the intra- or interset variables (Ter

Braak 1986) (Table 3.5) show that the first CCA axis is mostly related to slope

angle (negative relationship), and the second CCA axis to altitude. This means that

slope angle and altitude are significantly related to species composition in the

zonal páramo vegetation, and appear more important than other variables such as

pH, and soil depth and humus thickness.

Association

#

Families # spp Most diverse families

1. Ruilopezio paltonioides -

Neurolepidetum glomeratae 20 41

Ericaceae (7), Asteraceae (5), Clusiaceae,

Cyperaceae, Myrsinaceae and Poaceae (3) 2. Disterigmo acuminatum -

Arcytophylletum nitidum 27 61

Ericaceae (8), Asteraceae (6), Clusiaceae

(4)

3.Cortaderio hapalotrichae - Hypericetum juniperinum

22 50 Asteraceae (8), Cyperaceae, Ericaceae, Poaceae (5), Clusiaceae (4)

4. Puyo aristeguietae -

Ruilopezietum lopez-palacii 22 45

Asteraceae, Ericaceae and Poaceae (6),

Cyperaceae (5) 5. R. gollmeri - Ruilopezietum

jabonensis 13 22

Cyperaceae (4), Asteraceae and

Clusiaceae (3)


95

Table 3.4. Number of species for life forms (a) and growth forms (b) for each vegetation

association registered from line-intersect data from páramo vegetation of Ramal

de Guaramacal. 1. Ruilopezio paltonioides - Neurolepidetum glomeratae; 2.

Disterigmo acuminatum - Arcytophylletum nitidum; 3. Cortaderio hapalotrichae

- Hypericetum juniperinum; 4. Puyo aristeguietae - Ruilopezietum lopez-palacii;

5. Rhynchosporo gollmerii - Ruilopezietum jabonensis.

(a) Life forms Number of species by vegetation type

(1) (2) (3) (4) (5) Total spp

phanerophyte 6 5 3 3 1 8

microphanerophyte 6 11 8 9 1 15

nanophanerophyte 2 4 4 2 2 4

phanerophytic lignified grass 1 1 1 2 1 2

rosullate phanerophyte 5 5 7 5 2 8 hemicryptophyte 4 9 5 5 3 13

caespitose hemicryptophyte 6 7 12 12 8 16

climbing hemicryptophyte 1 3 - - - 3 chamaephyte - - 1 - - 2

frutescent chamephyte 7 6 5 5 2 8

reptant chamaephyte - 3 9 4 2 2 Epiphyte - 1 2

Total phanerophytes 20 26 23 21 7 37

Total hemicriptophytes 11 19 17 17 11 32 Total chamaephytes 10 15 10 7 4 20

Total spp 41 61 50 45 22 91

Total life forms 10 12 10 9 9 13

(b)

Growth forms*

Number of species by vegetation type

(1) (2) (3) (4) (5)

Total

spp

upright shrubs 9 12 10 9 2 18

tussocks 7 8 13 13 8 17

erect herbs 6 9 5 7 3 14 dwarf trees 6 9 6 5 2 10

prostrate herbs 2 8 4 2 2 9

ground rosettes 2 3 5 3 2 6 prostrate shrubs 5 5 3 2 1 6

cushions - - 1 1 1 3

stem rosettes 3 3 3 3 1 3 trailing herbs 1 3 - - - 3

epiphitic herbs - 1 - - - 2

Total spp 41 61 50 45 22 91

* Adapted from Ramsay & Oxley, 1997 ad Hedberg & Hedberg, 1979

The ordination diagram of the first CCA axis against the second CCA axis with the

samples (transect lines) labeled by vegetation types (Fig. 3.9) shows a fairly good

separation of vegetation communities established on the basis of the

phytosociological table (Table 3.1). Vegtype 1 (Ruilopezio paltonioides -

Neurolepidetum glomeratae), and to a lesser degree Vegtype 2 (Disterigmo

acuminatum - Arcytophylletum nitidum), are separated from the others towards the

left, suggesting that these vegtypes are associated with higher slope angles.

Similarly, vegtypes 3, 5, and to a lesser degree 4, must have rather low values of

slope angles. Vegtype 4 separates well along CCA axis 2, which suggests that this

vegtype occurs at the lowermost positions along the slopes.


96

Table 3.5. Standard canonical coefficients and interset variables of CCA ordination axis for

páramo vegetation of Ramal de Guaramacal.

Figure 3.9. CCA ordination diagram of 91 vascular species recorded in 91 páramo

vegetation samples (labeled by vegetation types) in Ramal de Guaramacal,

Andes Venezuela. Vegtypes: 1. Ruilopezio paltonioides - Neurolepidetum

glomeratae; 2. Disterigmo acuminatum - Arcytophylletum nitidum; 3. Corta-

derio hapalotrichae - Hypericetum juniperinum; 4. Puyo aristeguietae –

Ruilopezietum lopez-palacii; 5. Rhynchosporo gollmerii - Ruilopezietum

jabonensis.

