ZZZ(T)-Flora, Vegetation and Ecology in the Venezuelan Andes-A Case Study of Ramal de Guaramacal (2010)

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Flora, vegetation and ecology in the Venezuelan Andes: a case study of Ramal de
Guaramacal
Cuello Alvarado, N.L.
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Cuello Alvarado, N. L. (2010). Flora, vegetation and ecology in the Venezuelan Andes: a case study of Ramal
de Guaramacal. Amsterdam: Universiteit van Amsterdam, Institute for Biodiversity and Ecosystem Dynamics
(IBED).
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Download date: 21 jun 2020
FLORA, VEGETATION AND ECOLOGY
IN THE VENEZUELAN ANDES:
A CASE STUDY OF RAMAL DE GUARAMACAL
Nidia Lourdes Cuello Alvarado
ISBN 978-90-76894-87-4
Publisher: Universiteit van Amsterdam / IBED. Amsterdam.
Cover design: Karim Rodriguez / www.wix.com/masdesign/studio
Illustrations: Angelina Licata
FLORA, VEGETATION AND ECOLOGY
IN THE VENEZUELAN ANDES:
A CASE STUDY OF RAMAL DE GUARAMACAL
ACADEMISCH PROEFSCHRIFT
ter verkrijging van de graad van doctor
aan de Universiteit van Amsterdam
op gezag van de Rector Magnificus
prof. dr. D.C. van den Boom
ten overstaan van een door het college voor promoties ingestelde
commissie, in het openbaar te verdedigen in de Agnietenkapel
op donderdag 9 september 2010, te 14.00 uur
door Nidia Lourdes Cuello Alvarado
geboren te Barquisimeto, Venezuela
Promotor:
Prof. Dr. A. M. Cleef
Prof. Dr. H. Hooghiemstra
Copromotor:
Dr. J. F. Duivenvoorden
Overige leden:
Prof. Dr. O. Huber
Prof. Dr. P. J. M. Maas
Prof. Dr. J. H. J. Schaminée
Prof. Dr. J. H. D. Wolf
Faculteit der Natuurwetenschappen, Wiskunde en Informatica
CONTENTS
Chapter 1
Introduction
1
Chapter 2
The forest vegetation of Ramal de Guaramacal in the Venezuelan Andes.
Nidia L. Cuello A. and Antoine M. Cleef
Phytocoenologia 39(1): 109-156 (2009)
7
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)
65
Chapter 4
The páramo vegetation of Ramal de Guaramacal, Trujillo State,
Venezuela. 2. Azonal vegetation.
Nidia L. Cuello A. and Antoine M. Cleef
Phytocoenologia 39(4): 389-409 (2009)
111
Chapter 5
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
(Andes, Venezuela) and its ties to other páramo floras.
Nidia L. Cuello A., Antoine M. Cleef and Gerardo A. Aymard C.
The text of Chapter 5 has been submitted to Flora (general part; to be
accepted after review) and to Anales del Jardin Botánico de Madrid
(Venezuelan part, under review)
141
Chapter 6
Functional diversity of Andean forests in Venezuela changes with altitude.
Joost F. Duivenvoorden and Nidia L. Cuello A.
submitted to Global Ecology and Biogeography
161
Chapter 7
Synthesis
175
References
187
Appendix
Summary
Samenvatting
Resumen
207
239
243
247
Acknowledgements
Curriculum Vitae
Publications
251
253
254
Chapter 1
Introduction
Nidia L. Cuello A.
Introduction
_______________________________________________________
1.1 INTRODUCTION
The Venezuelan Andes belongs to the northernmost bioregion of the Andes. With
regard to biodiversity this area represents one of the most prominent areas on a
global scale. At a regional scale it is one of the zones with highest priority for
conservation (Dinerstein et al. 1995; Mittermeier et al. 1999; Myers et al. 2000).
The Andes of Venezuela is a continuation of the Colombian Cordillera Oriental
which ends at the Táchira Depression on the Colombian-Venezuelan border. The
northern extension is a small range, the Serranía de Perijá. The northeastern
extension is the Cordillera de Mérida, mainly referred to as the Venezuelan Andes,
and, includes the Páramo de Tamá which is part of the Cordillera Oriental.
Another mountain system, the Coastal Cordillera of Venezuela, is considered as a
system separated from the Andes (Schubert 1980; González de Juana et al. 1980;
Pouyllau 1989; Vivas 1992).
The Cordillera de Mérida is composed of several ranges including the Sierra
Nevada de Mérida, Sierra de la Culata, Sierra de Santo Domingo, Sierra de Tovar,
Sierra de Uribante, and the Sierra de Trujillo. This complex of ranges is about 100
km wide and extends in northeastern direction over 450 km. The highest altitude is
reached at the Pico Bolívar (5007 m) in the Sierra Nevada de Mérida. Most of the
Cordillera de Mérida is covered by montane forest while the land over 3000 m, at
places even over 2500 m, is covered by páramo.
During the past two decades the significant taxonomic and ecological diversity of
the northern Andes, as well as the ecological importance of these mountain
ecosystems, has been recognized (Van der Hammen et al. 1983, 1986, 1989, 2003,
2005, 2008; Henderson et al. 1991; Ramsay 1992; Churchill et al. 1995; Gentry
1995; Luteyn & Churchill 2000; Rangel 2000a, 2000b, 2007; Kappelle & Brown
2001; Lauer et al. 2001; Young et al. 2002; Beck et al. 2008; Gradstein et al.
2008). However, the number of studies on biodiversity of Venezuelan mountain
ecosystems, particularly of montane forests, are limited (Ataroff 2001). The
botanical knowledge of the Venezuelan Andes is still low (Olson et al. 1997; Dorr
et al. 2000) compared to other areas, such as the Guayana Tepuis. The Tepuis area
has been always attractive for botanical exploration and there we find the greatest
efforts of botanical knowledge (Huber 1995) and the nine volumes of the Flora of
the Venezuelan Guayana (Steyermark et al. 1995).
Most studies on the flora and vegetation of the Venezuelan Andes have been
carried out in the State of Mérida. Pionering studies are those of the forests of La
Mucuy (Lamprecht 1954) and of La Carbonera (Veillon 1965). Classic studies on
the páramo vegetation are the ‘Flora de los Páramos de Venezuela’ by Vareschi
(1970) and the ecological studies on páramos by Monasterio and collaborators
(Monasterio 1980).
Recent floristic analysis in the Venezuelan Andes of montane forest (Kelly et al.
1994, 2004; Schneider et al. 2000; Schneider 2001) and páramos (Yánez 1998;
Berg 1998; Berg & Suchi 2000) are restricted to local areas or preliminary (Ortega
et al. 1987; Cuello 1996, 1999, 2002; Dorr et al. 2000). More comprehensive
studies of the flora and vegetation of larger areas, including phytogeographical
analyses of páramo flora are few (Bono 1996; Ricardi et al. 1997, 2000) and
3
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
floristic lists are up to date limited to the flowering plants of the páramo (Briceño
& Morillo 2002, 2006).
The scarcity of information on the floristic composition of the Andean ecosystems
of Venezuela is hindering a good estimation of the floristic diversity, the botanical
composition of most forests and páramos, and the status of conservation of the
endemic flora in the protected areas. In the Mérida Andes mostly dry páramos
have been studied, but almost no knowledge is available for bamboo páramos. The
descriptions of the different páramo and high Andean forest communities by
Monasterio (1980) provide a first overview. Phytosociological studies describing
associations and higher syntaxonomical units are only known for the high Sierra
Nevada de Mérida páramos (Berg 1998) and for the transect study in the Andean
and high Andean forests (Schneider 2001).
These phytosociological studies triggered our interest to learn more about the
montane forest and páramo under wet climatic conditions in the Cordillera de
Mérida. Studies on the wet Andean ecosystems has been carried out before in
Colombia (Van der Hammen et al. 1984, 2005, 2008; Rangel Ch. & Lozano C.
1986; Rangel Ch. 1994; Cleef 1981) where bamboo páramos were described for
the first time.
As is the case in other South American countries, the Venezuelan Andes are
suffering an increased human intervention and many forests and páramos have
been converted into agricultural land. The conversion of areas with montane forest
into pasture land is a common feature in the Venezuelan Andes. This practice is
changing the water flows and erosive processes in the uplands, affecting
dramatically soil stability, and the supply of water to people both in the upland and
lowland areas (Ataroff 2001).
Fortunately, a relatively large part of the montane forests and páramo ecosystems
in the Venezuelan Andes are preserved by a net of thirteen national parks that in
total cover about 31% of the whole mountain system. One of such protected areas
is Ramal de Guaramacal, located at the northeastern end of the Venezuelan Andes.
It lies within the Guaramacal National Park and strong human intervention is not
present. The Páramo of Guaramacal has been reported as an important center of
diversification of the Espeletiinae genus Ruilopezia (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).
The presence of a road traversing the Guaramacal range from the drier
northwestern slope (Andes-facing) to the moister southeastern (llanos-facing)
slopes at lower elevation, offered an excellent opportunity to initiate in 1995 a
research project to study floristically the vegetation along an altitudinal gradient.
It has been widely recognized that altitudinal gradients in mountain ecosystems
have considerable impact on the distribution of biodiversity. Altitudinal gradients
in temperature, precipitation, and other parameters are known to influence
vegetation and diversity patterns over relatively short distances (Whittaker 1960;
Gentry 1982, 1988, 1995; Lomolino 2001). Temperature can be the principal
driver of ecosystem functioning (Chapin & Körner 1995; Colwell et al. 2008;
Svenning & Condit 2008).
4
Introduction
_______________________________________________________
With support from the related project Flora of Guaramacal, jointly conducted by
Basil Stergios (UNELLEZ) and Laurence Dorr (NMNH of Smithsonian
Institution), the current project combined geobotanical exploration and floristic
surveys of the vegetation types. The aim was to explore the structure, botanical
composition, and the diversity of forests and páramo, and to relate these results to
environmental gradients. A specific goal was to examine the relationship between
altitudinal patterns of diversity and plant functional traits, particularly in the
montane rain forests. In addition, the phytogeographical patterns of the wet
páramo flora were elucidated and compared to páramo areas in Ecuador,
Colombia, and the Talamancas of Central America. The results of this study are
basic for conservation and biodiversity management in the region.
The study of vegetation along the altitudinal gradient in Ramal de Guaramacal was
done using a floristic, phytosociological and a plant functional approach. A
growing amount of literature is illustrating the value of functional approaches to
understand biodiversity, pointing out the importance of analyzing changes of
functional traits along an altitudinal gradient for predicting the effects of
environmental changes on ecosystem functioning, such as those induced land use,
cover changes or by global warming (Díaz & Cabido 1998; Díaz et al. 1999;
Duckworth et al. 2000; Lavorel & Garnier 2002; McGill et al. 2006). Recently,
some studies focused on the importance of traits associated with animal-plant
interactions to analyse the relationship between species diversity and functional
diversity with the aim to explore the ecosystem responses to environmental
change, e.g. caused by deforestation (Mayfield et al. 2005) and fragmentation
(Girao et al. 2007). In mountain ecosystems it is expected that with increasing
elevation temperature and available land surface decrease leading to more
environmental stress and more ecological filtering as a consequence (Mayfield et
al. 2005). In this study, the change of composition and diversity of some functional
(energy balance-related, reproductive/fragmentation-related) traits of undisturbed
mountain forest of Ramal de Guaramacal was analyzed along an altitudinal
gradient with the aim to contribute with benchmark information to studies of
degraded tropical Andean ecosystems (see Chapter 6).
Several topics on the diversity of flora and vegetation ecology of Ramal de
Guaramacal along altitudinal gradients have been addressed, including issues of
phytosociology, altitudinal zonation, floristic diversity, phytogeography and
functional diversity. The observed patterns were interpreted on the basis of
comparisons with other regional and extra-regional studies.
This Ph.D. thesis consists of several published articles and manuscripts in review
at international journals.
Chapter 2 includes the characterization of the physical environment of the study
area of Ramal de Guaramacal and gives an overview of the geobotanical
exploration and botanical research conducted there. This information serves as
reference information for all chapters. This chapter gives a description of the
floristic diversity and structure of the montane rain forest vegetation of Ramal de
Guaramacal and presents a syntaxonomic scheme to classify the montane rain
forests of this part of the Venezuelan Andes. It is based on quantitative data and
analysis of physiognomy, floristic composition, ecological relations, and spatial
5
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
distribution of the different vegetation communities. An altitudinal zonation is
presented and a comparison is made to montane forests elsewhere in the
Venezuelan Andes. Finally human influence and issues of conservation are
discussed.
Chapters 3 and 4 deal with the study of the wet bamboo páramo vegetation on top
of the Guaramacal mountain range. Chusquea dominated bamboo páramos are
distributed along the humid upper forest line on the Llanos facing slopes of the
Venezuelan Andes. In Venezuela this type of vegetation was not studied before.
Chapter 3 covers the phytosociological classification and description of zonal
páramo vegetation communities and examines the affinities to páramo vegetation
elsewhere in the tropical Andes. In this chapter we also address the mosaic-like
distribution of shrub páramo, grass páramo and dwarf forest vegetation
communities which are present on the summits of Ramal de Guaramacal. These
aspects are based on the analysis of physiognomy, composition, floristic diversity,
and relationships of the vegetation communities and the environmental variables
involved.
Chapter 4 comprises the classification of azonal wet páramo vegetation
communities found so far in páramo areas of Ramal de Guaramacal. Patches of
azonal bunchgrass, Sphagnum bogs, aquatic communities, and boggy bamboo
páramo are described and their floristic relationships with similar páramo
communities elsewhere in Colombia and Venezuela are discussed.
Chapter 5 is an introduction to the analysis of the phytogeographical patterns and
affinities of the lowermost and the wet páramo vegetation of Ramal de
Guaramacal. It provides an analysis of the floristic connections with the
neighboring dry páramos of the Sierra Nevada de Mérida, and the floras of other
páramo areas in the northern Andes and in Central America. We describe how
phytogeographical components change among different páramos. Using
ordinations, we explored whether the phytogeographical patterns of the páramo
flora of Ramal de Guaramacal are determined by temperature, or more by the
overall permanent humidity which characterizes the Guaramacal bamboo páramo.
Chapter 6 encompasses the analysis of functional diversity of the mountain forest
of Ramal de Guaramacal as a function of altitude. This study is based on the
vascular plant species composition of forest plots sampled along an altitudinal
gradient in the study area (chapter 2) and their linked functional traits related to
energy balance and fragmentation, obtained by means of literature and herbarium
studies. Using DCA ordination and Fourthcorner analysis, we explored the
relationships between the studied traits and environment and discuss the
implications of climate change in temperature on functional changes. This study
shows the advantage of functional approach above species approach for the
analysis of the effect of environmental changes on mountain forest ecosystems.
Finally, a synthesis of the results and conclusions based on all chapters are
presented in chapter 7.
6
Chapter 2
The forest vegetation of Ramal de Guaramacal in the
Venezuelan Andes
Nidia L. Cuello A. and Antoine M. Cleef
PHYTOCOENOLOGIA 39(1):109-156. 2009
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
2.1 INTRODUCTION
Montane forests of the northern Andes are fragile ecosystems of significant
biological and ecological diversity with a complex biogeographical history, and
playing a major role in the regional hydrological balance (Gentry 1995; Cavalier &
Goldstein 1989; Cavelier et al. 1996; Holder 2006; Kappelle & Brown 2001).
Despite increased attention and conservation interest of the northern Andean forest
ecosystem over the past two decades (Henderson et al. 1991; Churchill et al. 1995,
Gentry 1995; Luteyn & Churchill 2000; Kappelle & Brown 2001; Van der
Hammen et al. 1984), studies on montane forests of the Venezuelan Andes remain
limited in area, subject and time. The majority of studies have been carried out in
the montane forests of the state of Mérida. The silvicultural studies of La Mucuy
and La Carbonera (Lamprecht 1954; Veillon 1965, 1985), vegetation ecology
(Vareschi 1953, 1956; Yánez 1998); floristic analysis (Kelly et al. 1994, 2004);
and several studies focusing on different aspects of ecophysiology, population
ecology and hydroecology of cloud forests (Brun 1979; ICAE 2005), aspects of
diversity, structure and biogeography on a succesional and mature forest stands
close to the town of Mérida (Schneider et al. 2000; Schneider 2001) are
particularly noteworthy. Few Andean montane forest areas outside of Mérida State
have been studied (Bono 1996; Ortega et al. 1987; Cuello 1996, 1999, 2002 and
Dorr et al. 2000). Beyond the Andes, other montane forest areas previously studied
are Coastal Cordillera (Huber 1986; Howorth & Pendry 2006); Cerro El Avila
(Vareschi 1955; Steyermark & Huber 1978; Meier 2004) and Cerro Copey in
Margarita Island (Sugden 1985).
On tropical mountains, the altitudinal limit of forest formations varies with latitude
(Troll 1959, 1968) or in response to local or regional peculiarities of topography or
climate (Grubb & Whitmore 1966; Monasterio & Reyes 1980; Van der Hammen
& Cleef 1986; Van der Hammen 1995; Lauer et al. 2001; Richter 2003). In the
tropical Andes, the distribution of vegetation types and their qualitative and
quantitative composition are thought to be determined largely by gradients of
temperature, rainfall, and relative humidity (Van der Hammen & Cleef 1986; Van
der Hammen 1995), and horizontal precipitation and mist deposition (Bendix et al.
2006; Richter & Moreira-Muñoz 2005). Gradients of temperature have
pronounced effects on the pattern of vegetation zonation, especially at the limits of
the upper forest line (Troll 1973; Rundel 1994).
In the northern Andes, altitudinal vegetation zonation has been distinguished as
lowland tropical forests from 0-1000 m, lower montane (LMRF) or subandean
forests from 1000-2300 m, upper montane (UMRF) or Andean forests from 23003500 (3200-3600) m and high Andean forests from 3500 (3000-3500)-3900 m.
Open paramo vegetation is found over 3200-3900 m up to the nival zone (>48005000 m) (Cuatrecasas 1934, 1958; Van der Hammen 1974; Cleef et al. 1984; Van
der Hammen & Hooghiemstra 2001).
The existence of altitudinal zonation with discrete vegetation belts in the northern
Andes (Cuatrecasas 1958; Van der Hammen 1974) versus a continuous change in
species composition in tropical mountains, have been subject of discussion. Some
quantitative studies in other tropical mountain areas support the existence of
9
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Flora, vegetation and ecology in the Venezuelan Andes
discrete vegetation belts (Kitayama 1992; Hemp 2006), while other studies have
been less conclusive about zonation (Nakashizucha et al. 1992; Kappelle et al.
1995; Lieberman et al. 1996; Vásquez & Givnish 1998). Differences in their
scope, sampling methods, and analytical techniques may account for the
discrepancy in their conclusions (Cuello 1996, 2000; Hemp 2006).
In the Venezuelan Andes, six ecological altitudinal zones have been recognized
(Sarmiento et al. 1971; Monasterio 1980; Monasterio & Reyes 1980; Ataroff &
Sarmiento 2004): (1) a Basal zone from piedemont to 1000 m, (2) a Subandean
zone between 1000 and 2000 m, (3) a Lower Andean zone between 2000 and 3000
m, (4) an Upper Andean zone between 3000 and 4000 m, (5) a High Andean zone
between 4000 and 4800 m and (6) a Snow zone above 4800 m. The distribution of
vegetation types along altitudinal zones in the Venezuelan Andes differs between
humid and dry slopes (Sarmiento et al. 1971; Monasterio 1980; Monasterio &
Reyes 1980; Ataroff & Sarmiento 2004).
As elsewhere, the Venezuelan Andes are suffering increased human intervention.
Many areas of montane forests continue to be converted into areas of agricultural
or other land uses, while basic biodiversity studies remain scarce (Ataroff 2001).
Fortunately, due to their steep slopes and inaccessibility there are still large parts
of the Venezuelan Andes, with or without legal protection, where natural
vegetation remains relatively undisturbed. One of such areas is Ramal de
Guaramacal, the larger part of which is enclosed within a national park (Cruz
Carrillo or Guaramacal National Park), not presently subject to a strong human
intervention.