3.5 DISCUSSION

Phytosociological classification and methodological constraints

The phytosociological classification of zonal páramo vegetation of the Guaramacal

range resulted in a provisional order (Ruilopezio lopez-palacii - Chusqueetalia

angustifoliae prov.), two new alliances and five associations. Four new

Canonical Coefficients

Standardized Original Units

Variable Axis 1 Axis 2 Axis 3 Axis 1 Axis 2 Axis 3 S.Dev

1 Alt 0.135 0.389 -0.124 0.002 0.005 -0.002 0.774E+02

2 Slope angle -0.495 0.018 -0.059 -0.053 0.002 -0.006 0.940E+01

3 Soils depth -0.054 -0.012 -0.326 -0.002 0.000 -0.011 0.302E+02

4 pH 0.075 -0.100 -0.035 0.269 -0.361 -0.125 0.278E+00

5 Humus depth -0.054 -0.031 0.181 -0.006 -0.003 0.019 0.952E+01


97

subassociations are described for two associations, two for each. Some variants

have also been recognised. The zonal subpáramo plant communities of Ramal de

Guaramacal are summarized in Table 3.6.

A class cannot yet be defined on the basis of the Guaramacal relevés alone (Table

3.1) and the complete lack of data from other Chusquea angustifolia bamboo

páramo areas in the region and from elsewhere in Venezuela and Colombia.

Regional comparison, therefore, presently remains impossible. However, in order

to evaluate the pattern of associated plant species and their dominancy a

comparison with zonal Chusquea tessellata páramos of the Colombian Cordillera

Oriental (Cleef 1981) has been undertaken (Table 3.7). The relevés are from the

Colombian data set of the second author. Typical Sphagnum bogs with Chusquea

tessellata have been avoided.

Inspection of Table 3.7 learns that zonal Chusquea angustifolia bamboo páramo of

Guaramacal shares about half of the vascular genera with the zonal Chusquea

tessellata bamboo páramo of Colombia. Most important, however, is that there is

no general agreement in generic pattern between both bamboo páramos, except for

Chusquea. Apparently the Guaramacal bamboo páramo has more woody species,

also because of its low altitude. The Colombian relevés span an altitudinal range

between about 3200 and 4040 m. In conclusion, the Chusquea angustifolia

bamboo páramo of Guaramacal represents a proper vegetation type not studied

elsewhere.

The páramo vegetation of the Guaramacal study area has been described on the

basis of a relatively low number of relevés (ninety one 5 m-line surveys).

Sampling effort in páramo areas of Ramal de Guaramacal was concentrated in the

by road accessible sector of Las Antenas of Páramo de Guaramacal. Las Antenas

area evidences most different physiognomic formations in relatively close

proximity, and with a larger altitudinal range (2820~3130 m). Only a limited

number of surveys were conducted in the remote areas of Páramo El Pumar, where

the zonal vegetation appears more homogeneous over large areas. There, little

variation in vegetation types, with a constant species composition, was observed

over a shorter altitudinal range (2880~2990 m).

As indicated in the methods section, line transects were laid out in apparently

homogeneous and representative páramo vegetation patches. A line of 10 m was

employed. The classical Zürich Montpellier approach uses plots of different size

according to the structure and diversity of the vegetation. The minimum area has to

be established for the different vegetation types (see also Westhoff & van der

Maarel 1973; Cleef 1981). In the case of the zonal páramo of Ramal de

Guaramacal, with its limited total of vascular species and few different páramo

vegetation types, the line of 10 m has always been employed for documenting the

presence of different species under the line with their cover abundance.

To our surprise, no apparent discrepancies appeared in the TWINSPAN analysis

and the final classification of the páramo plant communities. We believe a similar

result would appear when plot sampling has been used. This method has, in fact,

been chosen by the first author following a 1990 field course in the savannas of

Bolivia organized by Tratado de Cooperación Amazónica (Cuello et al. 1991). The


98

method of line-intercept transects has been widely used in vegetation ecology

since the papers by Canfield (1941) and McIntyre (1953). This method has been

tested for laboratory teaching (Cummings & Smith 2000; Kercher et al. 2003). It is

possible that the line intercept technique used here could yield higher cover

estimates but lower species richness estimates than the plot method, since plots (of

generally 5 m x 5 m) would cover larger area than single 10 m lines.

We consider, however, the resulting páramo classification is clearly visible for

Páramo de Guaramacal. Páramo communities at association level may be

representative of most páramo areas of Ramal de Guaramacal. A greater sampling

effort, in a balanced way, over the study area would be necessary to refine the

classification into infra association level.

Table 3.6. Presence degree table of zonal subpáramo plant communities of Ramal de

Guaramacal. I (0–20 %), II (21–40 %), III (41–60 %), IV (61–80 %) and V (81–

100 %). Community group 1 2 2.1 2.2 3 3.1 3.2 4 5

Number of relevés 10 28 13 15 25 12 13 17 11

1. Ruilopezio paltonioides - Neurolepidetum

glomeratae

Ruilopezia paltonioides IV II II I I I I I .

Disterigma alaternoides II II I . . . . I . Nertera granadensis II . . . I I II . .

Pentacalia greenmaniana I . . . . . . . .

Sphyrospermum buxifolium I . . . . . . . . Cybianthus laurifolius I . . . . . . . .

2. Disterigmo acuminatum - Arcytophylletum

nitidum

Disterigma acuminatum II V V V III . V II .

Gaultheria hapalotricha I III IV III I I I . .

Arcytophyllum nitidum I III II IV II . III . I Ageratina theifolia . II II I I II . I .

Galium hypocarpium . I I I . . . . .

Polypodium funckii . I I . . . . . . Eriosorus flexuosus . I I I . . . . .

Hymenophyllum myriocarpum . I I I . . . . .


Pentacalia cachacoensis . II IV I . . . . .

Vaccinium corymbodendron I II III I II II II I . Melpomene moniliformis . II III . I I . . .

Gaultheria anastomosans . II III I I II I I .

Themistoclesia dependens . I I I . . . . . Hesperomeles sp. . I I . I I . . .

Huperzia amentacea . I I . . . . . .

2.2. typicum Ugni myricoides II II I III I . I I .

Rubus acanthophyllos . II . II . . . . .

Ilex guaramacalensis . I . I . . . . Valeriana quirorana . I . I I . I I .

Gaultheria erecta . I . I . . . . .

Hymenophyllum sp. . I . I . . . . . Melpomene flabeliformis . I . I . . . . .

Melpomene xiphopteroides . I . I . . . . .

Polypodium sp. . I . I . . . . .

HYPERICO PARAMITANUM –

HESPEROMELETION OBTUSIFOLIAE


99

Community group 1 2 2.1 2.2 3 3.1 3.2 4 5

Blechnum schomburgkii V IV IV IV . . II II .

Hypericum paramitanum V V V IV I II I II I Neurolepis glomerata IV II III II I . I I I

Cybianthus marginatus II II IV I I I I I .

Hesperomeles obtusifolia II III IV II III I IV . . Libanothamnus griffinii II I II I I . II I .

Elaphoglossum cf. lingua I II I II . . . I .

Puya sp. II I I I . . . . . Miconia tinifolia I I I . . . . . .

Muehlenbeckia tamnifolia I I I . . . . . .

Epidendrum frutex I I I . . . . I .

Myrsine dependens . I II I . . . I .

Diplostephium obtusum I I I I I II I . .

Rhynchospora sp. I I I . I I I I .

3.Cortaderio hapalotrichae- Hypericetum

juniperinum

Hypericum juniperinum . II II I V V V II II Orthrosanthus acorifolius . I I I II III I . .

Calamagrostis sp. A . . . . I II . . .

Paepalanthus pilosus . . . . I I I . . Greigia sp. . . . . I . I . .

4. Puyo aristeguietae - Ruilopezietum lopez-

palacii

Puya aristeguietae I I . I I . I IV .

Chusquea tessellata . . . . . . . II .

Castilleja fissifolia . . . . . . . II .

Festuca guaramacalana . . . . . . . I .

Monnina sp. . . . . . . I . Bejaria aestuans . . . . . . . I .

Rhynchospora lechleri . . . . . . . I .

Oreobolus venezuelensis . . . . . . . I II Festuca sp. . . . . . . . I .

Utricularia alpina . . . . . . I .

5. R. gollmeri - Ruilopezietum jabonensis

Ruilopezia jabonensis . . . . I I I . V

Rhynchospora gollmeri . . . . I . . . IV

Isidrogalvia robustior . . . . I . I I I Gentianella nevadensis . . . . . . . . I

Calamagrostis planifolia . . . . . . . . I

HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE Xyris subulata var. acutifolia . I . I III V II IV V

Hypericum cardonae I I I . III V II I III

Carex bonplandii . . . . I III I . . Ruilopezia viridis . . . . I I I I .

RUILOPEZIO LOPEZ-PALACII - CHUSQUEETALIA ANGUSTIFOLIAE

Cortaderia hapalotricha III V V V V V V V V

Chusquea angustifolia IV IV III V IV IV IV IV V

Lycopodium contiguum IV IV IV V V V V V V

Ruilopezia lopez-palacii III IV III IV III IV III V I Geranium stoloniferum . III III III IV IV V II IV

Pernettya prostrata V V IV V V V V V II

Rhynchospora guaramacalensis III II II III I . II III .

Rhynchospora macrochaeta I II I III III IV II II V

Jamesonia imbricata III II I III II I III IV II

Chaetolepis lindeniana II IV IV III III III II I . Daucus montanus I III II III II II II I .

Hieracium avilae . I I I I I II II I

Hymenophyllum trichomanoides . I I I I I I . . Hypericum sp. I I I . I I . . .