The main goal of the present study is to identify, define and characterize the
montane rain forest vegetation of Ramal de Guaramacal and to establish a
syntaxonomic scheme or classification, based on analysis of the physiognomy,
floristic composition, ecological relations and spatial distribution of the different
vegetation communities. This work was conducted within the framework of a
larger project aiming to study the floristic and vegetation diversity of the
Guaramacal National Park (Cuello 1999, 2000, 2002, 2004). The classification of
the paramo vegetation of the Guaramacal summit area is described separately
(Cuello & Cleef 2009 b, c).
2.2 STUDY AREA
Geography
Ramal de Guaramacal is a mountain range extending approximately 30 kilometers
northeast towards the eastern end of the Venezuelan Andes between 9° 0521‟ N
and 70° 0020‟ W. (Fig. 1). Parts of the Boconó Municipality in the State of
Trujillo and Sucre Municipality of the Portuguesa state are included. This
formation, in its larger extension, reaches altitudes over 2000 m. The Guaramacal
range includes summits of 3130 m in Páramo de Guaramacal; 2970 m in Páramo
El Pumar; 2800 m in Páramo Agua Fría and 2600 m in Páramo Los Rosarios.
10
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Much of the surface area of the Ramal de Guaramacal is protected by the Gral.
Cruz Carrillo Nacional Park (or Guaramacal National Park), which includes, from
the lowermost level of 1600 m, an approximate surface area of 21,466 hectares.
Climate
The climate of the Venzuelan Andean cordillera, as in the whole country is largely
determined by the Intertropical Convergence Zone (ITCZ). The great altitudinal
interval with respect to the Llanos, in combination with full exposition to trade
winds, causes high precipitation of at least 3000 mm/year from low altitude up to
approximately 2400 m on the southeastern slopes. From this position upwards and
northwards, precipitation decreases to around 2000 mm on the northern slopes
(Reaud-Thomas 1989). The climate is characterized by a dry season from
November to March and a rainy season from April to October. Maximum
precipitation occurs during June and July (Cuello & Barbera 1999). Temperatures
remain low throughout the year, averaging around 18 to 20 oC between 1000 to
1500 m, and 9 to 12 oC in the zone above 2500 m. According to Grubb (1977) and
Sarmiento (1986) the decrease of temperature with altitude (lapse rate) is around
0.6 oC/100 m. Above 2000 m the nights are cold and seasonal frosts may occur at
altitudes over 2,500 m (Reaud-Thomas 1989; Urriola 1999).
Geology
Guaramacal mountain range is aligned to the South to the Boconó fault (Schubert
1980) and to the North of the Calderas fault. It constitutes a homogeneous block
separated from the rest of the mountain range (and limiting in the North and West)
by the interandean valley of Boconó River. To the South (Llanos slope) steep
slopes descend towards the Calderas fault, from there, the smaller mountain
formations decrease in altitude towards the piedemont at 400 m (Urriola 1999).
This mountain chain is the product of orogenic processes which built the
equatorial Andes. Some of the most important geological formations of the
Venezuelan Andean Cordillera are displayed, from the oldest bedrock of the
Precambric and the Paleozoic, to more recent deposits of the Neogene (Tertiary
and Quaternary period). The most predominant geologic formations in Ramal de
Guaramacal are: Sabaneta (Paleozoic), Palmarito (Paleozoic) y El Santuario or
Gobernador (Tertiary), and, in the North slope at the end of the Boconó Fault, the
Sierra Nevada formation (Grupo Iglesias, Upper Precambric) (Ministerio de Minas
e Hidrocarburos, s/f; Urriola 1999) is present.
In the Páramo de Guaramacal area (Sector A), the Sabaneta formation
predominates. This old formation consists of a sequence of gray to brown
sandstones intercalated with limolites and red to red-violet sandstones. Around
Páramo El Pumar (Sector C), the Santuario, or Gobernador, formation (Tertiary)
arises. This formation consists of 80% friables to hardened gray quartzeous
sandstones becoming brownish under weathered conditions, being locally
conglomerated in thick layers with intercalations of light colored limolites, and
laminations of dark lutites, being occasionally calcareous and making up to 20% of
the formation (Urriola 1999).
11
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Flora, vegetation and ecology in the Venezuelan Andes
Soils
The soils of Guaramacal are determined by a set of climatic, geologic, and
topographic conditions as well as vegetation characteristics for this mountainous
landscape (Marvez & Schargel 1999). Periglacial phenomena are also an important
landscape feature at the highest altitudes with páramos and high Andean forests.
This is also true for the Guaramacal range. Glacial lakes, different types of
morraines, roches mouttonnées, etc. account for the past presence of a glacial snow
and ice cap in the summit zone during past glacial times.
The high precipitation of the region favors intense lixiviation and acidification of
the soils; thus, acid soils of low pH (<5) predominate. The soils under forests and
páramos generally display high organic matter content due to low temperatures
and acid conditions which markedly reduce the activity of soil microorganisms.
The steep slopes and the elevated water content of soils on weathered rocks, favor
mass movements and landslides at different spatiotemporal intervals; also
determining substantial variability in soil depth and coarse fragment content
(diameter > 2 mm). The predominance of sandstone of the Sabaneta and El
Santuario formations accounts for a high sand content of the soils. Nevertheless,
the presence of fine grained sedimentary lutites causes clayey soils in some
localities. The soils of the study area are predominantly of the Entisols, Inceptisols
and Ultisols order (Marvez & Schargel 1999). Morphological and chemical
characteristics of some of the soil profiles representative of Ramal de Guaramacal
(from Marvez & Schargel 1999) are presented in Appendix 3.
Vegetation
The vegetation of Guaramacal Park area is predominantely represented by
montane rain forests with height and density decreasing with altitude (Cuello
2002). Subpáramo and of páramo vegetation is locally evident from 2700 m;
however, forest vegetation patches mixed with open páramo can be found to
elevations of up to 2900-3000 m in the summit zone.
The montane forests of Ramal de Guaramacal are in the Tropical Lower Montane
Very Wet Forest and Tropical Montane Rain Forest zones of the Holdridge
climatic life zone system (Ewel et al. 1976). According to Huber & Alarcon (1988)
the forests of Guaramacal are classified as „Bosques ombrófilos
submontanos/montanos siempreverdes‟. According to the bioclimatic classification
proposed by Costa et al. (2007) for the Andes of Merida, both forest and páramo
zones of Guaramacal correspond to the Mesotropical belt.
Previous geobotanical explorations and flora research
Botanical exploration of Ramal de Guaramacal started in the 1960‟s when a rural
road between Mosquey (Boconó - North slope) and the village of Guaramacal
(South slope) was constructed, to provide access to a complex of communication
antennas installed in the summit of Páramo de Guaramacal (Ortega et al. 1987,
Dorr et al. 2000). The first floristic exploration was conducted by G.C.K.
Dunsterville in 1963, who exploited this new track to collect orchids while
workers still were felling trees for the new road (Dunsterville & Garay 1965, Dorr
12
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
1999). Julian Steyermark visited Guaramacal at a later date, accompanied by
Marvin Rabe in 1966; in 1970, with Basset Maguire and Celia K. Maguire; and
finally alone in 1971 (Dorr 1999).
During the 1970‟s several botanists visited the Guaramacal range; the late Luis
Enrique Ruiz-Terán, being among them in 1973. Dana Griffin III collected
bryophytes in 1975; José Cuatrecasas in 1978 for Espeletiinae (Asteraceae), and
James L. Luteyn in 1978 mainly for Ericaceae and other species of the páramo.
Manuel López-Figueiras and Mason Hale also collected lichens in Guaramacal in
1979 (Dorr 1999).
During the 1980‟s, staff of Herbario Universitario (PORT) of the Universidad
Nacional Experimental de los Llanos „Ezequiel Zamora‟ (UNELLEZ) in Guanare,
headed by Francisco Ortega, Basil Stergios and Gerardo Aymard, made the first
effort to catalogue the Guaramacal flora (Ortega et al. 1987, Rivero & Ortega
1989). In many of their visits they were accompanied by botanists from the United
States of America, among them were Ronald Liesner and Henk van der Werff
from the Missouri Botanical Garden; Alan R. Smith of the University of
California, Berkeley; and Laurence J. Dorr, previously of the New York Botanical
Garden (Dorr 1999).
In 1995, the first author initiated a floristic research project studying vegetation
across an elevational gradient using plots for the vegetation analysis (Cuello
1996), consequently initiating explorations in other, more remote, areas of Ramal
de Guaramacal. Simultaneously, Basil Stergios (UNELLEZ) and Laurence Dorr
(Smithsonian Institution) initiated a project to document the Flora of Guaramacal.
This project initially served as a supporting framework for the present study of the
Guaramacal vegetation. The combined efforts of botanical exploration and floristic
survey, together with contributions from an integrated multidisciplinary team of
researchers from UNELLEZ, generated publication of a reference book dealing
with several aspects of the nature of the Guaramacal National Park (Cuello 1999).
In addition to the first results detailing forest composition and diversity (Cuello
1996, 2000, 2002, 2004), a first „Catalogue of the vascular plants of Guaramacal
National Park‟ (Dorr et al. 2000) has also been published. This catalogue accounts
for a total 147 families, 517 genera and 1227 species of vascular plants.
During the last twenty years, as a result of continued integrated survey and
botanical exploration, ca. 40 species new to science have been discovered in the
Guaramacal range (Morillo 1988; Axelius & D‟ Arcy 1993; Badillo 1994,
Carnevali & Ramírez 1998; Aymard et al. 1999; Taylor 2002; Stergios & Dorr
2003; Stancik 2004; Niño et al. 2005; Cuello & Aymard 2008, for the species list
see also Stergios 1999 and Dorr et al. 2000).
The combined intensive vegetation surveys by the first author and botanical
explorations by B. Stergios, L. Dorr and M. Niño (UNELLEZ) over the past ten
years, have resulted in the addition of several new records to the Dorr et al. (2000)
catalogue.
Collections of non-vascular plants have been neglected in Guaramacal to date.
Despite the collections made during the 70‟s, a published list of bryophytes and
lichens from Guaramacal does not yet exist. Only a few of the most prominent and
13
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
conspicuous of bryophyte and lichen species present have been recently collected
by first author, mainly from páramo vegetation. Many of the collections are
undetermined at PORT, with only a present account of ca. 55 bryophyte species
and 20 species of lichens.
2.3 METHODS
Field surveys
The study was carried out during 1995, 1996, 1999, 2003, 2005 and 2006 (see
Appendix 2.). Montane forest community composition of Ramal de Guaramacal,
Venezuelan Andes, was studied along the altitudinal gradient on both sides of the
range with different slope expositions. The study area was divided into three
sectors: (A) Guaramacal, (B) Agua Fría and (C) El Santuario (Fig. 2.1). Sector A
corresponds to both slopes (N and S) in the central and higher part of the mountain
range. This area is crossed North-South by a road that leads up to the
telecommunication repeater antennas at the top of the mountain (Páramo de
Guaramacal) and then descends to the village of Guaramacal on the South slope.
Sector B comprises both slopes of the northeastern end of Ramal de Guaramacal
with. This sector includes samplings sites named after nearby villages and rivers.
Such sites are “La Peña” and “Río Frío” on the South slope, Trujillo state; “La
Divisoria” and “El Alto”, at the border of Trujillo and Portuguesa states on the
South slope; and “Laguna Negra” in Trujillo state and “El Mogote”, at the border
of Trujillo and Portuguesa states on the North slope. Sector C corresponds to the
north-western end of the Park, and includes the site known as “Qda. Honda – El
Santuario”.
Within the park access remains limited to the only existing road in the Guaramacal
sector, whereas the other sites (La Divisoria, El Alto, El Mogote, Agua Fría, Río
Frío, Laguna Megra and El Santuario) could only be reached on foot using new or
existing pathways in the forest. The old paths crossing the mountain are also
locally known as “caminos reales”; having previously served as commercial
connections between towns located on the South side of Ramal de Guaramacal and
the city of Boconó and its surroundings.
On the South slope of Agua Fría sector it was also possible to survey forests below
the Park boundaries (1350 - 1550 m). The discovery of these natural forests at low
altitude provided the opportunity to document information on this region of the
country. No other forest inventories are known from slopes at these altitudes. On
the North slope of the same sector, the lowest plot was at 1650 m, near the locality
of El Mogote, also located outside the Park boundary. For a more complete
description of the study area the reader is referred to Cuello (1999).
The forest survey contained a total of 44 samples (total area 3.705 hectares)
located at different altitudes between 1300 and 3000 m, distributed throughout the
different Park sectors (Fig. 2.1, Appendix 2). The samples include 35 plots of 1000
m2 (20 m x 50 m, divided in subunits of 10 m x 10 m) each. The forests located at
higher elevations, due to access difficulties, low stature and low diversity of
species, were surveyed in smaller plots. These include one plot of 500 m2, one of
14
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
400 m2, one of 300 m2, three of 200 m2, two of 100 m2 and one of 50 m2 (see
Table 2.2(b), Appendix 2).
B
A
C
Figure. 2.1. Geographic position of Ramal de Guaramacal in the Andes of Venezuela, with
the outline of the National Park. Park sectors: A-Guaramacal, B-Agua Fría, CQda. Honda-El Santuario.
15
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
The altitudinal intervals between sampling sites were variable between 30 to 150
meters. The plots were selected taking into account accessibility, topography and
vegetation physiognomy. Thus, plots were laid out in relatively accessible sites of
less steep slope where possible and under a rather continuous canopy. In the
highest and steepest part of the South slope, in the “Guaramacal” sector, the plots
were laid out in sites where forest had been affected by nearby landslides. This
was inevitable reflecting natural conditions, since the South slope is steeper than
the North slope (Fig. 2.11). On the South slope of Agua Fría sector, the maximum
altitude surveyed was 2125 m. Traveling by foot to this altitude was extremely
difficult as the mountain relief in this sector is comprised of contiguous small
ranges with low summits (1900-2000 m). From 1300 m (altitude of base camp),
these lower summits had to be crossed (with lower valleys between at 1700 m) in
order to reach the forest limits at around 2450 m. Unfortunately, the opportunity to
arrive at this location remained unrealised at this time.
Within each plot, all rooted individuals – trees, shrubs, lianas, tall and thick-stem
or climbing terrestrial herbs and hemiepiphytes– ≥ 2.5 cm dbh (diameter at breast
height, taken at 1.3 m from the base of the trunk, or lower for shrubs and thickstemmed herbs) were recorded, labeled with numbered aluminum tags and their
dbh and height recorded. The 2.5 cm dbh minimum size was chosen to include
most of the small woody understory species, as well as lianas and hemiepiphytes,
and to make samples comparable with the studies of Gentry (1982, 1992, 1995).
Epiphytes, non-vascular plants, small herbs and other growth forms with stems <
2.5 cm were not surveyed.
Height of trees was estimated using a 2 m clipper pole as reference. Multistemmed
species were counted as single individuals, but entire stem diameters were
recorded for calculation of basal area. The same criterion was applied to multiple
aerial roots of hemiepiphytes, such as Clusia.
Individuals were assigned to morphospecies; a voucher sample of each
morphospecies collected from each plot. For ambiguous species multiple vouchers
were collected. Morphospecies were later matched for all plots. As voucher
samples from plots were mostly sterile, general collections of fertile specimens
outside the plots were also made.
Some individuals (mainly lianas or very tall trees) could not be vouchered. In these
cases, only registry of growth form, dbh and height were taken. For each site,
collections and observations of other species not included in the surveys, herbs and
epiphytes for instance, were made. In total over 2000 botanical numbers were
collected under the number of N. Cuello (et al.) from nr. 915 to 2900 and A. Licata
from nr. 150 to 690 (see Appendix 1).
Data processing and analysis
Identification and processing of botanical specimens was made at Herbarium
PORT of the Universidad de los Llanos (UNELLEZ) in Guanare, Venezuela.
Other herbaria, such as MO and US, were also consulted. Some specimens were
sent to specialists at other institutions to confirm identification. All specimens
16
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
collected have been deposited at PORT, some duplicates have been sent to VEN,
MER, MERF, MO and US.
All the information and field data were stored and handled using Microsoft Excel.
The total listing of the inventoried species together with their respective collection
numbers appears in Appendix 1.
For the physiognomic characterization of the forests structural profiles of 20 m x
10 m (Fig. 2.2-2.10) were elaborated in the direct neighbourhood of some of the
surveyed plots. The sites selected for the elaboration of profiles are georeferenced
in Appendix 2.
A data matrix of the relative abundance of 360 species and 44 plots was processed
with TWINSPAN (Hill 1979) using the program PC-Ord 4 (McCune & Mefford
1999). The resulting TWINSPAN was interpreted in terms of syntaxonomical
classification of the vegetation, on the basis of floristic affinities, according to the
Zürich-Montpellier approach (Braun-Blanquet 1979, Westhoff & van der Maarel
1973).
For forest descriptions we followed Cuatrecasas‟s (1934) classification of
subandean, Andean and high Andean forest. However, for discussion and
comparison with other montane forests, we also referred to the equatorial
montane rain forest zonation of LMRF, UMRF and SARF by Grubb (1977) used
also elsewhere in the equatorial tropics.
2.4 RESULTS
Flora diversity
A total of 388 morphospecies with dbh ≥ 2.5 cm, corresponding to 189 genera and
78 families of vascular plants, were recorded from the 45 forest plots in Ramal of
Guaramacal. These include: 4 families, 6 genera and 13 species of pteridophytes; 5
families, 13 genera and 19 species of monocots; 68 families, 170 genera and 355
species of dicots; and 1 gymnosperm species Podocarpus oleifolius var.
macrostachyus. From the total of 388 morphospecies, 309 were identified to
species level, 55 to genus, 9 to family and 15 were not identified. An additional
177 species of vascular epiphytes, herbs and small shrubs were annotated and
collected for forest description, but not documented in the plot surveys. All the
species registered and collected from the plots are listed in Appendix 1. Table 2.1
presents the most speciose families and genera based on the plot data from this
study. Six families were represented by 20 or more species, while 3 genera were
found with 10 or more congeners.
Forest structure
Table 2.2 (a and b) summarizes the structural parameters of the different plots by
sector, slope exposure and elevation. A total of 14,895 individuals with dbh ≥ 2.5
cm were recorded in a total of 3,705 ha of accumulated forest plot. The number of
individuals per 0.1-ha plot varied from 154 to 602, with an average density of 372.
17
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
On the North slope, density in 0.1 ha plots increased with altitude up to 2480 m,
decreasing towards 2890 m. However, in forest plots (SARF) over 2900 m altitude
a high density of low diameter individuals occurs with an average density of
110.02 individuals in 100 m2 plots (about to 1100 individuals extrapolated to 0.1
ha). The canopy height was variable within plots, but generally decreased with
altitude. Taller emergent trees were found at every altitude. Maximum and mean
diameter was variable among plots, the individual with the greatest diameter
(127.3 cm) found at 2480 m.
Table 2.1. Most diverse woody plant families with dbh ≥ 2.5 cm (>10 species) and genera
(>5 species) in the plots of the montane forests of Ramal de Guaramacal, Andes,
Venezuela.
Families
LAURACEAE
RUBIACEAE
MELASTOMATACEAE
MYRTACEAE
ASTERACEAE
MYRSINACEAE
ERICACEAE
EUPHORBIACEAE
Species
34
30
27
24
20
12
12
10
Genera
Ocotea
Miconia
Eugenia
Piper
Persea
Palicourea
Psychotria
Cybianthus
Species
21
19
13
8
7
6
6
5
Table 2.2(a) Summary of structural parameters for each forest 0.1 ha plot by slope and
Sector of Ramal de Guaramacal, Venezuela. Species richness, number of
individuals, basal area, mean and maximum height, canopy height, mean and
maximum diameter (in cm).