100

Table 3.7. Table of presence of the zonal Guaramacal Chusquea angustifolia bamboo

páramo associations combined with that of the zonal Chusquea tessellata

bamboo páramo community of the Colombian Cordillera Oriental based on

unpublished releves of the second author. The predominant genera are

underlined. I (0–20 %), II (21–40 %), III (41–60 %), IV (61–80 %) and V (81–

100 %)

.

Number of relevés 10 28 25 17 11 25

Association 1 2 3 4 5 Cord. Oriental

Colombia

Ageratina . II I I . .

Aragoa . . . . . I

Arcytophyllum I III II . I III

Azorella . . . . . I

Bartsia . . . . . V

Bejaria . . . I . .

Blechnum V IV . II . I

Breutelia I I I I IV

Calamagrostis . . I . I V

Campylopus I I I V

Carex . . I . . II

Castilleja . . . II . I

Castratella . . . . . I

Chaetolepis II IV III I . .

Chusquea IV IV IV V V V

Cortaderia III V V V V II

Cybianthus II II I I . .

Cyperus . . . . . I

Daucus I III II I . .

Diplostephium I I I . . I

Disterigma III V III II . II

Elaphoglossum I II . I . .

Epidendrum I I . I . .

Eriosorus . I . . . .

Eryngium . . . . . I

Espeletia . . . . . IV

Festuca . . . I . II

Galium . I . . . .

Gaultheria I III I .I .

Gentiana . . . . . II

Gentianella . . . . I IV

Geranium . III IV II IV I

Greigia . . I . . .

Halenia . . . . . I

Hesperomeles II III III . . .

Hieracium . I I II I I

Huperzia . I . . . I

Hydrocotyle . . . . . I

Hymenophyllum . I I . . .

Hypericum V V V III V I

Hypochaeris . . . . . III

Ilex . I . . . .

Isidrogalvia . . I I I .


101

Number of relevés 10 28 25 17 11 25

Association 1 2 3 4 5 Cord. Oriental

Colombia

Jamesonia III II II IV II I

Laestadia . . . . . I

Libanothamnus II I I I . .

Lycopodium IV IV V V V II

Lysipomia . . . . . I

Melpomene I II I . . .

Miconia I I . . . .

Monnina . . . I . .

Muehlenbeckia I I . . . .

Myrsine . I . I . .

Nertera II . I . . II

Neurolepis IV I I I I .

Niphogeton. . . . . . I

Oreobolus . . . I II III

Oritrophium . . . . . III

Orthrosanthus . I II . . .

Paepalanthus . . I . . I

Pentacalia I II . . . III

Pernettya V V V V II I

Plantago . . . . . I

Polypodium . I . . . .

Puya II I I IV . I

Rhacocarpus . . . . . II

Rhynchospora V IV V V V I

Rubus . II . . . .

Ruilopezia V IV V V V .

Scirpus . . . . . I

Sisyrinchium . . . . . I

Sphagnum III III I I III

Sphyrospermum I . . . . .

Themistoclesia . I . . . .

Ugni II II I I . .

Utricularia . . . . .

Vaccinium I II II I . I

Valeriana . I I I . I

Xenphyllum . . . . . I

Xyris . I III IV V .

Páramo flora composition and diversity

From a total of fifty 10 m-line intersect surveys, it was possible to register at least

48.2% from a total of 193 vascular species known to date, from páramo areas of

Ramal de Guaramacal. With a limited altitudinal span (2820-3130 m), the Páramo

de Guaramacal exceeds a total surface area of not more than 10 km2. Most species

are, in general, located in the lower part of the páramo belt. However, taking into

account the actual degree of isolation (presently separated ca. 30 km Southwest

and 35 km Northeast from the nearest páramo zones), the limited surface area and

altitudinal span, the presence of only some 200 vascular páramo species (alpha


102

diversity), compared to the number of 1544 vascular species reported from

Venezuelan páramos [1437 angiosperms species reported by Briceño & Morillo

(2002, 2006) plus 107 fern species reported by Luteyn (1999)] is quite

understandable. Judging from periglacial evidence in the Guaramacal páramo it is

clear that glaciation took place during the Last Glacial Maximum (LGM), and that

the páramo zone extended downslope. Connectivity to other páramos of the

Cordillera de Mérida was probably more functional during the LGM than is the

case today. Repeated isolation during interglacials in the past has triggered a

number of endemic species, and maybe even the highest species diversity of

Ruilopezia rosettes, thus far, reported in the Venezuelan Andes to date. Up to date,

about 50 endemic vascular species are known from Ramal de Guaramacal which

represent ca. 4% from a total of about 1400 vascular species.

Physiognomy: life forms and growth forms

Páramo vegetation of Ramal de Guaramacal is dominated mainly by woody

growth forms, particularly upright shrubs with bamboo groves and clumps, which

give an overall appearance of a mostly shrub páramo vegetation. Two out of five

associations are dominated by the presence of upright shrubs, and two out of the

three bunchgrass dominated associations, also contain a high number of shrub

species. The only grass páramo community almost devoid of shrubs is that of the

low diverse Rhynchosporo gollmerii - Ruilopezietum jabonensis. The only two

shrubby species registered in this association may be a consequence of sampling

near the border with the surrounding shrubby páramo of Cortaderio hapalotrichae -

Hypericetum juniperinum. The high relative humidity and the low altitudinal

range, coupled with the close proximity of the dwarf forests of the upper forest line

zone, may explain in part the dominance of shrubby growth forms in páramo

vegetation of Ramal de Guaramacal. From other extremely wet páramos the

predominance of shrubs has also been reported, e.g. the Biosphere reserve of

Podocarpus in South Ecuador (Bussmann 2002; Richter 2003; Becking et al. 2004

and the Tatamá páramo in the West Cordillera of Colombia (Cleef et al. 2005).

Phytosociological classification and environmental variables

Twinspan classification of Páramo de Guaramacal (Table 3.1) arranges vegetation

types in a sequence from shrub páramo to open páramo. This sequence could be

directly related to a decrease in temperature with increasing altitude. Additionally,

the CCA ordination analyses show that species composition in the zonal páramo

vegetation is foremost related to slope angle and altitude. On a later occasion

(Cuello in prep.) the results of the ordinations will be more detailed.

In the studied altitudinal range from 2800-3100 m in Páramo de Guaramacal, it is

generally observed that different vegetation types can be found occupying the

same altitude, with the exception of the grass páramo of Rhynchosporo gollmeri–

Ruilopezietum jabonensis, which is always found above 2900 m. Other vegetation

types, however, can be present above this altitudinal range; particularly, the shrub

páramo of Cortaderio hapalotrichae - Hypericetum juniperinum, which is always

present at the top of slopes.


103

In the sector surrounding Las Antenas area, as shown in Fig. 3.10, on North to East

slope expositions of Páramo de Guaramacal, and upslope the edges of the high

Andean dwarf forest association of Libanothamnetum griffinii (around 2800-3000

m) (Cuello & Cleef 2009a, Chapter 2), the ecotonic shrub páramo of Ruilopezio -

Neurolepidetum association is generally present on either convex or concave

slopes with relatively deep soils of predominantly loamy textures. Next, the grass

páramo of Puyo aristeguietae - Ruilopezietum lopez-palacii is found anywhere

from c. 2800 m to ~3040 m, alternating with the shrub páramo of the Cortaderio

hapalotrichae - Hypericetum juniperinum. Probably it belongs to the upper

subpáramo, but by burning incidences the original woody component has

decreased. The open grass páramo of Rhynchosporo gollmerii - Ruilopezietum

jabonensis follows in altitude to that of Puyo aristeguietae - Ruilopezietum lopez-

palazii. The lower grass páramo association is present predominantly on concave

areas with coarse sandy soils close to the upper sections of slopes. Finally, the

vegetation of Cortaderio hapalotrichae - Hypericetum juniperinum is present at the

top of the slope.

The effect of past disturbance, such as fires and the disruption of vegetation cover

during or after the trail construction and installation of the telecomunication

antennas, may explain the current distribution patterns of páramo vegetation in the

Antenas sector. There is a fragmentation of the high Andean dwarf forests (SARF),

evidenced by the current presence of some remnant islands surrounded by shrub

páramo and open páramo vegetation. The grass páramo of Puyo aristeguietae -

Ruilopezietum lopez-palacii seems to be a derived vegetation type from a past

burning of the apparently original and extensive Cortaderio hapalotrichae -

Hypericetum juniperinum shrub páramo, which currently occurs on the borders of

little valleys or near the top of slopes. The presence of a continuous cover of the

open páramo, with single-stemmed Hypericum juniperinum shrub (in fact a dwarf

tree) of the Cortaderio hapalotrichae - Hypericetum juniperinum, towards the

apparently pristine areas of Páramo El Pumar, at the West of the summit of Ramal

de Guaramacal, is indicative of a possible formerly more extensive presence in the

Las Antenas area. Both the Cortaderio hapalotrichae - Hypericetum juniperinum

and the Puyo aristeguietae - Ruilopezietum lopez-palacii associations share similar

species composition; the Cortaderio - Hypericetum being typically more speciose.

In Las Antenas area the vegetation of the Cortaderio - Hypericetum shows lower

species richness than in El Pumar area, and the páramo of the Puyo -

Ruilopezietum shows an absence, or very low presence of individuals of

Hypericum juniperinum shrubs.

On steeper South and Southwest slopes away from Las Antenas and along the

mountain ridge towards the West, the altitudinal sequence of vegetation types that

is contiguous upslope of the Libanothamnus griffinii dwarf forest, or that of

Gaultheria anastomosans - Hesperomeles obtusifolia (see Cuello & Cleef 2009a,

Chapter 2), is an alternation of shrub páramos of the Disterigmo - Arcytophylletum

association on concave or protected slopes, followed upwards by the shrub páramo

of the arcytophylletosum nitidum subassociation of the Cortaderio hapalotrichae -

Hypericetum juniperinum characteristic on steeper and wind exposed expositions.