Slope
Sector
Guaramacal
North
Agua Fría
North- (Qda.
west Honda)
18
Num
Alt. m Plot Nr. spp Num Ind
Basal
Area
1850
1960
2070
2100
2170
2300
2350
2400
2480
2580
2750
2870
2890
1830
1900
2100
2260
1880
2100
2250
4.57
2.66
6.64
3.31
5.00
8.57
5.19
6.91
4.59
5.67
5.2
4.35
3.65
4.94
7.08
5.15
5.43
4.65
5.35
4.03
5
1
19
2
18
3
20
4
17
16
39
37
35
14
25
26
27
21
22
23
36
41
46
35
41
50
60
59
36
33
41
18
27
53
60
44
61
43
55
35
182
358
446
401
316
377
479
547
602
413
458
231
423
390
320
492
438
227
324
257
Tree height
Max
4
20
24
18
26
21
22
19
23
19
17
16
12
24
28
26
24
32
30
18
Diameter
Med Canopy Max Med
7.3
10-20
64.0
9.2
8.0
10-15
38.2
7.2
7.6
10-18 114.6
7.9
6.5
10-15
62.1
7.1
9.4
10-20
90.0
9.3
7.1
8-15
111.4 10.5
8.1
8-15
60.0
7.9
6.5
6-15
108.2
8.3
7.0
6-14
127.3
6.3
8.6
6-14
69.0
9.4
9.8
7-14
42.7
8.6
8.2
6-12
44 10.1
6.7
5-10
40
8
7.3
10-20
64.0
9.2
10.8
8-24
116.5
7.8
9.4
9-23
61.0
6.9
9.1
7-18
105
8.2
11.6 14-29
83
9.9
10.4 13-26
70
8.5
8.6
7-14
50
9.5
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Slope
Sector
Guaramacal
South Agua Fría
Num
Alt. m Plot Nr. spp Num Ind
Basal
Area
1950
2100
2300
2470
2580
1330
1450
1550
1600
1770
1800
1875
1880
1950
2125
4.62
2.82
2.98
2.98
3.78
3.76
3.63
5.44
4.95
4.37
5.96
3.73
4.01
5.96
5.37
4.78
167.3
7
9
8
6
24
28
29
13
10
31
11
30
15
12
32
36
29
31
31
34
40
45
47
52
45
42
38
43
42
42
Average
Total
500
301
306
378
309
154
191
265
328
376
482
342
472
506
420
371.74
13011
Tree height
Max
22
20
21
13
18
28
27
24
25
28
26
26
25
18
24
Diameter
Med Canopy
8.9
9-18
7.7
9-13
7.9
9-16
6.5
8-11
8.6
8-14
10.4 10-24
8.8
10-24
11.7 13-22
7.3
12-20
11.5 11-24
7.2
12-20
11.5 11-22
7.2
15-22
6.7
9-13
9.9
9-18
Max Med
76 6.32
76 7.84
41.4
8.3
50
7.7
56
8.0
69 11.7
82
8.6
123 8.68
105 7.93
34
9.1
90 6.73
61
8.9
66 6.82
85 7.28
56
6.5
8.29
Table 2.2(b) Summary of structural parameters for forest plots (< 0.1 ha) in Ramal de
Guaramacal, Andes, Venezuela.
Slope
Sector
North
South
Alt. m
Average
Tree height
Diameter
Total
Plot Plot Num
Num Basal
Num
Nr. area spp
Ind/100 Area Max. Med. Canopy Max. Med.
Ind/plot
m2
2474 33
300
29
175
58.3
0.97
6.1
6-10
27
6.8
Guaramacal 2810 38
200
21
407
203.5
0.91 8.3 4.5
12
3-6
13
4.7
2830 PL3 100
10
41
41.0
0.19 4.3 2.8
2-4
16
6.8
2870 36
500
21
172
34.4
1.71
5.8
5-10
3050 34
200
18
263
131.5
0.81 6.5 4.1
3-5
21
5.3
3050 44
100
12
131
131
0.26
6
3.5
3-4
17
4.1
2950 41
400
22
272
68.0
1.77
10
6.2
5-8
25
6.9
Guaramacal 2950 40
200
19
277
138.5
1.1
6
3.6
3-5
27.5 5.6
3060 43
50
14
92
184
0.25
5
3.1
3-5
12.8 5.2
12
41.5 7.7
Forest classification
The interpretation of the TWINSPAN table, based on affinities of floristic
composition and relative species abundance, allowed recognition of seven
vegetation communities at association level, grouped in three alliances and one
major group equivalent to order level (Table 2.3). Three subandean forest (LMRF)
communities and four Andean - high Andean forest (UMRF-SARF) communities
are distinguished. The classification and description of the forest vegetation
communities of Ramal de Guaramacal are presented below.
19
20
Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23
Area 1x10 (m2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100
No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35
E 1 1 1 1 1
2 1 1
1
1 2 2 2 1 1 1 2 2 2 1 1
1 2 2 2 2
l 4 3 4 8 8
1 8 6
6
9 1 3 1 9 9 9 0 1 1 8 8
7 4 4 3 2
e 5 3 5 5 8
0 0 5
0
0 0 0 7 6 5 5 7 0 2 8 7
7 5 8 5 5
v 0 0 0 0 0
0 0 0
0
0 0 0 0 0 0 0 0 0 5 0 5
0 0 0 0 0
Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO
Park sector B B B A C C B B B B A A A A A B A B B B B B A A A C
Meliosma tachirensis - Alchornea grandiflora montane forest order group
Order
Alliance
Geonomo undatae -Posoq. coriaceae
Farameo killipii - Prunion moritzianae
1
2
Association
3
1.2
3.1
Subassociation
1.1
4.1
Variant
3.2
Subandean forests (LMRF)
Assoc. 1. Simiro erythroxylonis - Quararibeetum magnificae
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
1 5 3 1 3
Simira erythroxylon
.
.
.
.
.
. .
.
. .
.
.
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.
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.
.
Parathesis venezuelana
1 3 1 2 1
.
.
.
.
.
. .
.
. .
.
.
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.
.
Urera caracasana
2 1 1 4 .
4 3 . 4
1
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
Quararibea magnifica
.
1 3 . 3
.
.
.
.
.
. .
.
. .
.
.
.
.
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.
.
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.
.
Cuatresia riparia
.
1 1 . 1
1
.
.
.
.
. .
.
. .
.
.
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.
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.
.
Picramnia sp.
.
.
1 2 .
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
Aegiphila floribunda
1
.
.
. 3
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
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.
.
Diplazium hians
1
1
. 1 .
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
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.
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.
Inga edulis
.
1
. 1 .
.
.
.
.
.
. .
.
. .
.
.
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.
.
.
.
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.
.
Vasconcella microcarpa
.
.
.
. 1
.
.
.
.
.
. .
.
. .
.
.
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.
.
.
.
.
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.
.
Ocotea cernua
1
.
. 1 .
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
Huertea glandulosa
1
Subassoc. 1.1. typicum
.
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
Aphelandra macrophylla
3 1 2 .
.
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
Psychotria trichotoma
3 2 1 .
.
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
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.
Alchornea glandulosa
1 1 1 .
.
1 .
.
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
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.
Cecropia sararensis
3
3 2 .
.
.
.
.
.
.
. .
.
. .
.
.
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.
.
.
.
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Ocotea sp. C
.
2 3 .
.
.
.
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. .
.
. .
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Stylogyne longifolia
.
.
2 .
.
.
.
1
.
.
. .
.
. .
.
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.
.
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.
Hippotis albiflora
2
1 1 .
.
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.
. .
.
. .
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.
Matisia sp.
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. .
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. .
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Nectandra aff. membranacea
1 1
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4
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.
2
4
7
4
N
B
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.
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.
2
4
7
0
S
A
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.
4.2
2
1
0
0
S
A
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.
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.
2
5
8
0
S
A
34
.
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.
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.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
3
0
0
S
A
31
2
8
9
0
N
A
27
2
8
7
0
N
A
21
2
8
7
0
N
A
18
100
2
8
1
0
N
A
17
20
3
0
6
0
S
A
9
5
3
0
5
0
N
A
10
10
2
8
3
0
N
A
10
10
3
0
5
0
N
A
18
20
2
9
5
0
S
A
19
20
.
.
.
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.
2
9
5
0
S
A
22
40
40 41
Ruilopezio paltonioides - Cybianthion marginati
5
6
7
5.1
2
7
5
0
N
A
41
2
5
8
0
N
A
29
2
2
6
0
N
B
31
29
40
39 35 36 37 38 43 44 PL3 34
33
8
61
24
30
9
6
100 100 100 100 100 100
27 16 33
100 100
Table 2.3. Phytosociological table of montane forests of Ramal de Guaramacal, Venezuela. 1: Simiro erythroxylonis - Quararibeetum magnificae; 1.1: typicum; 1.2: bunchosietosum armeniaceae. 2: Conchocarpo larensis - Coussaretum moritzianae. 3: Croizatio brevipetiolatae - Wettinietum praemorsae; 3.1: hedyosmetosum cuatrecazanum; 3.2: var. Protium tovarense . 4: Schefflero ferrugineae - Cybianthetum laurifolii; 4.1: typicum; 4.2: miconietosum
suaveolentis. 5: Geissantho andini - Miconietum jahnii; 5.1: Subcommunity of Freziera serrata . 6: Libanothamnetum griffinii. 7: Gaultherio anastomosantis - Hesperomeletum obtusifoliae.
Flora, vegetation and ecology in the Venezuelan Andes
1 1 .
.
.
Trichilia pallida
.
.
.
.
.
.
Zygia bisingula
2
.
.
.
.
.
Ficus sp.
2
.
.
.
.
.
Paullinia capreolata
2
2
.
.
.
.
Tammsia anomala
.
2
.
.
.
.
Trichilia hirta
.
.
2 .
.
.
Piper hispidum
.
Subassoc. 1.2. bunchosietosum armeniaceae
.
. 4 .
.
Acalypha macrostachya
.
.
. 4 1
2
Psychotria fortuita
.
.
. 2 1
.
Bunchosia armeniaca
.
.
1 1 4
.
Pleurothyrium costanense
.
.
1 1 1
.
Ficus tonduzii
.
.
. 3 .
.
Saurauia tomentosa
.
.
.
. 2
.
Diplazium celtidifolium
.
.
.
. 2
2
Piper s p. (Liana)
.
.
. 2 .
.
Hydrangea aff. peruviana
.
.
.
. 1
1
Cestrum bigibbosum
.
.
.
. 1
1
Solanum nudum
.
Assoc. 2. Conchocarpo larensis - Coussaretum moritzianae
.
.
.
.
2
Coussarea moritziana
.
.
.
.
.
.
Conchocarpus larensis
4
.
.
.
.
1
Alsophila erinacea
.
.
.
. 1
1
Sloanea guianensis
.
.
.
.
.
1
Miconia lonchophylla
.
.
.
.
.
.
Meliosma pittieriana
.
.
.
.
.
1
Cyathea kalbreyeri
.
.
.
.
.
.
Eschweilera perumbonata
.
.
.
.
.
.
Chrysophyllum cf. cainito
1
.
.
. 1
1
Sloanea rufa
.
.
.
.
.
2
Mouriri barinensis
1
.
.
.
.
.
Asplundia vagans
.
.
.
.
.
.
Pseudolmedia rigida
.
.
.
. 1
1
Dussia coriacea
.
.
.
.
.
.
Eugenia grandiflora
.
.
.
.
.
.
Eugenia sp. 1
.
.
.
.
.
1
Inga aff. densiflora
.
.
.
.
.
.
Machaerium cf. floribundum
.
.
.
.
.
.
Picramnia sp. A
1
.
.
.
.
.
Ocotea rubrinervis
.
.
.
.
.
1
Stylogyne sp. A
.
.
.
. 1
3
Eugenia sp. 3
.
.
.
.
.
2
Chionanthus sp.
.
.
.
.
.
.
Tocoyena costanensis
.
.
.
.
.
.
.
.
.
.
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.
.
.
.
.
.
1
1
4
1
1
1
2
.
.
.
.
3
1
.
.
.
.
1
1
.
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.
2
.
.
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.
.
.
.
.
.
1
5
.
1
1
1
.
1
1
.
.
.
.
1
1
1
1
.
.
1
.
.
.
.
3
5
.
1
1
1
1
1
2
1
1
.
1
.
1
.
.
.
.
1
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.
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.
.
.
1
.
.
.
.
.
.
3
.
1
1
3
.
.
2
1
1
.
.
1
.
.
1
.
1
1
.
1
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1
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1
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.
1
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1
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1
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.
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.
1
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.
3
.
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.
2
1
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1
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.
1
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.
1
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.
1
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.
1
1
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.
1
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1
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2
.
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.
1
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1
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1
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1
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1
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1
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.
3
1
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1
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1
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The forest vegetation of Ramal de Guaramacal
21
Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23
Area 1x10 (m2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100
No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35
E 1 1 1 1 1
2 1 1
1
1 2 2 2 1 1 1 2 2 2 1 1
1 2 2 2 2
l 4 3 4 8 8
1 8 6
6
9 1 3 1 9 9 9 0 1 1 8 8
7 4 4 3 2
e 5 3 5 5 8
0 0 5
0
0 0 0 7 6 5 5 7 0 2 8 7
7 5 8 5 5
v 0 0 0 0 0
0 0 0
0
0 0 0 0 0 0 0 0 0 5 0 5
0 0 0 0 0
Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO
Park sector B B B A C C B B B B A A A A A B A B B B B B A A A C
Meliosma tachirensis - Alchornea grandiflora montane forest order group
Order
Geonomo undatae -Posoq. coriaceae
Farameo killipii - Prunion moritzianae
Alliance
Association
1
2
3
Subassociation
1.1
1.2
3.1
4.1
Variant
3.2
.
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2
.
. .
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. .
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Petrea pubescens
.
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2
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. .
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. .
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Piper arboreum
.
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2
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. .
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. .
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Eugenia sp.
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1
.
1 . .
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. .
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Salacia aff. cordata
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1 . .
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1 .
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Clusia sp. 1
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. 2 .
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.
1
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. .
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. 1 .
1
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Cecropia telenitida
.
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1
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. .
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1
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Eugenia sp. 2
.
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1
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. .
. 1 .
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Prunus cf. skutchii
.
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1 . .
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. .
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1
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Simira lezamae
.
Alliance 1. Geonomo undatae - Posoquerion coriaceae
2
.
1
1
1 . 1 1 . .
.
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Guarea kunthiana
1 1 3 2 2
.
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. 1
1 2 1
2
2 . .
. 2 .
.
1
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1
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Pouteria baehniana
5
.
1 4
1
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. .
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. .
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1
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Geonoma undata
1 1 4 . 1
.
1 . 3
5 1 2
.
1 1 .
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. .
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Calatola venezuelana
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.
2 1 2
2
.
2
1
1 . .
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Posoqueria coriacea
1
.
1 . 2
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2
3
2 . .
.
. .
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Rudgea nebulicola
.
.
1 1
1
1 . .
.
. .
.
1
.
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.
1
2
.
.
.
.
Matayba camptoneura
1 1 1 . 1
.
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2
3
.
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. .
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Mabea occidentalis
3
.
1 . 1
1 1 1
.
1 1 . 1 . .
.
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Persea peruviana
.
1 1 2 .
2
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Eugenia moritziana
.
.
1 . 1
1
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Marcgravia brownei
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1
1 . .
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. .
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Tapirira guianensis
1
.
. 1 .
.
1
.
1
.
. 1 .
. .
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Gordonia fruticosa
.
.
. 1 .
.
.
1
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. .
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. .
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Psychotria longirostris
.
.
1 .
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.
.
1
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. .
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. .
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Rollinia mucosa
.
1
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1
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. .
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Trigynaea duckei
.
1
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1
.
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. .
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. .
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Hydrangea cf. preslii
.
1
.
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.
1
.
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. .
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. .
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Ficus nymphaeifolia
.
1
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1
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. .
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. .
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Nectandra aff. purpurea
.
22
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4
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2
4
7
4
N
B
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2
4
7
0
S
A
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4.2
2
1
0
0
S
A
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2
5
8
0
S
A
34
.
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.
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.
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.
2
3
0
0
S
A
31
2
8
9
0
N
A
27
2
8
7
0
N
A
21
2
8
7
0
N
A
18
100
2
8
1
0
N
A
17
20
3
0
6
0
S
A
9
5
3
0
5
0
N
A
10
10
2
8
3
0
N
A
10
10
3
0
5
0
N
A
18
20
2
9
5
0
S
A
19
20
.
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2
9
5
0
S
A
22
40
40 41
Ruilopezio paltonioides - Cybianthion marginati
5
6
7
5.1
2
7
5
0
N
A
41
2
5
8
0
N
A
29
2
2
6
0
N
B
31
29
40
39 35 36 37 38 43 44 PL3 34
33
8
61
24
30
9
6
100 100 100 100 100 100
27 16 33
100 100
Flora, vegetation and ecology in the Venezuelan Andes
.
1 .
.
.
Paullinia cf. latifolia
.
Assoc. 3. Croizatio brevipetiolatae - Wettinietum praemorsae
.
.
.
.
.
Croizatia brevipetiolata
.
.
.
.
.
.
Wettinia praemorsa
.
.
.
.
.
1
Meriania grandidens
.
.
.
.
.
.
Aniba cf. cinnamomiflora
.
.
.
.
.
.
Cybianthus cuspidatus
.
.
.
.
.
.
Miconia cf. minutiflora
.
.
.
.
.
1
Elaeagia ruiz-teranii
.
.
.
.
.
.
Ocotea sp. A
.
.
.
.
.
.
Miconia lucida
.
.
.
.
.
.
Hedyosmum cf. gentryi
.
.
.
.
.
.
Faramea guaramacalensis
.
.
.
.
.
.
Maytenus sp. A
.
.
.
.
.
.
Ocotea aff. puberula
.
.
.
.
.
.
Myrcia acuminata
.
3.2. var. Protium tovarense
.
.
.
.
.
Protium tovarense
.
.
.
.
.
.
Coccoloba cf. llewelynii
.
.
.
.
.
.
Aiphanes stergiosii
.
.
.
.
.
.
Persea meridensis
.
.
.
.
.
.
Miconia sp. B
.
.
.
.
.
.
Myrcia sp.1
.
Subassoc. 3.1. hedyosmetosum cuatrecazanum
Hedyosmum cuatrecazanum
.
.
.
.
.
.
.
. 1 .
.
Palicourea demissa
.
.
.
.
.
.
Sapium stylare
.
.
. 1 .
.
Aegiphila ternifolia
.
.
.
. 2
3
Casearia tachirensis
.
.
.
.
.
.
Palicourea puberulenta
.
.
.
. 2
1
Meriania macrophylla
.
.
.
.
.
.
Perrottetia quinduensis
.
.
.
.
.
1
Guettarda crispiflora
.
.
.
.
.
1
Turpinia occidentalis
.
.
.
.
.
.
Cestrum darcyanum
.
.
. 1 .
.
Miconia amilcariana
.
Andean Forest (UMRF)
Assoc. 4. Schefflero ferrugineae - Cybianthetum laurifolii
.
.
.
.
.
Cybianthus laurifolius
.
.
.
.
.
.
Myrsine coriacea
.
.
.
.
.
.
Schefflera ferruginea
.
.
.
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.
.
Hedyosmum crenatum
.
.
.
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.
.
Miconia ulmarioides
.
.
.
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.
Ilex laurina
.
1
.
3
.
1
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1
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1
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5
2
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2
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1
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1
1
1
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1
2
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1
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2
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1
3
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1
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1
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2
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1
4
.
3
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1
1
1
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2
2
2
1
1
2
2
1
1
1
.
1
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5
2
1
1
.
1
1
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1
.
2
3
1
1
1
3
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1
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5
.
2
.
1
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4
1
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3
2
2
1
3
.
2
3
1
1
1
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5
2
1
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1
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1
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2
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1
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.
2
5
.
3
2
1
1
.
2
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2
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2
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5
1
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1
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1
1
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1
1
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4
5
1
1
1
1
1
1
1
.
1
1
.
2
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.
1
.
1
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1
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4
4
2
2
.
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1
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1
1
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2
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2
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1
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5
5
1
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1
1
1
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.
1
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1
1
.
2
.
2
5
1
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1
.
2
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1
1
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1
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4
2
2
1
.
2
.
2
.
2
3
.
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1
1
1
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1
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5
1
1
2
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.
1
.
2
.
1
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.
1
2
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1
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2
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.
4
2
2
2
.
1
1
1
.
2
.
1
.
.
.
2
1
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.
2
.
2
1
1
3
3
1
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3
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1
2
2
4
2
1
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1
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1
1
1
3
3
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1
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4
1
1
2
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1
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2
1
1
2
2
1
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1
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3
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1
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1
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4
1
1
2
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2
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4
1
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The forest vegetation of Ramal de Guaramacal
23
Plot No.