104

Figure 3.10. Gradient SARF – Zonal páramo, 3000-3050 m, North of „Las Antenas‟, Ramal

de Guaramacal, Andes, Venezuela 1. Ruilopezio paltonioides - Neurolepide-

tum glomeratae (var.1.1 Disterigma alaternoides); A. Libano-thamnetum

griffinii; 3. Cortaderio hapalotrichae - Hypericetum juniperinum; 4. Puyo

aristeguietae - Ruilopezietum lopez-palacii; 5. Rhynchosporo gollmeri -

Ruilopezietum jabonensis.

In Páramo El Pumar, at c. 2.5 km West from Las Antenas, the open shrub páramo

of Cortaderio - Hypericetum abounds all over the altitudinal range from ~2880 to

3000 m. The continuity of the Cortaderio - Hypericetum dominated landscape of

Páramo El Pumar is interrupted with the presence of some (azonal) bogs around

glacial lakes (Cuello & Cleef, 2009c, Chapter 4). Shrub páramo of the Disterigmo

acuminatum - Arcytophylletum nitidum association is further present on concave,

or protected slopes, as well as high Andean forest patches of the Geissantho andini

- Miconietum jahnii on sites with apparent local variation in topography and soils

(Cuello & Cleef 2009a, Chapter 2). Open páramo of Rhynchosporo gollmerii -

Ruilopezietum jabonensis also occurs in small patches in depressions at borders or

near the top of slopes over 2900 m, but is always surrounded by the shrub páramo

of Cortaderio hapalotrichae - Hypericetum juniperinum.

Glacial morphology and páramo vegetation

Evidence of the last glaciation is apparent nearly everywhere on the around the

3000 m ridge of Ramal de Guaramacal. The summit zone is generally narrow but

slightly wider and highest near Las Antenas. The ridge in the area of Pumar is

widest with a few small glacial lake basins and terminal moraines. Here a large

glacier has been descending along the Llanos slope. Remnants of ground moraines

and periglacial sediments are found outside the area of inclinated bedrock which is

the most salient feature of the landscape. Roche moutonnée has also been locally

observed. During the Last Glacial Maximum (LGM), the páramo zone probably

extended to around 2000 m when interpolated from Laguna Pedro Palo from the

Andes near Bogotá (Hooghiemstra & Van der Hammen 1993). The snow and

glaciers would possibly have been restricted mainly to the ridge area; the


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Guaramacal páramo zone of the present. Slopes were too steep for the support of

snow and ice, which probably collected at the base of these steep slopes; covered

mainly today by upper montane and the subalpine dwarfed rain forests.

Looking at the páramo landscape of the Guaramacal ridge, we can observe that the

zonal vegetation of Cortaderio hapalotrichae - Hypericetum juniperinum is most

important in terms of the cover of the Guaramacal páramo (Photo 3.2). This open

shrubby vegetation also covers most of the rocky surfaces of Páramo de

Guaramacal with, in general, limited soil thickness ranging from between 5-10 to

115 cm. The vegetation of both associations of the alliance Hyperico paramitanum

- Hesperomeletion obtusifoliae are contiguous to the upper forest line, the humid

shrub páramo of Ruilopezio paltonioides - Neurolepidetum glomeratae association

is based on deeper soils (up to ca. 105 cm) and is closer to the UFL and the shrub

páramo of the association Disterigmo acuminatum - Arcytophylletum nitidum is

contiguous to that of the latter.

The nature of the large surface of exposed bedrock, and the climatic characteristics

of the top effect, mean this area cannot support subalpine forest or upper montane

rain forest, not even under a warming climate.

Comparison with other páramos

As detailed in the introduction, Chusquea bamboo páramos have not yet been

studied in Venezuela. They are distributed along the humid UFL on the Llanos

slope of the Venezuelan Andes. Páramo de Guaramacal is also part of this unit. It

is unknown if Chusquea bamboo páramos also occur along the UFL on the

Maracaibo slope of the Cordillera de Mérida. Although Chusquea angustifolia has

also been reported also from páramo areas in Zulia (Briceño & Morillo 2006) and

specimens collected from Perijá are listed in MBG W3Tropicos database.

On Avila and Naiguatá, Vareschi (1953, 1955) and Aristeguieta & Ramia (1951)

described Chusquea spencei bamboos from the Libanothamnus neriifolius

community (see also Steyermark & Huber 1978). Chusquea spencei has also been

reported in the páramos of Cendé, Jabón and Las Rosas in Trujillo-Lara states

border, North to Northeast of the Guaramacal range, as well as in Páramo El

Zumbador and Tamá in Táchira, and in Páramo Los Conejos (La Culata) near

Mérida (Monasterio 1980b). In humid areas of Páramo de Tamá, Bono (1996)

describes the presence of Chusquea formations (a „Chusqueetum‟ community of

Chusquea angustifolia and Ch. tessellata) along small streams. It seems that

Chusquea spencei prefers a drier páramo habitat (Monasterio 1980b) than

Chusquea angustifolia, which determines the aspect of the Páramo de Guaramacal.