Area 1x10 (m2)
No. of Species (DBH ≥ 2.5 cm)
E
l
e
v
Slope exposure
Park sector
Order
Alliance
Association
Subassociation
Variant
Drimys granadensis
Prestoea acuminata
Palicourea apicata
Weinmannia glabra
Rhipidocladum geminatum
Podocarpus oleifolius
Ilex myricoides
Persea aff. mutisii
Weinmannia fagaroides
Byrsonima sp.
Ocotea jelski
Ilex sp.2
Persea sp.1
Byrsonima karstenii
Viburnum tinoides
Arthrostylidium venezuelae
Palicourea jahnii
Pentacalia vicelliptica
Subassoc. 4.1. typicum
Calyptranthes cf. meridensis
Brunellia cf. integrifolia
Panopsis suaveolens
Myrcia aff. guianensis
Dioicodendron dioicum
Gaiadendron punctatum
Ilex truxillensis var. bullatissima
Meliosma venezuelensis
Symplocos bogotensis
Ocotea sericea
24
1
3
3
0
S
B
1
4
5
0
S
B
1 1
2
8 8
1
5 8
0
0 0
0
N NO NO
A C C
11 14
1
8
0
0
S
B
42
1
6
5
0
N
B
53
100 100
10
1
6
0
0
S
B
52
100
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1
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1
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Geonomo undatae -Posoq. coriaceae
1
2
1.1
1.2
1
4
5
0
S
B
43
55
36
40
47
45
22
100
5
21
13 28 29
100 100 100 100 100
25
2
3
18
1
7
12 19 26 32 15 30
35
50
41
41
36
42
46
44
42
43
38
31
4
45
59
36
60
35
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1
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2
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2
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1
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1
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3
1
1
1
1
1
1
1
1
.
2
1
1
1
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1
1
1
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1
1
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1
1
1
3
1
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1
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1
1
2
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1
1
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1
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1
2
2
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1
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1
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2
2
6
0
N
B
61
2
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1
.
3
3
3
2
2
2
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1
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2
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4
2
5
8
0
N
A
33
.
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1
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2
.
3
.
2
3
1
2
1
1
.
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2
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1
.
2
4
7
4
N
B
29
30
17 20 23 27 16 33
100 100 100 100 100 100 100
1 2 2 2 1 1 1 2 2 2 1 1
1 2 2 2 2
9 1 3 1 9 9 9 0 1 1 8 8
7 4 4 3 2
0 0 0 7 6 5 5 7 0 2 8 7
7 5 8 5 5
0 0 0 0 0 0 0 0 0 5 0 5
0 0 0 0 0
N N N N N S S N N S S S S N N N NO
B A A A A A B A B B B B B A A A C
Meliosma tachirensis - Alchornea grandiflora montane forest order group
Farameo killipii - Prunion moritzianae
3
3.1
4.1
3.2
.
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1 1 1 1
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1
3 . .
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1 .
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2 1 2 1
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1
.
60
100 100 100 100 100 100 100 100 100 100 100 100
6
9
24
8
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1
1
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2
4
7
0
S
A
31
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1
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1
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4.2
2
1
0
0
S
A
29
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1
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2
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1
4
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2
5
8
0
S
A
34
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1
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1
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2
3
0
0
S
A
31
2
8
9
0
N
A
27
2
8
7
0
N
A
21
2
8
7
0
N
A
18
100
2
8
1
0
N
A
17
20
3
0
6
0
S
A
9
5
3
0
5
0
N
A
10
10
2
8
3
0
N
A
10
10
3
0
5
0
N
A
18
20
2
9
5
0
S
A
19
20
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2
2
1
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2
9
5
0
S
A
22
40
40 41
Ruilopezio paltonioides - Cybianthion marginati
5
6
7
5.1
2
7
5
0
N
A
41
40
39 35 36 37 38 43 44 PL3 34
100 100 100 100 100 100
Flora, vegetation and ecology in the Venezuelan Andes
.
.
.
Ocotea sericea
.
Subassoc. 4.2. miconietosum suaveolentis
.
.
.
Critoniopsis paradoxa
.
.
.
.
Hedyosmum sp. A
.
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.
Miconia suaveolens
.
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.
Hyeronima scabrida
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Aegiphila moldenkeana
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.
Chusquea purdieana
.
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.
Ocotea sp. B
.
.
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.
Monnina meridensis
.
Alliance 2. Farameo killipii - Prunion moritzianae
.
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.
Faramea killipii
.
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.
Clethra fagifolia
.
.
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.
Anaectocalyx bracteosa
.
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.
Cyathea pauciflora
.
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.
Prunus moritziana
.
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.
Zanthoxylum melanostictum
.
.
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.
Cyathea caracasana
1
.
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.
Cybianthus iteoides
.
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.
Eugenia cf. tamaensis
.
Weinmannia aff. balbisiana
.
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Rudgea tayloriae
.
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.
Aiouea dubia
.
.
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.
Miconia mesmeana
.
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.
Miconia tovarensis
.
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.
Myrcia cf. sanisidrensis
.
.
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.
Ocotea vaginans
.
.
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.
Hieronyma moritziana
.
.
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.
Symbolanthus vasculosus
.
.
.
.
cf. Elaeoluma nuda
.
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.
.
Ocotea leucoxylon
.
.
.
.
Mikania banisteriae
.
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.
Citronella costaricensis
.
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.
Geonoma jussieuana
.
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.
Diogenesia tetrandra
.
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.
Ocotea cf. hexanthera
.
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.
Saurauia yasicae
.
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Dicksonia sellowiana
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Mikania nigropunctulata
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Eugenia albida
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2
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1
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1
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1
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3
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1
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The forest vegetation of Ramal de Guaramacal
25
Plot No. 13 28 29 5 21 22 11 14 10 25 2 3 18 1 7 12 19 26 32 15 30 31 4 17 20 23
Area 1x10 (m2) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100
No. of Species (DBH ≥ 2.5 cm) 47 40 45 36 43 55 42 53 52 60 35 50 41 41 36 42 46 44 42 43 38 45 59 36 60 35
E 1 1 1 1 1
2 1 1
1
1 2 2 2 1 1 1 2 2 2 1 1
1 2 2 2 2
l 4 3 4 8 8
1 8 6
6
9 1 3 1 9 9 9 0 1 1 8 8
7 4 4 3 2
e 5 3 5 5 8
0 0 5
0
0 0 0 7 6 5 5 7 0 2 8 7
7 5 8 5 5
v 0 0 0 0 0
0 0 0
0
0 0 0 0 0 0 0 0 0 5 0 5
0 0 0 0 0
Slope exposure S S S N NO NO S N S N N N N N S S N N S S S S N N N NO
Park sector B B B A C C B B B B A A A A A B A B B B B B A A A C
Meliosma tachirensis - Alchornea grandiflora montane forest order group
Order
Geonomo undatae -Posoq. coriaceae
Farameo killipii - Prunion moritzianae
Alliance
Association
1
2
3
Subassociation
1.1
1.2
3.1
4.1
Variant
3.2
Montane forest order group of Meliosma tachirensis and Alchornea grandiflora
1
.
.
.
1
.
.
.
.
. 1 .
. 1 1
.
.
2 .
2
.
2 2 3 3
Clusia trochiformis
.
1
. 2 1
2 1 2
1
1 1 1 1 3 1 1 2 1 .
1 3
2 1 .
3 1
Alchornea grandiflora
.
.
. 1 1
3 1 3
1
2 3 3 3 2 . 1 2 1 2 1
.
1 1 .
1
.
Piper longispicum
.
.
. 1 .
.
1
.
1
.
. . 1 . . 1 1
.
.
1
.
1
. 2
.
2
Clusia alata
.
.
. 1 .
.
1 1
2
1 2 1 2 . 2 .
.
.
1 .
.
.
1 .
.
.
Miconia theaezans
.
2 2 .
.
.
.
1
.
.
2 3 1 . 2 1 1 2 3 1
.
1 1 .
.
.
Ruagea pubescens
.
.
2 2 .
1 1
.
1
1 2 1 1 1 2 1 1
.
.
1
.
1
.
.
1 1
Geissanthus fragrans
.
.
1 .
.
.
1 1
1
1 . .
. 3 .
.
2 1 2 3 2
2
. 1
.
.
Besleria pendula
.
.
.
.
.
.
.
.
.
2
.
2
1
.
.
3
2
1
.
1
2
3
.
.
1
.
Cyathea fulva
.
.
.
.
.
1 1
.
.
1 1 2 1 1 1 2 1 1 .
1 1
1
.
.
2
.
Beilschmiedia tovarensis
.
.
.
. 1
1 1
.
.
1 . 1 . 1 1 1 1 1 1 1 1
1 1 .
1 1
Billia rosea
1
.
.
.
.
1 1
.
1
1 . 1 1 1 2 1 2 1 2 .
.
.
1 .
1
.
Hieronyma cf. oblonga
.
.
.
1 .
.
.
.
.
.
1 1 .
.
. . 1
.
1 .
1 1
.
1 .
2 2
Psammisia hookeriana
.
1 .
.
1 1 1
.
2 . . 1 2 1 1 1 1 .
1 1
.
.
.
.
.
Blakea schlimii
.
.
.
.
.
2
.
.
.
1 1 1 1 . .
.
.
1 .
.
1
2 1 .
1
.
Meliosma tachirensis
.
.
1 . 1
.
.
.
1
.
. .
.
. .
.
.
.
.
5 1
.
.
.
.
1
Sphaeradenia laucheana
.
.
. 1 1
.
.
2
.
.
1 1 1 1 .
.
1
.
1 .
.
.
.
.
1
.
Tetrorchidium rubrivenium
1
.
.
.
.
1
.
1
.
1 . .
. 3 .
.
3
.
.
.
.
1
.
.
.
.
Palicourea angustifolia
.
.
.
.
.
.
.
1
2
1 . .
. 1 .
.
1
.
.
1
.
2
.
.
.
.
Dendropanax arboreus
1
.
. 1 .
.
.
.
.
.
. .
. 1 1 1 1
.
.
.
.
.
1 1 1
.
Ocotea karsteniana
.
.
.
.
.
.
.
1
1
2 . .
.
. .
.
.
.
.
.
2
.
.
.
.
.
Eugenia cf. oerstediana
1
.
.
.
.
1 . . 1 . .
.
.
.
.
.
.
.
.
.
1
.
Miconia cf. dolichopoda
1 1 1 1 .
.
.
. 1
1
.
.
.
.
1 1 1 . .
.
.
.
.
.
.
.
. 1
.
.
Trichilia septentrionalis
.
.
.
.
.
.
.
.
1
.
1 2 .
. .
.
.
1 .
.
.
.
2 .
1
.
Ocotea floribunda
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
2
2
.
.
.
.
Weinmannia sorbifolia
.
.
.
.
.
.
.
1
.
1 . .
.
. .
.
.
.
.
1
.
.
1 .
1 1
Ocotea macropoda
.
.
.
.
.
.
1
.
.
.
. .
.
. .
.
.
2 .
.
.
.
.
.
2
.
.
Elaeagia karstenii
.
.
.
.
.
.
1
1
.
. .
.
. .
.
.
.
.
.
.
.
.
.
1
.
Tabebuia guayacan
1
26
2
1
.
1
1
.
.
.
1
1
1
.
.
.
.
2
.
.
.
1
.
1
.
.
.
.
.
.
3
.
.
1
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
.
.
4
4
.
.
2
.
.
.
.
.
.
.
.
.
.
.
3
.
.
.
.
.
.
.
.
.
.
.
.
2
4
7
4
N
B
2
.
.
3
1
.
.
.
.
.
.
.
.
.
1
.
.
1
.
.
.
.
.
.
.
.
.
.
2
4
7
0
S
A
4
2
.
2
3
.
1
.
.
.
.
1
.
.
.
.
1
.
.
.
2
.
.
.
.
.
.
.
4.2
2
1
0
0
S
A
3
.
.
4
1
.
.
.
1
.
.
.
1
.
.
.
.
.
.
.
.
.
1
.
.
.
.
.
2
5
8
0
S
A
34
3
.
.
3
3
2
.
.
.
.
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
.
.
2
3
0
0
S
A
31
2
8
9
0
N
A
27
2
8
7
0
N
A
21
2
8
7
0
N
A
18
100
2
8
1
0
N
A
17
20
3
0
6
0
S
A
9
5
3
0
5
0
N
A
10
10
2
8
3
0
N
A
10
10
3
0
5
0
N
A
18
20
2
9
5
0
S
A
19
20
.
.
.
1
.
.
.
.
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
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.
.
.
.
.
.
.
.
.
1
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1
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2
9
5
0
S
A
22
40
40 41
Ruilopezio paltonioides - Cybianthion marginati
5
6
7
5.1
2
7
5
0
N
A
41
2
5
8
0
N
A
29
2
2
6
0
N
B
31
29
40
39 35 36 37 38 43 44 PL3 34
33
8
61
24
30
9
6
100 100 100 100 100 100
27 16 33
100 100
Flora, vegetation and ecology in the Venezuelan Andes
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Hedyosmum translucidum
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Freziera serrata
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Weinmannia auriculata
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Weinmannia karsteniana
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Macrocarpaea bracteata
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Ocotea calophylla
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Assoc. 6. Libanothamnetum griffinii
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Libanothamnus griffinii
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Assoc. 7. Gaultherio anastomosantis - Hesperomeletum obtusifoliae
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Hesperomeles obtusifolia
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Diplostephium obtusum
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Gaultheria anastomosans
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Pentacalia greenmanniana
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Ageratina theifolia
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Alliance 3. Ruilopezio paltonioides - Cybianthion marginati
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Miconia tinifolia
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Cybianthus marginatus
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Myrsine dependens
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Vaccinium corymbodendron
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Pentacalia cachacoensis
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Ruilopezia paltonioides
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Blechnum schomburgkii
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Monochaetum discolor
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Symplocos tamana
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Ilex guaramacalensis
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Themistoclesia dependens
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Monnina sp.
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Clusia sp. A
1
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Sloanea brevispina
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Macleania rupestris
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Mikania stuebelii
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Celastrus liebmannii
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Cyathea pungens
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Geonoma orbignyana
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Assoc. 5. Geissantho andini - Miconietum jahnii
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Weinmannia lechleriana
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Geissanthus andinus
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Miconia jahnii
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Oreopanax discolor
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Disterigma alaternoides
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Thibaudia floribunda
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Pentacalia theifolia
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Subcommunity 5.1. Freziera serrata
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3
3
2
1
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5
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5
4
4
1
1
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2
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The forest vegetation of Ramal de Guaramacal
27
28
Bredemeyera sp. 25(1)
Brunellia acutangula 6(1)
Calyptranthes sp. 27(1)
Cecropia sp. 25(1)
Cestrum buxifolium 24(1)
cf. Escallonia hispida 27(1)
Chrysophyllum sp. 22(1)
Cissus trianae 15(1)
Citharexylum venezuelense 13(1)
Coccoloba sp. 28(1)
Coffea arabica 10(1)
Alsophila angelii 27(1)
Anthurium eminens 29(1)
Anthurium ginesii 24(1)
Anthurium humboldtianum 27(1)
Anthurium nymphaeifolium 15(1)
Anthurium smaragdinum 21(1)
Bacharis brachylaenoides 39(1)
Ageratina neriifolia 20(1)
Allophylus cf. glabratus 18(1)
Plot No.
Area 1x10 (m2)
No. of Species (DBH ≥ 2.5 cm)
E
l
e
v
Slope exposure
Park sector
Order
Alliance
Association
Subassociation
Variant
Chusquea angustifolia
Chaetolepis lindeniana
.
.
0
0
0
0
1
3
3
0
S
B
1
4
5
0
S
B
1 1
2
8 8
1
5 8
0
0 0
0
N NO NO
A C C
11 14
1
8
0
0
S
B
42
1
6
5
0
N
B
53
100 100
10
1
6
0
0
S
B
52
100
.
.
0
0
0
0
.
.
0
0
1
1
1
7
12 19 26 32 15 30
35
50
41
41
36
42
46
44
42
43
38
31
4
45
59
36
60
35
.
.
0
1
1
0
1
4
2
5
8
0
N
A
33
.
.
0
1
1
0
1
2
4
7
4
N
B
29
6
9
24
8
.
.
0
1
1
1
0
2
4
7
0
S
A
31
.
.
0
1
1
1
0
4.2
2
1
0
0
S
A
29
.
.
0
1
1
1
0
2
5
8
0
S
A
34
.
.
0
1
1
1
1
2
3
0
0
S
A
31
2
8
9
0
N
A
27
2
8
7
0
N
A
21
2
8
7
0
N
A
18
100
2
8
1
0
N
A
17
20
3
0
6
0
S
A
9
5
3
0
5
0
N
A
10
10
2
8
3
0
N
A
10
10
3
0
5
0
N
A
18
20
2
9
5
0
S
A
19
20
1
1
1
0
1
.
.
1
0
1
2
.
1
0
1
.
.
1
1
0
0
.
.
1
1
0
1
.
.
1
1
0
1
.
.
1
1
0
1
.
1
1
1
1
.
.
1
1
1
1
.
1
1
1
A: Guaramacal; B: Agua Fria, C: El Santuario
Solanum aturense 18(1)
Solanum confine 3(1)
Sphaeropteris sp. 3(1)
Talauma sp. 13(1)
Ternstroemia acrodantha 4(1)
Ternstroemia sp. A 17(1)
Ternstroemia sp.B 16(1)
Vismia baccifera 7(1)
Zanthoxylum acuminatum 6(1)
Rhamnus sphaerosperma 27(1)
Roupala barnettiae 32(1)
Ruagea glabra 1(1)
Ruellia tubiflora 3(1)
Schlegelia spruceana 14(1)
Sloanea laurifolia 1(1)
Smilax kunthii 20(1)
Randia cf. dioica 14(1)
.
.
1
0
0
2
9
5
0
S
A
22
40
40 41
Ruilopezio paltonioides - Cybianthion marginati
5
6
7
5.1
2
7
5
0
N
A
41
40
39 35 36 37 38 43 44 PL3 34
100 100 100 100 100 100
Panopsis sp. 31(1)
Paragynoxis cuatrecasasii 39(1)
Paragynoxis venezuelae 6(1)
Persea ferruginea 23(1)
Persea sp.2 27(1)
Persea sp.3 14(1)
Piper aduncum 5(1)
Piper phytolaccifolium 13(1)
Piper sp. 18(1)
Piper veraguense 22(1)
Psychotria amita 22(1)
Psychotria cf. lindenii 10(1)
Neea sp. 13(1)
Ocotea aff. tarapotana 14(1)
Ocotea auriculata 5(1)
Ocotea sp. 33(1)
Ocotea terciopelo 20(1)
Oreopanax sp. 38(1)
Ossaea micrantha 21(1)
Nectandra sp. 13(1)
.
.
0
1
1
0
0
2
2
6
0
N
B
61
30
17 20 23 27 16 33
100 100 100 100 100 100 100
1 2 2 2 1 1 1 2 2 2 1 1
1 2 2 2 2
9 1 3 1 9 9 9 0 1 1 8 8
7 4 4 3 2
0 0 0 7 6 5 5 7 0 2 8 7
7 5 8 5 5
0 0 0 0 0 0 0 0 0 5 0 5
0 0 0 0 0
N N N N N S S N N S S S S N N N NO
B A A A A A B A B B B B B A A A C
Meliosma tachirensis - Alchornea grandiflora montane forest order group
Farameo killipii - Prunion moritzianae
3
3.1
4.1
3.2
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
.
. .
.
. .
.
.
.
.
.
.
.
.
.
.
.
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 0
0 1 1 1 1
1 0 0 0 1 1 1 1 1 1 1 1
1 0 0 0 0
1
0 0 0 0 0 0 1 1
1 0 0 0 0
60
Markea sp. 20(1)
Mascagnia sp. 10(1)
Meliosma meridensis 4(1)
Mendoncia tovarensis 14(1)
Miconia donaeana 29(1)
Miconia elvirae 41(1)
Miconia sp. C (hibrido) 3(1)
Mikania houstonians 22(1)
Mikania sp.1 20(1)
Monnina sp.2? 41(1)
Morus insignis 28(1)
Myrcianthes sp. 3(1)
.
.
0
0
1
1
0
1
Endlicheria sp. 25(1)
Eschweilera sp. nov. 13(1)
Eugenia sp. 33(1)
Eugenia triquetra 37(1)
Eugenia sp. 2 39(1)
Ficus tovarensis 25(1)
Fuchsia membranacea 37(1)
Gaultheria erecta 38(1)
Greigia albo-rosea 20(1)
Hedyosmum racemosum 1(1)
Heisteria acuminata 28(1)
Henriettella cf. verrucosa 14(1)
.
.
0
0
1
1
0
18
Henriettella tovarensis 10(1)
Hesperomeles sp. 36(1)
Hoffmannia pauciflora 21(1)
Hydrangea sp.1 22(1)
Hypericum paramitanum 40(1)
Ilex sp.1 5(1)
Ladenbergia cf. buntingii 13(1)
.
.
0
0
1
0
3
Henriettella sp. 25(1)
.
.
0
0
0
1
2
Cyathea aff. straminea 27(1)
Cybianthus stapfii 38(1)
Dichapetalum pedunculatum 25(1)
Disterigma sp. 27(1)
Drymonia crassa 18(1)
Elaeagia myriantha 5(1)
Elateriopsis oerstedii 28(1)
.
.
0
0
0
1
25
100 100 100 100 100 100 100 100 100 100 100 100
Cupania cf. scrobiculata 13(1)
.
.
0
0
0
0
Geonomo undatae -Posoq. coriaceae
1
2
1.1
1.2
1
4
5
0
S
B
43
55
36
40
47
45
22
100
5
21
13 28 29
100 100 100 100 100
Flora, vegetation and ecology in the Venezuelan Andes
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
MONTANE FOREST ORDER GROUP OF MELIOSMA TACHIRENSIS AND ALCHORNEA
GRANDIFLORA
Physiognomy: This group of forests concerns humid montane cloud forest
communities belonging to both subandean and Andean forest. These forests are
dense, with a high number of thin-stemmed individuals and a medium-high canopy
(25-30 m) in subandean forests to medium-low (15-20 m) in Andean forests. The
presence of a high bryophyte cover on tree trunks is characteristic.
Composition and syntaxonomy: Among the characteristic large tree canopy
species of this forest group are Alchornea grandiflora, Beilschmiedia tovarensis,
Billia rosea, Elaeagia karstenii, Hieronyma cf. oblonga, Miconia cf. dolichopoda,
Ruagea pubescens, Tetrorchidium rubrivenium. Common hemiepiphitic trees are
Clusia trochiformis and Clusia alata. Also are frequent the lianas and vines Blakea
schlimii, Celastrus liebmannii, Macleania rupestris, Mikania stuebelii and
Psammisia hookeriana. Besleria pendula and the tree ferns Cyathea fulva and C.
pungens are also common among the smaller trees of up to 6 m tall. Diagnostic
species of the subcanopy are Geissanthus fragans, Meliosma tachirensis, Miconia
theaezans, Piper longispicum var. glabratum
This group of forests with Meliosma tachirensis and Alchornea grandiflora could
be considered as a provisional order, in which, the following two alliances are
recognized: Geonomo undatae - Posoquerion coriaceae and Farameo killipii Prunion moritzianae.
Ecology and distribution: The forest communities belonging to the montane
forest order group of Meliosma tachirensis - Alchornea grandiflora are found from
1350 m on the South slope, from 1650 m on the North slope and up to around 2600
m on Ramal de Guaramacal.
GEONOMO UNDATAE – POSOQUERION CORIACEAE Cuello & Cleef 2009
Typus: Simiro erythroxylonis – Quararibeetum magnificae. Table 2.3
Subandean forests of the Geonoma undata - Posoqueria coriacea alliance
Physiognomy and composition: The forest communities of this alliance are
humid forests of medium to high stature, up to 25-30 m tall, characterized by the
presence of high trees of: Rubiaceae, Euphorbiaceae, Lauraceae, Sapotaceae,
Melastomataceae, Moraceae, Bombacaceae, Meliaceae and Rutaceae being among
the most important according to abundance, frequency and basal area. The most
diverse families by species represented are Rubiaceae, Lauraceae, Melastomataceae, Myrtaceae, Euphorbiaceae and Meliaceae.
Among the canopy species can be found Calatola venezuelana, Ficus
nymphaeifolia, Gordonia fruticosa, Matayba camptoneura, Mouriri barinensis,
Nectandra aff. purpurea, Persea peruviana, Posoqueria coriacea, Pouteria
baehniana, Tapirira guianensis and Trigynaea duckei. In the subcanopy are
common: Eugenia moritziana, Geonoma undata, Guarea kunthiana, Mabea
29
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Flora, vegetation and ecology in the Venezuelan Andes
occidentalis, Psychotria longirostris, Rollinia mucosa and Rudgea nebulicola.
Some lianas can also be found in these forests, such as: Hydrangea cf. preslii,
Marcgravia brownei, Paullinia cf. latifolia and Salacia aff. cordata.
Syntaxonomy: This alliance is defined on the basis of ten 0.1-ha plots that include
211 species with dbh ≥ 2.5 cm, corresponding to 123 genera and 60 families of
vascular plants. The diagnostic species in the canopy are Posoqueria coriacea and
Pouteria baehniana. In the subcanopy small trees of Matayba camptoneura and
Guarea kunthiana, and the palm Geonoma undata are also diagnostic.
This alliance includes two associations: Simiro erythroxylonis - Quararibeetum
mag-nificae and Conchocarpo larensis - Coussareetum moritzianae.
Ecology and distribution: The Geonoma undata - Posoqueria coriacea alliance
of subandean forest groups the forest communities located on the lower slopes of
Ramal de Guaramacal, in particular areas of remnant forests near and beyond the
border of the National Park.
1. Simiro erythroxylonis – Quararibeetum magnificae Cuello & Cleef 2009
Typus: Cuello Plot No. 28. 1880 m, Table 2.3
Subandean forests of Simira erythroxylon and Quararibea magnifica
Physiognomy and composition: The subandean forests of Simira erythroxylon
var. meridensis and Quararibea magnifica display a medium stature and density,
mainly composed of mature trees with an average diameter greater than 10 cm and
a few thin individuals. The canopy of the forest is composed of trees of between 10
to 28 m with a dense cover; Simira erythroxylon var. meridensis, Quararibea
magnifica, Ocotea cernua and Posoqueria coriacea being the most abundant
species. In some areas there are some emergent trees of up to 32 m, such as:
Pleurothyrium costanense, Casearia tachirensis, Sloanea aff. guianensis and
Simira erythroxylon var. meridensis, being among the most abundant.
Simira erythroxylon var. meridensis is also common in the subcanopy (5-10 m),
together with treelets of Parathesis venezuelana, Aegiphila floribunda, Inga edulis,
Miconia cf. dolichopoda, Trichilia pallida, Vasconcella microcarpa, the small
trees, Cuatresia riparia, Picramnia sp. C. and the tree ferns Cyathea pungens and
C. caracasana. Among the most abundant lianas and climbers present are:
Anthurium eminens, A. smaragdinum, Campyloneuron ophiocaulon, Elateriopsis
oerstedii, Paullinia capreolata, Piper sp., Smilax spinosa, Sphaeradenia laucheana and Trichomanes radicans. Epidendrum unguiculatum, Guzmania mitis,
Maxillaria nigrescens, Mezobromelia capituligera, Peperomia ouabianae, P.
peltoidea, P. portuguesensis and Polytaenium lineatum stand out among the
epiphytes.
The understory is rich in terrestrial ferns, such as: Asplenium alatum, Didymochlaena truncatula, Diplazium celtidifolium, D. hians, Polystichum muricatum,
some of them reaching heights of up to 2 m. The terrestrial orchid Corymborkis
flava, the palm Chamaedorea pinnatifrons, and shrubs like Urera caracasana,
30
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Psychotria fortuita and several species of the genus Piper, such as P. hispidum, P.
dilatatum and P. aduncum, are also present, among others.
Syntaxonomy: This association is defined on the basis of ten 0.1-ha plots with 127
species with dbh ≥ 2.5 cm. The diagnostic species are Simira erythroxylon in the
canopy, and Parathesis venezuelana in the subcanopy. Two subassociations can be
recognized: subass. typicum and bunchosietosum armeniacae.
Ecology and distribution: The subandean forest of the association Simiro
erythroxylonis – Quararibeetum magnificae can be found on the southern slope of
Ramal de Guaramacal, sector Agua Fría, in the surroundings of Río Frío, (13001500 m); in the northwestern sector of Qda. Honda (1800-2100 m); and on the
northern slope, around the recreative area Laguna de los Cedros (1800-1900 m)
(Photo 2.1).
Photo 2.1. Aspect of the subandean forest of the North slope of Ramal de Guaramacal.
Forests with whitish canopies of Cecropia telenitida above of the recreative area
“Laguna de los cedros” at 1800 m.
Simiro erythroxylonis – Quararibeetum magnificae
1.1 subassociation typicum Cuello & Cleef 2009
Typus: Cuello Plot No. 28, 1450 m. Photo 2.2
Subassociation of Simira erythroxylon
31
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Flora, vegetation and ecology in the Venezuelan Andes
Physiognomy and composition: The canopy trees reach 10-24 m with a dense
cover. The composition is as described for the association; Simira erythroxylon,
Pouteria baehniana and Quararibea magnifica, also being abundant, along with
other species like Cecropia sararensis, Eugenia moritziana, Hippotis albifora,
Tammsia anomala and Trichilia pallida. The subcanopy differs from that of the
association by the major presence of Alchornea glandulosa, Hippotis albiflora,
Rudgea nebulicola, Simira lezamae, Stylogyne longifolia and the palm Geonoma
undata, which can reach heights of 14 m. Among the small trees and shrubs of 3 to
6 m, Aphelandra macrophylla, Cuatresia riparia and Psychotria trichotoma are
prominent. Further occurances of the shrubs: Palicourea petiolaris, Piper
phytolaccifolium, Psychotria patria, Randia dioica, Ruellia tubiflora var.
tetrastichantha, Winterigia solanacea; stranglers like Dioscorea coriacea;
epiphytes as Dichaea sp., Elleanthus graminifolius, Jacquiniella teretifolia and
Peperomia peltoidea; ferns like Asplenium uniseriale; large perennial herbs like
Heliconia hirsuta and Sphaeradenia laucheana; and small herbs: Heppiella
viscida, Ichnanthus nemorosus, Solanum pentaphyllum, Sanicula liberta, and
Begonia trispathulata are also present in the ground layer.
Syntaxonomy: The subassociation is defined on the basis of three 0.1-ha plots
with 83 species with dbh ≥ 2.5 cm. The diagnostic species in the canopy are
Alchornea glandulosa, Cecropia sararensis and Ocotea sp. C. Aphelandra
macrophylla and Psychotria trichotoma are diagnostic in the subcanopy.
Other species found in the forest only of this subassociation, albeit at very low
density and frequency, are: Citharexylum venezuelense, Coccoloba sp., Cupania
cf. scrobiculata, Eschweilera sp. nov. (Cuello 1832), Heisteria acuminata, Ladenbergia cf. buntingii, Nectandra sp. (Cuello 1838), Neea sp. (Cuello 1851), Morus
insignis, Miconia donaeana and Talauma sp. (Cuello 1745).
Ecology and distribution: The forests of the Simiro erythroxylonis Quararibeetum magnificae subassociation typicum are located between 1300 and
1500 m, on the South slope, near the border of Portuguesa-Trujillo states and also
in the surroundings of the Río Frío (sector B - Agua Fría). These forests represent
the few remaining extensions of undisturbed mature forest of this altitudinal zone.
Simiro erythroxylonis – Quararibeetum magnificae
1.2. subassociation bunchosietosum armeniacae Cuello & Cleef 2009
Typus: Cuello Plot No. 5, Fig. 2.2
Subassociation of Bunchosia armeniaca
Physiognomy and composition: Physiognomy and composition as described for
the association; Acalypha macrostachya, Calatola venezuelana, Cecropia
telenitida, Ficus tonduzii and Pleurothyrium costanense being more abundant in
the canopy. This subassociation differs from the typicum subassociation by the
subcanopy presence of Bunchosia armeniaca, Cestrum bigibbosum, Hydrangea
aff. peruviana, Psychotria fortuita, Saurauia tomentosa and Solanum nudum, and
the high density of Diplazium celtidifolium in the understory.
32
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Syntaxonomy: The forests of the subassociation bunchosietosum armeniacae are
represented by two 0.1-ha plots, with a total of 67 species with dbh ≥ 2.5 cm.
The diagnostic species of the subassociation are Bunchosia armeniaca, Psychotria
fortuita and Pleurothyrium costanense.
Photo 2.2. Interior of plot 29 of the subandean forest of the association Simiro
erythroxylonis - Quararibeetum magnificae subassociation typicum at 1450 m
in the Agua Fría-Río Frío sector on the South slope.
Ecology and distribution: The forests of the subassociation of Bunchosia
armeniaca are those patches of subandean forest between 1800 and 1900 m on the
northern and northwestern slope of Ramal de Guaramacal. Their extension is
limited by the Park border. In general this altitudinal interval is occupied by human
activities throughout the North slope. This community can be recognized from
distance by presence of Cecropia telenitida (“yagrumo blanco”) due to its
conspicuous white color leaf pubescence (Photo 2.1). This species typically
33
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
constitutes a zone of forest on the North slope in Guaramacal, generally
corresponding to the zone of cloud cover accumulation at around 1800-2000 m.
Subandean forest with Cecropia telenitida or vicariant species can also easily be
observed elsewhere in the Andes of both Venezuela and Colombia (Cuatrecasas
1958; Smith 1985; Cleef et al. 2003).
Figure 2.2. Subandean forest of the association Simiro erythroxylonis - Quararibeetum
magnifcae subassoc. bunchosietum armeniacae. Plot 5: 1850 m, North slope.
Af: Aegiphila floribunda; Ba: Bunchosia armeniaca; Ct: Cecropia telenitida;
Em: Eugenia moritziana; Gk: Guarea kunthiana; Hp: Hydrangea peruviana;
Md: Miconia cf. dolichopoda; Pc: Pleurothyrium costanense; Pf: Psychotria
fortuita; Poc: Posoqueria coriacea; Pv: Parathesis venezuelana; Se: Simira
erythroxylon; Uc: Urera caracasana.
34
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
2. Conchocarpo larensis – Coussareetum moritzianae Cuello & Cleef 2009
Typus: Cuello Plot No. 10. Table 2.3, Fig. 2.3
Subandean forests of Conchocarpus larensis y Coussarea moritziana
Physiognomy and composition: The canopy height is between 10 and 25 m, with
emergent trees of up to 30 m and a species compositon as described for the
alliance.
Some of the most abundant species in the canopy are Coussarea moritziana,
Miconia lonchophylla, Aniba cf. cinnamomiflora, Protium tovarense, Eschweilera
perumbonata, Chrysophyllum cf. cainito, Tocoyena costanensis subsp. andina and
Sloanea guianensis.
Among the subcanopy trees (3-10 m tall), Piper longispicum var. glabratum,
Mabea occidentalis, Croizatia brevipetiolata, Hedyosmum cf. gentryi, Conchocarpus larensis, Meliosma pittieriana, Tabebuia guayacan, Rudgea nebulicola,
Faramea guaramacalensis, Petrea pubescens, Eugenia cf. tamaensis, and the
palms Geonoma undata and Wettinia praemorsa stand out.
Lianas are also abundant in this community. Among them are Clusia sp.1,
Dichapetalum pedunculatum, Hydrangea sp. 1 (Cuello 2211), Machaerium cf.
floribundum, Mascagnia sp. A, Mendoncia tovarensis, Mikania houstonians, Salacia aff. cordata, Schlegelia spruceana, the climber Asplundia vagans and
hemiepiphytic trees such as Clusia alata y C. trochiformis. The most common
epiphytic species are the ferns Asplenium raddianum, Microgramma percusa, the
bromeliads Guzmania mitis and Mezobromelia capituligera, and the orchids
Pleurothallis biserrula, Scaphyglottis summersii and Trichocentrum pulchrum.
Among the common small trees and shrubs less than 3 m tall are: Besleria
pendula, Dendropanax arboreus, Eugenia cf. oerstediana, Psychotria amita, P. cf.
lindenii, Piper aequale, and the tree ferns Alsophila erinacea, Cyathea kalbreyeri
and C. fulva. In some places there are also dense colonies of large-leaved perennial
herbs such as Sphaeradenia laucheana, Heliconia hirsuta and Renealmia nicolaioides. Small herbs like Heppiela viscida and ferns such as Danaea moritziana,
Arachniodes denticulada, and Asplenium radicans are present in the ground layer.
Syntaxonomy: The association of Conchocarpo larensis - Coussareetum moritzianae is defined on the basis of five 0.1 ha-plots, with 145 species with dbh ≥ 2.5 cm.
This forest association can be distinguished from the other two forest associations
of the alliance by the diagnostic presence of Conchocarpus larensis, Coussarea
moritziana, Meliosma pittieriana, Hedyosmum cf. gentryi, Pseudolmedia rigida
and Cyathea kalbreyeri.
Other diagnostic species in this association, although of lesser abundance and
frequency, but absent elsewhere are: Alsophila erinacea, Asplundia vagans,
Eugenia grandiflora, Machaerium cf. floribundum, Petrea pubescens, Piper
arboretum, Salacia aff. cordata and Tocoyena costanensis.
35
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
The canopy species Aniba cf. cinnamomiflora, Protium tovarense, Miconia
lonchophylla, Sloanea guianensis and Wettinia praemorsa present in this
association, are also absent from the previous association of the alliance, but are
common in the forest of the Croizatio brevipetiolatae - Wettinietum praemorsae of
the Farameo killipii - Prunion moritzianae.
Ecology and distribution: The forests of the Conchocarpo larensis - Coussareetum moritzianae are located mainly at the northeastern end of Ramal de Guaramacal (sectors Laguna Negra, El Mogote, El Alto) at altitudes between 1600 and
1900 m on both the North and South slopes, as well as at 2100 m in the
northwestern sector of Qda. Honda.
Figure 2.3. Subandean forest of the association Conchocarpo larensis - Coussaretum
moritzianae. Plot 10: 1650 m, South slope. Cc: Chrysophyllum caimito; Ccu:
Cybianthus cuspidatus; Cm: Coussarea moritziana; Cl: Conchocarpus
larensis; Da: Dendropanax arboreus; Gu: Geonoma undata; Ho: Hyeronima
oblonga; Mo: Mabea occidentalis; Mb: Mouriri barinensis; Mp: Meliosma
pittierana; Pp: Petrea pubescens; Pl: Psychotria cf. lindenii; Wp: Wettinia
praemorsa.
36
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
FARAMEO KILLIPII – PRUNION MORITZIANAE Cuello & Cleef 2009
Typus: Croizatio brevipetiolatae - Wettinietum praemorsae. Table 2.3
Subandean and Andean forests of the Faramea killipii and Prunus moritziana alliance
Physiognomy and composition: These are humid forests with a moderate to high
density of trees of medium to tall height. The most important trees with regards to
abundance, frequency and basal area belong to the families Euphorbiaceae,
Melastomataceae, Rubiaceae, Arecaceae, Clusiaceae, Lauraceae, Cyatheaceae,
Chloranthaceae, Myrtaceae and Cunoniaceae. The top ten most diverse families
are: Melastomataceae, Lauraceae, Rubiaceae, Euphorbiaceae, Myrtaceae,
Myrsinaceae, Cyatheaceae, Clusiaceae, Arecaceae and Chloranthaceae.
The main canopy species are the same as mentioned for the forest group of
Meliosma tachirensis and Alchornea grandiflora. Additionally, important canopy
species for the forest of this alliance are: cf. Elaeoluma nuda, Hieronyma
moritziana, Prunus moritziana, Ocotea leucoxylon, O. vaginans, Weinmannia
balbisiana and Zanthoxylum melanostinctum. Common subcanopy species are
Faramea killipii, Clethra fagifolia and Eugenia cf. tamaensis. In these forests
lianas and vines, like Mikania banisteriae, Diogenesia tetrandra, Mikania
nigropunctulata are also common. Very common shrubs in the understory are:
Cybianthus iteoides and Symbolanthus vasculosus.
Syntaxonomy: Two associations are recognized in this alliance, defined from 23
plots that include 228 species with dbh ≥ 2.5 cm belonging to 118 genera and 60
families.