In the Guaramacal páramo a few patches of Chusquea tessellata have also been

documented. Chusquea angustifolia is present close to the UFL along the Llanos

side of the Cordillera de Mérida and the Eastern Cordillera of the Andes in

Colombia: Páramo de Sumapaz representing thus far its southernmost distribution.

Chusquea angustifolia thrives in a clouded wet upper forest line habitat in

comparison to its high altitude adapted relative Chusquea tessellata, which is a

common species throughout the humid páramos of Colombia and Ecuador

extending southwards to Bolivia (Luteyn 1999; Clark 2000). Chusquea

angustifolia has smaller leaves but a greater density of leaves per branch than


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Chusquea tessellata. However, it is estimated that Chusquea angustifolia has an up

to three times greater leaf surface area than Chusquea tessellata. This factor may

also explain the dominance of Chusquea angustifolia in the wet Páramo de

Guaramacal, which is also at lower altitude than most other zonal bamboo páramos

as a function of top effect combined with the presence of bare rocky surfaces of

the Guaramacal ridge. The limited knowledge on the presence and composition of

Chusquea angustifolia bamboo páramos elsewhere, provides argument to rank the

order described here as provisional.

Species of Neurolepis bamboos are also highly indicative of wet environmental

conditions. Associated with Chusquea angustifolia, Neurolepis glomerata occurs

in the dwarf forests of the SARF-UMRF association of Gaultherio anastomosans -

Hesperomeletum obtusifoliae (Cuello & Cleef 2009) and in the zonal páramo of

Ruilopezio paltonioides - Neurolepidetum glomeratae association described here.

Neurolepis aristata is a low to tall bamboo occurring in association with Chusquea

tessellata in bamboo páramo or as groves in protected sites near the UFL (Cleef

1981; Bussmann 2002). In the Guandera summit area in northern Ecuador an

association of Neurolepis aristata bamboo vegetation was developed around 4000

m on the Amazon slope in an Espeletia pycnophylla - Calamagrostis effusa

bunchgrass páramo (Moscol & Cleef 2009a). The Podocarpus Park páramo in

South Ecuador is probably the world‟s most wet páramo. Bussmann (2002)

described a number of páramo communities from its northeastern extremity with

Neurolepis being present as the most dominant bamboo species: e.g.

Neurolepidetum laegaardii Bussmann 2002. A number of bamboo species of

Bussmann‟s Neurolepidion laegaardii alliance include: Chusquea tessellata,

Neurolepis weberbaueri, and further Chusquea loxensis, Ch. leonardiorum, Ch.

perligulata and Neurolepis nana. An association Neurolepidetum aristatae

Bussmann 2002 has also been described from this rain-swept páramo.

This is the first time that Chusquea angustifolia has been referred to in a

phytosociological context. Aside from the reference made by Bono (1996) in

Páramo de Tamá, we are not aware of the bamboo vegetation of this species

elsewhere or how this species interacts with the more common bamboo páramo

species, Chusquea tessellata.

The few clumps of Chusquea tessellata in Páramo de Guaramacal are supposed to

be relatively recent arrivals in a setting occupied entirely by Chusquea

angustifolia. Looking at the present-day distribution of Chusquea angustifolia, we

assume that other unnamed associations in UFL in Páramos of Lara-Trujillo,

Mérida, Táchira (Tamá), and Zulia (Perijá) where the species has been reported in

Venezuela (Clark 1990; Briceño & Morillo 2006), Arauca slope of Sierra Nevada

del Cocuy, Páramo de Pisba, Chingaza and Sumapaz among other localities on the

Llanos slope of the Colombian Eastern Cordillera are present.

From páramos of Trujillo-Lara states, Páramos Cendé, Jabón and Las Rosas to the

north easternmost Venezuelan Andes, Monasterio (1980b) described the shrub

páramo with a rosette community of Ruilopezia jabonensis or „Rosetal de

Ruilopezia jabonensis’, as the most important Andean páramo vegetation

“association” or community found within this area. This community was also

referred as the driest páramo area of the country, receiving scarcely 600 mm/year