Prunus moritziana and Zanthoxylum melanostictum are diagnostic in the canopy;
Faramea killipii and Clethra fagifolia in the subcanopy. For this alliance, Miconia
tovarensis, Rudgea tayloriae, the tree ferns Cyathea caracasana and C. pauciflora
and the treelets Anaectocalyx bracteosa and Cybianthus iteoides are also
considered diagnostic.
This alliance contains the subandean forest association of Croizatio brevipetiolatae
- Wettinietum praemorsae, and the Andean forest association of Schefflero
ferrugineae - Cybianthetum laurifolii.
Ecology and distribution: The alliance of Faramea killipii and Prunus moritziana
includes subandean and Andean forest communities present at altitudes between
1770 and 2600 m on the South slope, and from 1950 to ~2600 m on the North
slope.
3. Croizatio brevipetiolatae –Wettinietum praemorsae Cuello & Cleef 2009
Typus: Cuello Plot No. 1. Table 2.3, Fig. 2.4, 2.5.
Subandean forests of Croizatia brevipetiolata and Wettinia praemorsa
Physiognomy and composition: These forests are of medium stature and density.
They display a canopy that reaches between 15 to 25 m in height, with some
37
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
emergent trees of 26 to 30 m tall. Among the canopy species, besides those
mentioned for the alliance and higher group, Aniba cinnanomiflora, Elaeagia ruizteranii, Protium tovarense, Miconia lucida and Sloanea guianensis are more
prominent. The height of the subcanopy reaches between 5 and 15 m with the
dominance of the palm Wettinia praemorsa, occasionally reaching up to 25 m.
Croizatia brevipetiolata is very abundant in the subcanopy layer, together with
Palicourea angustifolia, Piper longispicum var. glabratum and Geissanthus
fragrans, among others. The parasitic Aethanthus nodosus is also a regular
occurance in the canopy.
Common epiphytic species are the ferns Asplenium auriculatum, A. cuspidatum, A.
harpeodes, Elaphoglossum cuspidatum, E. eximium, Hymenophyllum polyanthus,
Melpomene xiphopteroides, Polypodium fraxinifolium, P. funckii, Polytaenium
lineatum, Terpsichore subtilis, T. taxifolia and T. xanthotrichia. Bromeliads such
as Guzmania mitis, Mezobromelia capituligera and species of Aechmea, Racinaea
and Tillandsia are present. Epiphytic Philodendron fraternum is characteristic for
the canopy as well.
Among the species of the lower treelets layer (up to 5 m): Besleria pendula, the
little palms Geonoma jussieuana, Geonoma undata, the tree ferns Cyathea fulva,
C. pauciflora, C. kalbreyeri, C. caracasana, Dicksonia sellowiana and the
perennial large herbs Sphaeradenia laucheana and Anthurium nymphaeifolium
stand out.
Syntaxonomy: The association Croizatio brevipetiolatae - Wettinietum praemorsae is defined on the basis of twelve 0.1-ha plots which contained 154 species with
dbh ≥ 2.5 cm.
The diagnostic species in the canopy are: Wettinia praemorsa, Meriania
grandidens, and Miconia lucida; and Croizatia brevipetiolata as well as Wettinia
praemorsa in the subcanopy.
In this association one subassociation and one variant are recognized: the
subassociation of hedyosmetosum cuatrecazanum and the variant of Protium
tovarense.
Ecology and distribution: The forests of the Croizatia brevipetiolata and Wettinia
praemorsa association include a more or less homogenous zone between 1700 and
ca.~2300 m altitude along both slopes of Ramal de Guaramacal. The locally
known “mapora palm” (Wettinia praemorsa) is common, and these forests are
consequently known as “maporales”. These forests can display some variations in
composition and structure between 1700 and 1900 m on the southern slope (variant
of Protium tovarense), and between 2100 and ~2300 m on the northern slope
(subass. hedyosmetosum cuatrecazanum), generally however, they maintain a
more uniform composition between 1900 and 2200 m on both slopes.
Croizatio brevipetiolatae – Wettinietum praemorsae
3.1. subassociation hedyosmetosum cuatrecazanum Cuello & Cleef 2009
Typus: Cuello Plot No. 2. Table 2.3, Fig. 2.5.
Subassociation of Hedyosmum cuatrecazanum
38
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Physiognomy and composition: The canopy of the forest reaches between 10 and
25 m in height; emergent trees reaching up to 28 m. The composition is in
agreement with the description for the association. Casearia tachirensis,
Perrottetia quinduensis, Meriania macrophylla, Miconia theaezans var. longifolia,
Sapium stylare and Turpinia occidentalis are important in the canopy. Species that
may sometimes reach the canopy as well are: Aegiphila ternifolia, Hedyosmum
cuatrecazanum, Trichilia septentrionalis and the palm Wettinia praemorsa.
Among the smaller trees, 5 to 10 m tall, are: Croizatia brevipetiolata, Eugenia cf.
tamaensis, Geissanthus fragrans, Hedyosmum cuatrecazanum, Meriania
grandidens, Palicourea demissa, P. puberulenta, Piper longispicum var.
glabratum and Prunus moritziana. In the layer of small trees and shrubs from 2 to
5 m, the tree ferns Cyathea fulva and Sphaeropteris sp., as well as treelets of
Cestrum darcyanum, Gordonia fruticosa, Guarea kunthiana, Miconia
amilcariana, M. mesmeana subsp. longipetiolata, Persea peruviana, Saurauia
yasicae, Solanum confine and Trichilia hirta; the shrubs Cybianthus cuspidatus
and Ruellia tubiflora var. tetrastichantha and the low palms Geonoma orbigniana
and G. jussieuana were recorded.
Syntaxonomy: The subassociation of hedyosmetosum cuatrecazanum is defined
on the basis of three 0.1-ha plots with 72 woody species with dbh ≥ 2.5 cm.
The diagnostic species in the canopy are Casearia tachirensis, Hedyosmum
cuatrecazanum, Palicourea demissa and Sapium stylare. Diagnostic species in the
subcanopy are: Aegiphila ternifolia, Cestrum darcyanum, Croizatia brevipetiolata
and Trichilia septentrionalis.
Ecology and distribution: The subandean forests of the subassociation
hedyosmetosum cuatrecazanum are located between 2100 and 2300 m altitude on
the northern slope of Guaramacal. These forests (example at 2300 m) are on slopes
of 20-25% inclination, and on soils of variable depth (0 to 100 cm), with sandy
textures with 20-50% of coarse fragments, dark brown reddish colors in the
superficial layers (0 to 20 cm) and yellowish red colors in the deep layers (20-100
cm). pH increases with depth from 4 to 4.4; while the percentage of organic matter
diminishes from 7.5 to 3%.
3.2. Variant of Protium tovarense
Representative rel.: Cuello Plot No. 30. Table 2.3, Photo 2.3
Physiognomy and composition: The forests of the variant of Protium tovarense
are of medium stature and density. They display a dense canopy, of 15 to 25 m in
height with some emergent of trees of up to 30 m. Basically, they are made up by
the same species indicated for the association, but differing in the higher
abundance of Protium tovarense, Coccoloba cf. llewelynii, Weinmannia sorbifolia,
Hedyosmum cf. gentryii, Weinmannia glabra, Myrcia acuminata, Meliosma
tachirensis, Miconia lucida, among others. The subcanopy layer reaches between 8
and 12 m. Common species are Aiphanes stergiosii, Cyathea pauciflora, C. fulva,
C. kalbreyeri, Eschweilera perumbonata, Eugenia tamaensis, Matayba
camptoneura, Myrcia cf. sanisidrensis, Palicourea apicata and Weinmannia
39
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Flora, vegetation and ecology in the Venezuelan Andes
glabra. Further observed in this community are, the vine Cissus obliqua, the
epiphytes Columnea sanguinea, Mezobromelia capituligera; the climbing fern
Hypolepis nigricens, the shrubs Notopleura steyermarkiana, Symbolanthus
vasculosus, Cavendishia bracteata, the little palm Geonoma jussieuana and the
terrestrial clubmoss Huperzia reflexa.
Figure 2.4. Subandean forest of the association of Croizatio brevipetiolatae - Wettinietum
praemorsae. Plot 1: 1960 m, North slope. Ac: Aniba cinnamomiflora; Ag:
Alchornea grandiflora; Bp: Besleria pendula; Br: Billia rosea; Cb: Croizatia
brevipetiolata; Cc: Cybianthus cuspidatus; Er: Elaeagia ruiz-teranii; Gf:
Geissanthus fragrans; Mth: Miconia theaezans; Op: Ocotea puberula; Pa:
Palicourea angustifolia; Pp: Palicourea puberulenta; Pm: Persea meridensis;
Pl: Piper longispicum var. glabratum; Pb: Pouteria baheniana; Sc:
Symbolanthus calygonus; Sg: Sloanea guianensis; Wp: Wettinia praemorsa.
Syntaxonomy: The variant of Protium tovarense is defined on the basis of three
0.1-ha plots with a total of 75 species with dbh ≥ 2.5 cm. These forests are
distinguished by the presence of the character species Protium tovarense and
Weinmannia sorbifolia, as well as the palm Aiphanes stergiosii (6 to 12 m). This
40
The forest vegetation of Ramal de Guaramacal
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palm, although abundant in these forests, seems to have a very restricted
distribution since its existence is known only from this community from where the
species was originally described.
Ecology and distribution: The forests of the variant of Protium tovarense are
located between (1600) 1700 and 1900 m of altitude on the South slope of Ramal
de Guaramacal, sector Agua Fría, in the border zone between Portuguesa and
Trujillo states (“Alto de La Divisoria de La Concepción”) and above of the small
village “La Peña de Agua Fría”. These forests are on sites with very steep slopes
and shallow wet soils with a large amount of great rock fragments. Because of
their inaccesibility they remain very pristine forests. In the forests near Alto de La
Divisoria de La Concepción, trees of Podocarpus oleifolius could be observed
growing close to the top of the mountain to around 1900 m altitude.
Figure 2.5. Subandean forest of the association Croizatio brevipetiolatae – Wettinietum
praemorsae subassoc. hedyosmetosum cuatrecazanum. Plot 2: 2100 m, North
slope. Ag: Alchornea grandiflora; Bt: Beilschmiedia tovarensis; Ct: Casearia
tachirensis; Cb: Croizatia brevipetiolata; Dt: Dussia tessmannii; Er: Elaeagia
ruiz-teranii; En: Elaeoluma nuda; Gf: Geissanthus fragans; Hc: Hedyosmum
cuatrecazanum; Pd: Palicourea demissa; Pl: Piper longispicum var.
glabratum; Pm: Prunus moritziana; Pp: Palicourea puberulenta; Rp: Ruagea
pubescens; Th: Turpinia heterophylla; Wp: Wettinia praemorsa.
41
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Flora, vegetation and ecology in the Venezuelan Andes
Photo 2.3. Interior of the subandean forest of plot 30 of Croizatio brevipetiolatae –
Wettinietum praemorsae variant of Protium tovarense at 1875 m. South slope,
Agua Fría-La Peña sector.
4. Schefflero ferrugineae – Cybianthetum laurifolii Cuello & Cleef 2009
Typus: Cuello Plot No. 4. Table 2.3, Fig. 2.6, 2.7
Andean forests of Schefflera ferruginea and Cybianthus laurifolius
Physiognomy and composition: Humid cloud forests, with a canopy height of
between 8-18 m, with some emergent isolated trees up to 22 m. Among the most
common canopy species are: Eugenia cf. tamaensis, Ilex myricoides, Miconia
tinifolia, Ocotea aff. karsteniana, Podocarpus oleifolius var. macrostachyus,
Trichilia septentrionalis and Weinmannia glabra.
The community structure is dominated by a great abundance of small diameter
individuals (<10 cm dbh). Species with high abundance include Calyptranthes
meridensis, Clethra fagifolia, Critoniopsis paradoxa, Cybianthus laurifolius,
Eugenia cf. tamaensis, Faramea killipii, Hedyosmum crenatum, Ilex laurina,
Miconia ulmarioides, Myrsine coriacea, Palicourea apicata, Schefflera
ferruginea, Weinmannia glabra, among others. Hemiepiphytic trees like Clusia
alata and C. trochiformis are also common in the canopy. Epiphytes are very
abundant; among them: Pecluma divaricata, Peperomia peltoidea, Racinaea sp., a
diversity of orchid species of the genera Pleurothallis (P. semiscabra, P.
archidiaconi, P. siphoglossa, P. bivalvis, among others), and Stelis. The parasitic
Aetanthus nodosus and the hemiepiphytic climber Sphaeradenia laucheana are
also present, as well as several species of vines of the Ericaceae and Asteraceae
42
The forest vegetation of Ramal de Guaramacal
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families, such as Diogenesia tetrandra, Macleania rupestris, Mikania
nigropunctulata, M. stuebelii, Pentacalia vicelliptica and Themistoclesia
dependens.
In the lower layer, small melastomataceous trees (2-3 m tall) or shrubs abound,
such as: Anaectocalyx bracteosa, Miconia ulmarioides and M. suaveolens. The
tree ferns Cyathea fulva, C. caracasana, C. pauciflora and Dicksonia sellowiana
are very common. Colonies of the bamboo Rhipidocladum geminatum are also
very abundant.
Very common shrubs in the understory belong to Notopleura steyermarkiana. The
small palms Geonoma jussiaeana and G. orbigniana and the small shrub
Psychotria aubletiana are also frequent. Among the terrestrial herbs Lycopodium
jussiaei, Rhynchospora tuerckheimii, R. immensa and the terrestrial short-stemmed
fern Culcita coniifolia are distinguished.
Figure 2.6. Andean forest of the association Schefflero ferrugineae - Cybianthetum
laurifolii. Plot 4: 2450 m., North slope. Cl: Cybianthus laurifolius; Cf:
Cyathea fulva; Fk: Faramea killipii; Im: Ilex myricoides; Mt: Miconia
tinifolia; Po: Podocarpus oleifolius var. macrostachyus; Sf: Schefflera
ferruginea; Ta: Ternstroemia acrodontha; Wg: Weinmannia glabra.
43
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Flora, vegetation and ecology in the Venezuelan Andes
Syntaxonomy: The forest association Schefflero ferrugineae – Cybianthetum
laurifolii is defined based on eleven plots (ten of 0.1-ha and one of 0.03-ha) with
158 species with dbh ≥ 2.5 cm. Diagnostic species are: Cybianthus laurifolius,
Hedyosmum crenatum, Miconia ulmarioides, Myrsine coriacea and Schefflera
ferruginea.
Ecology and distribution: The forest community of the association Schefflero
ferrugineae - Cybianthetum laurifolii represents a transitional type between the
subandean and Andean forests, located over 2300 m altitude on the northern slope
and to even lower altitudes (2100 - 2250 m) in areas near crests or lower summits.
Characteristic of these sites is a greater density of thin-stemmed treelets with a low
canopy. A group of tree species of much smaller leaves like those of the Andean
forest, is also found intermingled with common large-leaved species of the
subandean forest. This transitional condition seems similar to that described by
Kelly et al. (1994).
Schefflero ferrugineae – Cybianthetum laurifolii
4.1. subassociation typicum Cuello & Cleef 2009
Typus: Cuello Plot No. 4. Table 2.3, Fig. 2.6
Subassociation of Cybianthus laurifolius
Physiognomy and composition: Forest structure and composition as described for
the association. Additionally important canopy species are: Dioicodendron
dioicum, Gaiadendron punctatum, Ilex truxillensis var. bullatisima, Meliosma
venezuelensis, Myrcia aff. guianensis, Ocotea sericea and Symplocos bogotensis.
Syntaxonomy: The forest Schefflero ferrugineae – Cybianthetum laurifolii
subassociation typicum is defined on seven plots, with 96 species with dbh ≥ 2.5
cm. The diagnostic species are Brunellia cf. integrifolia, Calyptranthes cf.
meridensis, Palicourea apicata and Panopsis suaveolens.
Ecology and distribution: The forests of the subassociation of Cybianthus
laurifolius are located between 2350 -2580 m on the North slope of Guaramacal
sector. This community was also observed throughout the mountain ridge between
2260 and 2570 m on the North slope of Agua Fría sector, and in the same
conditions at 2250 m on the North-West slope in El Santuario sector.
Schefflero ferrugineae – Cybianthetum laurifolii
4.2. subassociation miconietosum suaveolentis Cuello & Cleef 2009
Typus: Cuello Plot No. 6. Table 2.3, Fig. 2.7.
Subassociation of Miconia suaveolens
Physiognomy and composition: These forests have a very irregular canopy
between 8 and 15 m and a few emergent isolated trees that can reach up to 18 or 20
m. Some of the highest trees are: Alchornea grandiflora, Miconia theazans, M.
44
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
tinifolia, M. tovarensis and Prunus moritziana. Hemiepiphytic trees Clusia
trochiformis and C. alata, with multiple aerial roots are also abundant.
Common canopy species are Aegiphila moldenkeana, Clethra fagifolia,
Critoniopsis paradoxa, Cybianthus laurifolius, Hedyosmum translucidum,
Hedyosmum sp. A, H. crenatum, Ilex laurina, Weinmannia lechleriana and W.
fagaroides.
The presence of tree ferns is very common, being Cyathea pauciflora the most
abundant. Small trees of Melastomataceae, like: Miconia suaveolens, M.
mesmeana subsp. longipetiolata, M. theaezans and Anaectocalyx bracteosa; and
shrubs like Symbolanthus vasculosus, the bamboo Chusquea purdieana and the
palm Geonoma jussieuana, are frequent in the understory.
Figure 2.7. Andean forest of the association Schefflero ferrugineae - Cybianthetum
laurifolii. subassoc. miconietosum suaveolentis. Plot 6: 2500 m, South slope.
Ca: Clusia alata; Cf: Clethra fagifolia; Cp: Critoniopsis paradoxa; Cp:
Cyathea pauciflora; Gj: Geonoma jussieuana; Hs: Hyeronima scabrida; Mth:
Miconia theazans; Ms: Miconia suaveolens; Mto: Miconia tovarensis; Mv:
Meliosma venezuelensis; Hsp: Hedyosmum sp. A; Wl: Weinmannia
lechleriana. Rt: Rudgea tayloriae.
Syntaxonomy: The forests of subassociation miconietosum suaveolentis are
represented by four 0.1-ha plots with 73 species with dbh ≥ 2.5 cm. These forests
differ from those of the subassociation typicum by the diagnostic presence of
Critoniopsis paradoxa, Hedyosmum sp. A, Hyeronima scabrida and Miconia
45
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Flora, vegetation and ecology in the Venezuelan Andes
suaveolens, which, with the exception of Critoniopsis paradoxa, are absent from
that subassociation.
Ecology and distribution: The forests of the subassociation of Miconia
suaveolens are located between 2100 and 2600 m on the South slope of
Guaramacal sector. These rain forests are located in areas with very steep slopes
apparently affected more by landslides. Mass movements are frequent and widely
observed at the highest elevations of the South slope triggered by the high
precipitation.
In general these forests have a low diversity of species; some species common to
the stable sites at the same altitude on the North slope are absent. A greater
abundance of species with stilt roots, indicative of plant adaptation to steep
surfaces under a wet climate, are present (Cleef et al. 1984).
RUILOPEZIO PALTONIOIDES – CYBIANTHION MARGINATI Cuello & Cleef 2009
Typus: Geissantho andini - Miconietum jahnii. Table 2.3
High Andean forests of the Ruilopezia paltonioides - Cybianthus marginatus alliance
Physiognomy and composition: Dense and low rain forests with a canopy of 5 (3)
to 12 (14) m in height, conformed by thin-stemmed, small-leaved trees.
Asteraceae, Ericaceae, Myrsinaceae, Melastomataceae, Cunoniaceae and
Aquifoliaceae are the most diverse and predominant families. These forests share
the diagnostic canopy species Cybianthus marginatus, Ilex guaramacalensis,
Miconia tinifolia, Myrsine dependens, Symplocos tamana and the espeletinioid
species Ruilopezia paltonioides. Other common species are the small trees
Monochaetum discolor, Pentacalia cachacoensis, Vaccinium corymbodendron, the
tree fern Blechnum schomburgkii in the understory, and the liana Themistoclesia
dependens.
Syntaxonomy: Three associations are recognized in this alliance, defined from 11
variably sized plots that include 69 species with dbh ≥ 2.5 cm, belonging to 47
genera and 28 families of vascular plants.
The forest communities grouped in this alliance are (1) Geissantho andini Miconietum jahnii, with a possible subassociation of Freziera serrata; (2)
Gaultherio anastomosantis - Hesperomeletum obtusifoliae association, and (3) the
conspicuous dwarf forest association Libanothamnetum griffinii.
Ecology and distribution: The alliance of Ruilopezia paltoniodes and Cybianthus
marginatus groups Andean and high Andean forests located between 2750 and
2950 m direct under the summit zone of Ramal de Guaramacal. This altitudinal
zone is characterized by high relative humidity, permanent fogs and frequent rain
showers, indicated by a high cover of epiphytic mosses and liverworts.
5. Geissantho andini – Miconietum jahnii Cuello & Cleef 2009
Typus: Plot No. 37. Table 2.3, Fig. 2.8, Photo 2.5
46
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
Andean/high Andean forests of Geissanthus andinus and Miconia jahnii
Physiognomy and composition: These Andean/high Andean forests are of low
stature with a high density of individuals, having an open lowermost layer and a
thick litter layer. A high density of epiphytes, mainly ferns, orchids, mosses and
liverworts is a characteristic feature.
The canopy reaches up to 6 to 14 m in height with some emergent trees of up to 16
m, among them are: Ilex guaramacalensis, Miconia jahnii, Myrsine dependens and
Symplocos tamana, which are also the most abundant in the canopy together with
Cybianthus marginatus, Geissanthus andinus, Weinmannia lechleriana and
Oreopanax discolor, among others. In these forests lianas and vines are very
common, such as: Fuchsia membranacea, Pentacalia theaefolia, Thibaudia
floribunda, Disterigma alaternoides, Mikania stuebelii, Themistoclesia dependens
and Pentacalia vicelliptica. Striking are the vascular epiphytes: Guzmania
squarrosa, Odontoglossum schillerianum, Raccinaea tetrantha, and several fern
species of Polypodium and Asplenium. The bamboo Chusquea angustifolia can be
found inside the forest, forming dense clumps with multiple culms that can reach
up to 8 m in height and 3 cm in diameter.
The presence of the shrub Macrocarpaea bracteata, rosettes of Bromeliaceae
(Cuello 2816) and low shrubs like Psychotria dunstervillorum are common in the
understory. Trailing herbs like Hydrocotyle venezuelensis and Drymaria ovata; the
terrestrial orchid Cranichis antioquensis growing on the litter, and the tall erect
Cyrtochilum megalophium with inflorescences of up to 2 m in length; small
crawling herbs like Sibthorpia repens and species of Pilea and Rhynchospora
guaramacalensis and Carex jamesonii are additionally common occurances.
Syntaxonomy: The association of Geissantho andini - Miconietum jahnii is based
on four plots (three of 0.1 ha and one of 0.04 ha) with 53 species with dbh ≥ 2.5
cm. These low forests are distinguished by the presence of Miconia jahni,
Geissanthus andinus and Weinmannia lechleriana as diagnostic canopy species. In
this association, a subcommunity of Freziera serrata is distinguished.
Ecology and distribution: The humid dwarf forests of Geissanthus andinus and
Miconia jahnii cover considerable spatial extent in the zone of the Páramo El
Pumar to the center-west summit of Ramal de Guaramacal, at 2800-2950 m.
Geissantho andini – Miconietum jahnii
5.1. Subcommunity of Freziera serrata
Physiognomy and composition: The community of Freziera serrata concerns a
dense forest, which displays a higher canopy than the forests of the association
Geissantho andini - Miconietum jahnii, reaching 6 to 14 m with some emergent
trees up to 18 m. Besides of the listed ones for the association, the most common
canopy species are: Freziera serrata, Hedyosmum translucidum, Weinmannia
auriculata, W. karsteniana, and Podocarpus oleifolius var. macrostachyus.
47
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Flora, vegetation and ecology in the Venezuelan Andes
Under the canopy are dense colonies of the high bamboo Rhipidocladum
geminatum and small trees like Cybianthus laurifolius, Miconia mesmeana subsp.
longipetiolata, Baccharis brachylaenoides, Cestrum darcyanum, Oreopanax
discolor, the tree fern Cyathea fulva, and the shrub Macrocarpaea bracteata. Also
observed are the vine Passiflora truxillensis and the epiphytic fern Terpsichore
xanthotrichia. In the underbrush dense dwarf bamboo colonies of Neurolepis
glomerata and individuals of Rhynchospora guaramacalensis are present.
Syntaxonomy: The Andean forest of the Freziera serrata community could be
considered as a subassociation of the Geissantho andini - Miconietum jahnii. This
community is recognized from a single 0.1-ha plot with 40 woody species of dbh ≥
2.5 cm. The diagnostic canopy species are: Freziera serrata, Hedyosmum
translucidum, Weinmannia auriculata and W. lechleriana.
Figure 2.8. High Andean forest of the association Geissantho andini - Miconietum jahnii.
Plot 37: 2890 m. Páramo El Pumar. Cm: Cybianthus marginatus; Cha: Chusquea angustifolia; Da: Disterigma alaternoides; Dg: Drimys granadensis; Ga:
Geissanthus andinus; Ig: Ilex guaramacalensis; Md: Myrsine dependens; Mj:
Miconia jahnii; Mt: Miconia tinifolia; Vc: Vaccinium corymbodendron.
Ecology and Distribution: The subcommunity of Freziera serrata is found at
2750 m in an area of very steep slopes and difficult access on the North slope of
Ramal de Guaramacal. This forest stand is bounded on one side by the road
leading to the antennas at the summit, and by a clearing made for maintenance
below the electricity cables that lead to the antennas (Photo 2.4).
48
The forest vegetation of Ramal de Guaramacal
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Photo 2.4. (left) Aspect of the forest stand of the association Geissantho andini Miconietum jahnii subcommunity of Freziera serrata at 2750 m on the North
slope.
Photo 2.5. (right) Interior of plot 37 of the Andean forest of the Geissantho andini Miconietum jahnii at 2890 m in Páramo El Pumar.
6. Libanothamnetum griffinii Cuello & Cleef 2009
Typus: Cuello Plot No. 38. Table 2.3, Fig. 2.9, Photo 2.6, 2.7
High Andean dwarf forest of Libanothamnus griffinii
Physiognomy and composition: This community is represented by very dense
dwarf forests with a conspicuous broad-leaved white-grayish canopy conformed
by thin-stemmed small trees of 3 to 5 m high, and dominated by a great density of
the espeletinioid species Libanothamnus griffinii. Other species are scarce and include: Clethra fagifolia, Cybianthus marginatus, Miconia jahnii, M. tinifolia, Monnina sp., Monochaetum discolor, Palicourea jahnii, Weinmannia auriculata, W.
karsteniana and W. lechleriana.
The high cover of liverworts is noticeable, mainly comprising species of
Plagiochila (Cuello 3040, 3043), further species of Riccardia and of filmy ferns of
Trichomanes sp. (Cuello 2938), and Lellingeria myosuroides over tree trunks.
The understory is open and species poor. There are dispersed colonies of the
terrestrial fern Culcita coniifolia, individuals of the slender Eriosorus flexuosus
and the tree fern Blechnum schomburgkii. The orchid Brachionidium tuberculatum, the clubmoss Huperzia sp. (Cuello 2822), and the stoloniferous Psychotria dunstervillorum, as well as dense cushions of bryophytes like Plagiochila
49
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Flora, vegetation and ecology in the Venezuelan Andes
sp. (Cuello 3042) and Sphagnum meridense grow on top of a thick litter layer.
Towards the forest edges of the upper forest line the terrestrial orchid Cyrtochilum
ramosissimum, ground rosettes of species of Puya and Greigia and the endemic
sedge Rhynchospora guaramacalensis can be found.
Figure 2.9. High Andean dwarf forest of the association of Libanothamnetum griffinii. Plot
38: 2810, North slope. Cm: Cybianthus marginatus; Lg: Libanothamnus
griffinii; Mt: Miconia tinifolia; Pc: Pentacalia cachacoensis; Pj: Palicourea
jahnii; Wa: Weinmannia auriculata; Wk: Weinmannia karsteniana.
Photo 2.6. Exterior aspect of a high Andean dwarf forest of the Libanothamnetum griffinii
at the upper forest line (~2800-2900 m) on North slope of Ramal de
Guaramacal.
50
The forest vegetation of Ramal de Guaramacal
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Syntaxonomy: This association is based on four plots with 27 species with dbh ≥
2.5 cm, Libanothamnus griffinii being the character species.
Photo 2.7. Interior of plot 38 of the high Andean dwarf forest of the Libanothamnetum
griffinii at 2810 m on the North slope.
Ecology and distribution: The Libanothamnus griffinii dwarf forests are present
over large extensions along the upper forest line. They grow over convex slopes
with inclinations of between 20-30 degrees. On the North slope, they are observed
to extend continuously from lower elevations (2800 m) with a taller stature and
closer canopy cover than on the South slope, where their presence appears to start
at around 3000 m.
On the northern slopes, the Libanothamnus griffinii forests are observed to grow in
wind protected areas, on relatively deep soils with presence of small coarse
fragments (1 cm diameter) from 70 cm depth, and clay-sandy textures, with gray
colors in the upper layer, turning light to dark brown and dark reddish with depth.
On the wind exposed southern slopes, Libanothamnus griffinii treelets grow
shorter with altitude forming a more open canopy. Here they occur on relatively
shallow ground, with rock fragments from 45 cm depth; with clay to clay-sandy
51
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Flora, vegetation and ecology in the Venezuelan Andes
textures and gray colors turning from spotted orange to dark brown with depth.
Libanothamnus griffinii can also form shrub páramo communities with other páramo species as described for the Páramo vegetation of Guaramacal in Chapter 3.
7. Gaultherio anastomosantis – Hesperomeletum obtusifoliae Cuello & Cleef 2009
Typus: Cuello Plot No. 34. Table 2.3, Fig. 2.10, Photo 2.8, 2.9
High Andean dwarf forests of Gaultheria anastomosans and Hesperomeles obtusifolia
Physiognomy and composition: The dwarf forests of the Gaultheria
anastomosans - Hesperomeles obtusifolia association are very low in stature, very
dense in the number of individual treelets and exhibit an almost closed cover
allowing little light penetration. The canopy reaches between 4 and 6 m in height
with some emerging individuals reaching 8-10 m, such as: Miconia tinifolia and
Weinmannia lechleriana and the stem rosettes of Ruilopezia paltonioides. The
most abundant species of the canopy are (in order of abundance) Cybianthus
marginatus, Hesperomeles obtusifolia, Vaccinium corymbodendron, Myrsine
dependens, Gaultheria anastomosans, Diplostephium obtusum, Miconia tinifolia
and Pentacalia cachacoensis. Less abundant are, Ilex guaramacalensis, Monnina
sp., Oreopanax discolor, Pentacalia greenmanniana and Symplocos tamana. Vines
and lianas are very common, especially those of the Ericaceae family. The most
abundant include: Disterigma alaternoides, Psammisia hookeriana, Themistoclesia
dependens and Thibaudia floribunda. Epiphytic ferns are also very abundant;
Grammitis sp. and Melpomene flabelliformis grow among a dense cover of
bryophytes. Most prominent are Campylopus trichophorus, Herbertus acanthelius,
and species of Lepidozia and Plagiochila.
Figure 2.10. High Andean dwarf forest of the association of Gaultherio anastomosantis Hesperomeletum obtusifoliae. Plot 34: 3050 m. Bs: Blechnum schomburgkii;
Cha: Chusquea angustifolia; Cm: Cybianthus marginatus; Dv: Diplostephium obtusum; Ga: Gaultheria anastomosans; Gsp: Greigia sp.; Ho: Hesperomeles obtusifolia; Md: Myrsine dependens; Mt: Miconia tinifolia; Psp.
Puya sp.; St: Symplocos tamana.
52
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
The open understory, with a conspicuous thick litter layer, is poor in species. Only
in more open sites are colonies of the tree fern Blechnum schomburgkii and
Bromeliaceae (Cuello 2816). Further may be noted: dispersed individuals of the
terrestrial orchid Gomphichis costaricense, patches of stoloniferous dwarf shrub
Psychotria dunstervillorum growing in the litter layer, and bamboo clumps of
Chusquea angustifolia and Neurolepis glomerata. A diversity of bryophytes
growing on the base of trunks and over tree roots, are distinguished, such as Bryum
grandifolium, Campylopus pilifer, C. nivalis, Dicranum frigidum, Herbertus juniperinus, Leptodontium longicaule, Plagiochila cf. aerea, Plagiochila sp., Scapania
portoricensis, Sphagnum meridense and Tetraplodon mnioides, among others.
Syntaxonomy: The Gaultherio anastomosantis - Hesperomeletum obtusifoliae
association, is based on three plots with 31 species with dbh ≥ 2.5 cm.
The diagnostic species of this association are Diplostephium obtusum, Gaultheria
anastomosans and Hesperomeles obtusifolia. Cybianthus marginatus besides of
being diagnostic for the alliance is also diagnostic for the association.
Photo 2.8. Aspect of the humid high Andean dwarf forests on the South slope of Ramal de
Guaramacal. Southeast of „Las Antenas‟ area, 2900-3000 m.
Ecology and distribution: The dwarf forests of the Gaultherio anastomosantis Hesperomeletum obtusifoliae association are situated in the summit areas of Ramal
de Guaramacal, between 2950 and 3050 m. They form patches, or islands, of forest
among the páramo vegetation, especially over, wind protected concave areas on
the North slope. On South facing slopes these forests are observed growing on larger extensions of continuous forest on steep surfaces with slopes up to 22 degrees.
They are found on shallow soils with the presence of rock fragments from 15 cm
53
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Flora, vegetation and ecology in the Venezuelan Andes
depth and bed rock at 45 cm, with clayey-loamy textures, dark gray in color
turning to more clear with orange spots at greater depths, and pH 3.6 to 3.9 in the
upper soil layers rising to 4.1 to 4.9 at increased depth.
Together with the dwarf forest of Libanothamnetum griffinii these forests constitute the upper forest line on Ramal de Guaramacal
Photo 2.9. Interior of plot 34 of the high Andean dwarf forest of the Gaultherio
anastomosantis - Hesperomeletum obtusifoliae at 3050 m.
2.5 DISCUSSION
Forest phytosociological classification and methodology limitations
The phytosociological classification of the montane forests of Ramal de
Guaramacal has resulted in three new alliances and seven associations.
Subassociations are described for four associations. Variants are still to be
confirmed; only one variant was described.
Classes and orders cannot yet be defined on the basis of the present number of
relevés and the information available in Table 2.3, and, due to the lack of data
from montane forests in the region and elsewhere in Venezuela and adjacent
Colombia. It is to be expected that subandean forests (LMRF), and Andean - high
Andean forests (UMRF-SARF) respectively, as ecologically very different
ecosystems, will belong to separate orders and classes of equatorial montane rain
forests. A major forest group of Meliosma tachirensis - Alchornea grandiflora was
recognized, which could be considered as equivalent to order. However, the
species in common in that group could also belong to a supra-class unit (Table
54
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
2.3), which covers the wide-spread formation of equatorial montane forests of the
Andes. This is the main reason we do not propose a proper syntaxon for the order
level. A surprise was the Farameo killipii - Prunion moritzianae, which includes
forest associations of both LMRF and UMRF (see below). This is the first time
that LMRF in the lower part and UMRF in the upper part were included into one
phytosociological alliance. Thus far, as experienced in the seven studied altitudinal
transects of ECOANDES programme in Colombia, the alliances only accounted
for either LMRF plots or UMRF-SARF plots. There may be two reasons: (1) the
steep slopes of Guaramacal with bamboo páramo on top (3120 m) result in a type
of „compressed forest zonation‟. In a short altitudinal interval under almost
permanent high environmental humidity with only slight temperature change sharp
vegetation limits/borders may become obscured. We think this is the most likely
explanation. (2) The other possibility is that the alliance is an artifact due to a lack
of phytosociological resolution; more relevés in this altitudinal interval could
provide more information for the forest classification and altitudinal zonation.
The forest vegetation has been described on the basis of a relatively low number of
relevés. Only outside the Guaramacal sector are they not homogeneously
distributed over the altitudinal gradient. Although most plots were 1000 m2
(significantly larger as those of mostly 500 m2 in the ECOANDES transects in
Colombia), some of the forest plots between 2800-3000 m were of 100 to 400 m2.
The experience of the second author in the UMRF-SARF plots in Colombia and
North Ecuador suggests this corresponds rather to the minimum area. In tropical
montane rain forests species diversity decreases with altitude. The 0.1 ha plots of
this study most probably do not represent the minimum area for LMRF, but are
apparently sufficiently representative for the UMRF sampling (see also Westhoff
& Van der Maarel 1973). In the SARF dwarf forests, we used smaller plot size
also according to the conditions of the forest cover of predominantely very steep
terrain. The ECOANDES forest plot size of 500 m2 (0.05 ha) was established for
practical reasons. Because most of the sites were very remote and difficult to
reach, the plot size of 500 m2 corresponded more to the effort and time required to
sample a plot of this size in one day by 2-3 researchers, representing a more
practical approach (Cleef et al. 1984).
The methodology adopted includes only the census of woody species with
diameter ≥ 2.5 cm. Thus, forest communities are defined based mainly on
diagnostic tree species from understory and canopy, respectively, rather than on
other growth forms.
In our opinion however, the resulting forest classification is clearly visible for the
montane forests of Ramal de Guaramacal. It is the first attempt for
phytosociological classificaction of montane rain forests of the Venezuelan Andes
based on a quantitative data set for an entire mountain range.
Altitudinal zonation
The altitudinal zonation of the montane forests of Ramal de Guaramacal is
depicted in Fig. 2.11. TWINSPAN classification for montane forest plots of Ramal
de Guaramacal arranged forest types in Table 2.3 according to the altitudinal
55
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
gradient. Based on physiognomy and floristic composition, these forest types can
easily be grouped into zones corresponding to LMRF, UMRF and SARF classes of
Grubb (1977), or to subandean, Andean and high Andean forests, respectively,
according to Cuatrecasas (1934, 1958); also having been applied by Cleef et al.
(1984, 2003) and Rangel-Ch. et al. (2003, 2005, 2008) for the ECOANDES
transect studies. Elsewhere in the Colombian Andes, Rangel-Ch. & Franco-R.
(1985), Rangel-Ch. & Lozano (1989) and Rangel-Ch. (1994) have published
details on montane forest transects. For Ecuador, reference is made to forest
transect studies by e.g. Bussmann (2002), Lauer et al. (2001) and Moscol & Cleef
(2009b).
The first division of the TWINSPAN classification of Guaramacal montane forests
separates the less diverse Andean and dwarf high Andean forest (UMRF-SARF)
communities above 2750 m from the species-rich subandean-andean (LMRF and
UMRF) communities present up to 2600 m. The second division separates the
lower from the upper subandean (LM) forests and includes, in the second group, a
forest type belonging more to Andean (UMRF) forest.
LMRF of Ramal de Guaramacal can be found from 1350 m on the South slope
and from 1650 m on the North slope in some parts of the mountain range.
However, most LMRF extends from 1800 to about 2300 m. The limit of 1800 m is
determined by the Park boundaries, below that, disturbed areas occupy the LMRF
zone, especially on the North slope. UMRF is present from 2300 to ~2800 m on
the North slope of Guaramacal sector; on top of small ranges on South or NorthWest slopes UMRF is also present near 2100 m. SARF in Ramal de Guaramacal is
present at the same altitude as páramo vegetation, from 2800 to 3050 m. Most
probably exposition vs. protection to strong trade wind, low temperature and
extremes and availability of substrate to support a dwarf forest may be responsible
(see forthcoming paper Cuello et al. in prep.).
Forest zonation is variable between the North and South slope of Guaramacal. Fig.
2.11 shows that on the windward South slope, forest zones of UMRF tend to reach
lower elevations than on the opposite and drier North slope. In the first instance,
temperature is probably most accountable for this phenomenon. Almost permanent
humidity prevents higher temperatures and causes slightly lower values for the
medium annual temperature. Also the frequent landslides on the steeper and wetter
slopes at mid-high elevation may play a role. LMRF forests also display a lower
position as longer gradient and more forest extensions are present below 1800 m.