107

rainfall at 3000-3400 m altitude. There, the Ruilopezia jabonensis vegetation

community is present over large open areas and is surrounded by woodland

communities of Libanothamnus neriifolius, and a shrubby bamboo páramo

community dominated by Chusquea spencei and the endemic Pentacalia

rigidifolia. According to Monasterio (1980b), the Ruilopezia jabonensis páramos

of the Trujillo-Lara state border are composed mainly of a high ground rosette

cover (50-60%): with Arcytophyllum caracasanum, Hypericum caracasanum and

H. laricifolium shrubs in addition to tussocks of Cortaderia nitida and

Orthrosanthus chimboracensis. Niño et al. (1997), in a brief quantitative páramo

vegetation survey utilising a 50 m line intersect transect in Páramo Cendé at 3200

m, studied a community dominated by Ruilopezia jabonensis, characterized by a

high cover of Chusquea angustifolia bamboo, and a prominent abundance and

diversity of bunchgrass species, such as Agrostis meridensis, Aristida sp.,

Cortaderia nitida, Danthonia secundiflora and an orchid species, Stenorrhynchos

vaginatum (Niño et al. 1997). The silvery monocarpic rosettes of Ruilopezia

jabonensis appear well adapted to higher elevations and drier conditions than those

in Guaramacal. This may explain its limited presence on only small patches of the

shrubless páramo of the Rhynchosporo gollmeri - Ruilopezietum jabonensis

association, occurring over well-drained coarse sandy soils, and restricted by the

lower altitude of Ramal de Guaramacal from 2900 up to 3100 m. None of the

bunchgrass companion species of the Ruilopezia jabonensis community of Páramo

Cendé reported by Niño et al. (1997) are present in Páramo de Guaramacal. The

presence of silvery rosettes is also curiously observed in disturbed páramos, e.g.

Espeletia schultzii in Mérida, Venezuela; Espeletia argentea near Bogotá in the

Colombian Eastern Cordillera.

The humid shrub páramo communities of Guaramacal show some generic

compositional and physiognomic affinities with some of the humid páramos areas

of Táchira state (Monasterio 1980b; Bono 1996). From the shrub páramos of

Táchira state, Monasterio (1980b) refered to a low and diverse páramo community

of Ruilopezia jahnii - Puya aristeguietae, as being one of the most important

communities occurring in locally wet (boggy like) areas in Páramo El Zumbador at

3200-3400 m. In this community, both Ruilopezia jahnii and Puya aristeguietae

are codominant, forming patches surrounded by dense shrub páramo communities

dominated by Blechnum aff. schomburgkii stem rosettes (Bono 1996) and shrubs,

including: Arcytophyllum caracasanum, Clusia sp. and Hypericum caracasanum

(Monasterio 1980b). Puya aristeguietae has been also reported from páramos of

Trujillo (Guirigay), Lara, Mérida (El Tambor, Pico Bolívar and La Carbonera) and

Zulia (Holst 1994), where we also assume the presence of other unnamed

associations containing this species. This big ground rosette has also been

documented for the northern páramos of the Colombian Cordillera Oriental (Cleef

1981). In Guaramacal, Puya aristeguietae is associated with the locally endemic

Ruilopezia lopez-palacii in the Puyo aristeguietae - Ruilopezietum lopez-palacii,

and is also a dominant species in the Cortaderio hapalotrichae - Hypericetum

juniperinum.


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Natural disturbances, land use and conservation

The summit of Páramo de Guaramacal has been affected by the construction of the

road and the subsequent installation of the telecommunications antennas complex

since the 1960‟s. During those years, disturbance of the natural vegetation cover

and fires have occurred. Before the construction of the road there was a path

crossing the range North-South, located just between the current road (and to the

side of it) and the location of the antennas. This track provided a commercial

connection between the village of Guaramacal, located on the South slope of

Ramal de Guaramacal, and the city of Boconó. It is also known that villagers of

the past made extensive use of the páramo adjacent to the path as fields for pasture.

Natural fires may also have occurred elsewhere in the summit zone of Ramal de

Guaramacal, especially on the driest days of the year of high radiation, as was

recently observed in Páramo El Pumar.

Since 1988, Ramal de Guaramacal has been, and continues to be, protected as a

National Park. Thus far, this has proven effective in, keeping the majority of

human activities and their associated impacts outside the park borders. Only the

area occupied by the antenna infrastructure, as well as the road and electrical

pylons in Páramo de Guaramacal, are currently treated as a special use zone („Zona

de Uso Especial‟) where some limited (disturbance) activities are permitted. The

more extensive and remote remainder of the Ramal de Guaramacal páramo

ecosystem is free from human activities and very well conserved.

Conclusions

Regardless of some methodological limitations, problems with accessibility and

environmental conditions during the study of the páramo vegetation of Ramal de

Guaramacal, the results of this study represent the first attempt at syntaxonomical

classification and understanding of the floristic composition and patterns of

bamboo páramos communities of the humid Llanos slopes of Venezuelan Andes.

The mosaic-like distribution of shrub páramo, grass páramo and dwarf forest

vegetation communities present on the summits of Ramal de Guaramacal may be

the consequence of multiple factors, influenced by the top effect promoting a low

UFL, permanently high relative humidity, and past disturbance events and fire

dynamics.

With the exception of some generic floristic affinities and physiognomic

similarities, the páramo vegetation communities described for Ramal de

Guaramacal cannot be directly related to any other of the named communities

elsewhere in the Andes.


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