This asymmetric configuration of forest zones on equatorial mountains has also
been reported elsewhere (e.g. Kappelle et al. 1995; Cleef 1981). In general, there
are dry and a humid to wet slopes opposing each other.
Furthermore, it is noticeable that there is a low altitudinal upper limit of the forest
(Upper Forest Line or UFL) exists in Ramal de Guaramacal, apparently caused by
the “top effect” (Grubb 1971) with UMRF (including SARF) found at lower
altitude (Grubb 1977). Previously, this phenomenon was also known as the
„telescope‟ effect of mountain mass elevation (Van Steenis 1961, 1972) and, when
looked at from a different angle, the „Massenerhebung‟ effect (Schröter 1926). In
the Mérida Andes, the upper forest line is situated at an average elevation of
around 3400 m (Monasterio 1980; Schneider 2001), a vertical difference of some
56
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
350 m compared to Guaramacal UFL. With a lapse rate of 0.6 oC per 100 m
altitudinal interval would indicate a mean annual temperature of about 2 oC colder
in the summit zone of Guaramacal range. The first temperature records of the
Davis Pro 2 climate station installed near the summit of Guaramacal (3100 m) by
the first author since December 2006, registered a diurnal temperature variation
from 4-6 oC to 14-16 oC; the lowest temperatures recorded being between 1.3-4 oC;
the highest between 16-18 oC, with a mean temperature of 8.6 oC between January
and June 2007. In a forthcoming study (Cuello et al, in prep.) after the completion
of a year climate measurements, we hope to provide more detail on the low UFL
phenomenon.
Also, differences in humidity from the drier North and the wetter South slope
affect the altitudinal position of vegetation zones between slopes (Fig. 2.11).
Another effect of the low altitudinal position of the UFL is the compression of the
montane forest zones; they are situated in shorter vertical distances (Fig. 2.11).
The sequence of forest zones along the steep South slope is shortest in distance.
Figure 2.11. Semi-schematic profile of Ramal de Guaramacal, Andes,Venezuela. The
altitudinal zonation of montane forests and bamboo páramo along the North
and South slopes is depicted with the respective plots numbers. Vertical
exaggeration 5.0 x.
57
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
WSW of Bogotá, the UFL has been located at 1900-2000 m during the coldest
phase of the Last Glacial Maximum (Hooghiemstra & Van der Hammen 1993). It
is most probable that during the same cold period of the Last Glacial, conditions in
the Venezuelan study area were almost similar, likely resulting in a much more
compressed altitudinal montane forest zone relative to that of today.
Paleoecological studies of lake sediments, e.g. the promising peat land at c. 2000
m near the Park Rangers house of the Guaramacal National Park, may provide
more clues.
Forest composition and diversity
Montane forests of Ramal de Guaramacal show a floristic composition and
diversity that change along altitude. Family composition shows the same trend as
observed in other Andean forests (Gentry 1992, 1995; Rangel-Ch. 1991). In Lower
Montane Rain Forests of Guaramacal, Rubiaceae, Lauraceae and Melastomataceae
are the most speciose of woody families. In Upper Montane Rain Forests, the
Lauraceae family is still the most diverse, followed by Melastomataceae and
Myrtaceae, while in Subalpine rain forests the Asteraceae and Ericaceae are the
most species rich families (Table 2.4).
Table 2.4. Most species-rich families by forest zones.
LMRF
(1300~2300 m)
Family
Rubiaceae
Lauraceae
Melastomataceae
Myrtaceae
Euphorbiaceae
Piperaceae
Myrsinaceae
Cyatheaceae
Meliaceae
Arecaceae
Moraceae
Sapindaceae
Solanaceae
67 families
UMRF
~2350 ~2900 m
#
spp.
29
24
22
17
10
8
7
7
6
6
6
6
6
266
SARF
(2800 – 3060 m)
Family
Lauraceae
Melastomataceae
Myrtaceae
Asteraceae
Rubiaceae
Ericaceae
Myrsinaceae
Cyatheaceae
Euphorbiaceae
Cunoniaceae
Aquifoliaceae
#
spp.
19
13
13
11
9
8
7
6
6
6
5
51 families
169
Family
Asteraceae
Ericaceae
Myrsinaceae
Melastomataceae
Cunoniaceae
Araliaceae
Polygalaceae
Rosaceae
20 families
#
spp.
11
7
5
5
4
2
2
2
50
Species diversity and composition also change along the altitudinal gradient with
some variations caused by slope exposure and sectors. Species richness generally
decreases with elevation; however, local increase in species richness per 0.1 ha
58
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
plot was observed between 2300-2400 m on the North slope of Guaramacal in the
LMRF - UMRF limit zone (Table 2.2). This diversity trend and its relation to
increasing humidity with elevation from the dry interandean Boconó valley to the
top of the mountain has been previously discussed (Cuello 1996, 2002). However,
the mid slope diversity peak for bryophytes and lichens reported by Wolf (1993)
may not be ruled out. Here we also confirm, as Schneider (2001) reported for the
first time from a montane forest transect near Mérida, a mid slope diversity bulge
for vascular species at the transition from LMRF to UMRF.
Lower limit of LMRF is represented by the Simiro erythroxylonis - Quararibeetum
magnificae, which, on the North slope, shows a distinct set of species as in the
bunchosietosum armeniacae subassociation, while on the South slope the vicariant
forest type is represented by that of the typicum subassociation.
Characteristic is the LMR forest of the Croizatio brevipetiolatae - Wettinietum
praemorsae, which is present on both slopes of Guaramacal with the same
altitudinal range. However, a LMR forest variant of Protium tovarense is
obviously characteristic for the South slope and the LMR forest of the
subassociation hedyosmetosum cuatrecazanum is present at the uppermost limit of
LMRF zone on the North slope. The composition of the forest variant of Protium
tovarense is also indicative of the high atmospheric humidity occurring on the
South slope. The presence of some large-leaved species of Hedyosmum, of
Weinmannia and several species of Cyathea are some examples.
UMR forest is represented by the Schefflero ferrugineae - Cybianthetum laurifolii
and the subcommunity with Freziera serrata of the Geissantho andini Miconietum jahnii. On the South slope the lower limit of UMRF is observed at
2100 m, being represented by the miconietosum suaveolentis of the Schefflero
ferrugineae - Cybianthetum laurifolii; on the North slope at 2350 m by the typical
subassociation, which is also found at 2250 m on the North-West slope. The forest
of the typical subassociation on the North slope presents a mixed composition with
a set of species from both LM and UMRF converging in these forests.
Rain forest of the subcommunity of Freziera serrata of the Geissantho Miconietum at 2750 m, represents the upper limit of the UMRF on the North slope
in Guaramacal sector. UMR forest association of Geissantho - Miconietum can
reach altitudes of up to 2890 m, as observed in Páramo El Pumar at the center-west
summit of Ramal de Guaramacal.
Distinctive forest composition is also noticeable in the SARF zone (2800-3050 m)
of UMRF, where two low species diverse dwarf forest associations, one of
Libanothamnus griffinii and the other of Gaultheria anastomosans and
Hesperomeles obtusifolia with high density of Cybianthus marginatus combined
with bamboo páramo vegetation to characterize the upper forest line. SARF, in our
opinion, belongs as a subzone to the domain of UMRF, as not only ecology but
also floristics and soil characteristics are shared as have been shown by the
ECOANDES studies (Van der Hammen et al. 1983-2008).
59
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Forest structure
The structure of the montane forests of Ramal de Guaramacal becomes more
compressed towards higher elevations. With an increase of altitude, an increase in
stem density and a decrease in stem diameter and canopy height is also observed
(Table 2.2 a,b, Fig. 2.12). LMRF are dense and of medium height, with canopies
up to 25 m tall, while UMRF canopies can reach up to 18 m, and those of SARF
are only 6-8 (10) m tall.
Basal area was slightly increased on the North than on the South slopes and shows
different patterns against altitude between slopes. On the South slope basal area
decreases with altitude, while on the North slope still high values have been
documented between 2300-2400 m (Fig. 2.12). This lower basal area is probably
due to the effect of disturbance by landslides on the steeper South slope.
Diversity and density of growth forms also varies with elevation among vegetation
zones (Table 2.5). More diversity and density of palms, lianas and climbers is
clearly observed in LMRF. Although diversity and the density of lianas decrease
with altitude, an important, and substantial, percentage of the total species richness
of SARF forest is represented by liana species (16.3%). There might be a
relationship with the obviously increased forest dynamics as a consequence of
steep slopes and consequently landslides. Hemiepiphytic trees as Clusia are
present in both LMR and UMR forests, but with a greater density in UMRF.
Density of tree ferns decreases with elevation, yet more diversity of tree ferns is
observed in UMRF, but the tree ferns are definitely more conspicuous in LMRF.
Table 2.5. Number of species and individuals of different growth forms in altitudinal zones
of montane forests of Ramal de Guaramacal.
LMRF
UMRF
SARF
Growth forms
Total trees≥ 2.5 cm
Trees ≥10 cm
Tree ferns
Hemiepiphytic trees
(Clusia)
Large herbs (incl.
ferns and cyclant.)
Bamboos
Climbers
Lianas
Palms
Shrubs
Stem rosette
Total
Total
area
of
samples (ha)
Spp.
#
(%)
215
(77.9)
168*
78.1*
7
(2.5)
Indiv.
#
(%)
5615
(72.0)
1430
25.5*
325
(4.2)
#
145
94
8
Spp.
(%)
(77.5)
64.8*
(4.3)
Indiv.
#
(%)
3876 (81.8)
833 21.5*
216 (4.6)
#
37
27
2
Spp.
Indiv.
(%)
#
(%)
(75.5) 2128 (92.2)
73.0* 316 14.8*
(4.1)
63
(2.7)
3
(1.1)
113
(1.4)
3
(1.6)
339
(7.2)
-
-
-
-
4
1
3
37
6
**
276
(1.4)
(0.4)
(1.1)
(13.4)
(2.2)
**
-
149
1
39
223
1338
**
7803
(1.9)
(0.0)
(0.5)
(2.9)
(17.1)
**
-
2
3
2
19
4
1
187
(1.1)
(1.6)
(1.1)
(10.2)
(2.1)
(0.5)
-
33
136
4
88
44
3
4739
(0.7)
(2.9)
(0.1)
(1.9)
(0.9)
(0.1)
-
1
8
**
1
49
(2.0)
(16.3)
**
(2.0)
7
67
**
44
2309
(0.3)
(2.9)
**
(1.9)
2.2
1.13
0.36
*percentage of the total trees≥2.5 cm; ** shrubs present in LM and SARF were <2.5 cm diameter.
60
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
(a)
9
LMRF
UMRF
8
7
6
N
Basal area (m2)
S
5
4
3
2
1
13
30
14
50
15
50
16
00
17
70
18
00
18
50
18
75
18
80
19
50
19
60
20
70
21
00
21
25
21
70
23
00
23
50
24
00
24
70
24
80
25
80
27
50
28
70
28
90
0
Altitude (m)
(b)
700
UMRF
LMRF
600
N
S
Individuals
500
400
300
200
100
28
90
28
70
27
50
25
80
24
80
24
70
24
00
23
50
23
00
21
70
21
25
21
00
20
70
19
60
19
50
18
80
18
75
18
50
18
00
17
70
16
00
15
50
14
50
13
30
0
Altitude (m)
Figure 2.12. Distribution of basal area (a) and number of woody individuals (b) per 0.1-ha
plots of montane forest for the North and South slope of Ramal de
Guaramacal, Andes, Venezuela.
61
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Flora, vegetation and ecology in the Venezuelan Andes
Comparison with other montane forests in the Venezuelan Andes
Few studies on composition and diversity of montane forests in the Venezuelan
Andes are available for comparison with the montane forests of Ramal de
Guaramacal. No recent studies, with the exception of the few and local studies of
Vareschi (1953), address the phytosociological classification and description of
vegetation communities in the Venezuelan Andes. Only general descriptions of
UMRF-SARF of the Mérida Andes s.l. have been delivered by Monasterio (1980)
and Veillon (1955). Berg & Suchi (2000) also reported a forest of Podocarpus
oleifolius up to 3100-3200 m altitude, as well a dwarf forest (SARF) community of
Libanothamnus cf. lucidus, Ugni myricoides and Cybianthus marginathus in
Páramo La Aguada in Sierra Nevada National Park, Mérida state. Bono (1996)
delivered species lists of montane forests of Táchira state, mostly structured
according to vegetation layers.
The UMRF association of Schefflero ferrugineo - Cybianthetum laurifolii in
Guaramacal displays floristic and physiognomical affinities with UMRF in Mérida
state, as those forest characterized by the presence of Podocarpus oleifolius var.
macrostachyus described for Mérida state in La Mucuy around 3000 m (Vareschi
1953), Sierra Nevada National Park between 3000-3200 m (Berg & Suchi 2001)
and Valle San Javier at 2950-3000 m (Schneider 2001). Clear floristic affinity is
found with the forests of La Montaña (‘teleférico’ or cable car) Sierra Nevada
National Park at 2550-2650 m (Kelly et al. 1994), where at least 16 (41%) of the
39 tree species reported in La Montaña are present in the Schefflero ferrugineae Cybianthetum laurifolii of Guaramacal, among them some diagnostic ones such as
Brunellia integrifolia, Hedyosmum crenatum, Myrsine coriacea, Schefflera
ferruginea and Weinmannia glabra.
The forest subcommunity of Freziera serrata of the Geissantho andini Miconietum jahnii at 2750 m in Guaramacal Sector displays some floristic affinity
with a succesional forest stand at 2600-2700 m in Valle San Javier, Mérida
described by Schneider (2001). This forest community in Guaramacal must have
been affected by disturbance due to the proximity to the road, man made clear cuts
and steep slopes, and evidenced by the presence of some species (such as Brunellia
integrifolia, Clethra fagifolia and Freziera serrata) which are also common (but
not restricted) in secondary Andean forests of Sierra Nevada de Mérida (Vareschi
1953).
Dwarf forest of Libanothamnus griffinii could be comparable in physiognomy and
some companion species (of Hypericum, Vaccinium, Weinmannia) with the dwarf
forest dominated by Libanothamnus neriifolius described for the Cordillera de
Mérida and the Cordillera de la Costa (Vareschi 1953, 1955; Monasterio 1980), as
well with those of Libanothamnus glossophyllus from the Sierra Nevada de Santa
Marta, Colombia (Cleef & Rangel 1984) and the Cordillera de Perijá (Rangel &
Arellano 2007). In Venezuela, Libanothamnus neriifolius dwarf forests are present
in the lower limit (2700-3200 m) of dry páramos of the Sierra Nevada de Mérida
and páramos of the Lara-Trujillo state border (Cendé, Tuñame, Los Nepes) on
sedimentary rocks of lutites and sandstones (Monasterio & Reyes 1980;
Monasterio 1980). However, the Libanothamnus neriifolius/glossophyllus dwarf
forests of the Sierra Nevada de Santa Marta, Colombia, are present on wind
62
The forest vegetation of Ramal de Guaramacal
_______________________________________________________
protected steep slopes between 3700-3900 m in comparable ecological conditions
as Polylepis forest in the Eastern Colombian Cordillera and the Sierra Nevada de
Mérida (Cleef & Rangel 1984). The Libanothamnus griffinii dwarf forests of
Guaramacal are present on similar bed rock and altitudinal range as
Libanothamnus neriifolius dwarf forests; under much wetter conditions however.
Libanothamnus griffinii, originally described as endemic from Guaramacal, has
also been reported for Lara state (Briceño & Morillo 2002). In total there are 11
species of Libanothamnus reported, most of them for Venezuela (Luteyn 1999);
few of them however, constitute dwarf forests.
Human influence and conservation
Ramal de Guaramacal is surrounded by at least 12 small villages and towns (Fig.
2.1). There has been a long history of agricultural activity in the region, now
occupying premontane and part of lower montane forest zone mainly for coffee
plantation, slash and burn cultivation and extensive cattle ranging, among other
land uses (Barbera 1999). However, the high ridges and steep slopes of
Guaramacal have kept most of the montane forest areas with minimum
disturbance. Only few paths crossing the range North-South existed in the past,
providing commercial connections between towns located South of Ramal de
Guaramacal and the city of Boconó and surroundings on the North side. These
paths were soon abandoned during the 1960’s after the road for the installation of
the antennas complex near the summit (3080 m) of Ramal de Guaramacal
continuing to the village of Guaramacal (c. 1300 m) on the South slope of the
massif, was constructed.
Ramal de Guaramacal has been, and continues to be, protected as a National Park
since 1988, keeping most human activities and impacts outside the park borders.
Fires are known to have occurred in the past, especially in páramo areas close to
the antennas, as well as in an area known as Cerro El Diablo on the West side of
the Ramal where some cattle were kept ranging in an extensive way. Timber
extraction is known to be selective, occurring at very low intensity and generally
takes place in close proximity to the park limits.
Currently, Ramal de Guaramacal and its montane ecosystems is one of the best
conserved national parks in Venezuela. We can only hope this situation will
continue into the future.
63
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
67
Flora, vegetation and ecology in the Venezuelan Andes
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).
68
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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 (9 o 14‟
1.02” N; 70o 11‟ 6.47” W) and Páramo El Pumar (9o 12‟ 45.6” N; 70o 12‟ 5.55”
W), 2.5 km Southwest of 'Las Antenas'. Ramal de Guaramacal is an outlier of the
Venezuelan Andes, located South from the town of Boconó, Trujillo state,
approximately 120 km Northeast of Mérida, in the centre of the Sierra Nevada de
Mérida (Fig. 3.1).
The climatic characteristics of high humidity with permanent fog favour the
development of great ground cover of Sphagnum spp. characteristic of the zonal
shrub páramo vegetation associations and border of forests. This condition is very
common all over the páramo areas of Ramal de Guaramacal and is not considered
here as an azonality. First climatic records from a Davis Pro 2 climate station
installed near the summit of Guaramacal (3100 m) by the first author since
December 2006 to December 2007 (monthly precipitation in mm and monthly
temperature in Celsius), registered a total amount of yearly rainfall of at least
2995.4 mm (some data were lost during some days in the most rainy months of
june and july 2007). Relative humidity is extraordinary high, with a mean humidity
of 96.88% throughout the year. The lowest mean relative humidity was observed
in the month of February with a value of 92.35%. Mean temperature is 8.6oC, the
lowest temperatures of 1.3oC are recorded in December and January and the
highest temperature of 18.6oC in March. Detailed data of the Davis Pro 2 climate
station are intended to be published in a forthcoming paper on the upper forest line
(Cuello et al. in prep.). For a more complete description of the study area the
reader is referred to Chapter 2 and Cuello (1999).
69
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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 highAndean 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 subpáramo vegetation extends in greater proportion on low inclined convex slopes,
and is represented by grasspáramo of the Puyo aristeguietae - Ruilopezietum lopezpalacii; bordered by or combined, with the vegetation of the new association
Cortaderio hapalotrichae - Hypericetum juniperinum. There, the grasses Cortaderia 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 singlestemmed 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
Flora, vegetation and ecology in the Venezuelan Andes
Table 3.1. Phytosociological table of zonal páramo vegetation of Ramal de Guaramacal, Andes, Venezuela.
Releve number
1
Releve (field number)
2
3
4
5
6
7
8
9 10 11 12 13 14 15 16 17 18 19
47a 47b 48b 32b 48a 39a 39b 11a 32a 3a 12a 12b
2
2
2
32
33
3
3
2
3
3
2
2
2
2
36 37
38
17a 17b 37b 7b
2
3
3
2
3
2
3
3
3
2
3
3
2
3
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
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
NW NW
N
SE
N
NW NW NW
SE
SE
S
S
W
30
30
30
17
20
25
30
1
2
2
2
1
1
1
1
1
95 106 45
38
60 >90 25
34
40
18
2
34 35
7a
0
18
2
30 31
3
19
2
29
3
30
2
28
0
45
2
26 27
3
45
3
24 25
T
Slope angle (degrees)
2
23
L
Slope exposition
3
22
19a 19b 2b 46a 46b 3b 45a 45b 29b 37a 29a 18b 34a 34b 43a 43b 18a 31a 31b 49b
A
(m)
3
20 21
2a
3
3
4
0
0
0
0
0
0
5
5
5
0
0
0
0
0
0
0
0
0
0
0
0
5
0
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
13
37
22
22
22
20
35
36
10
29
24
25
25
37
23
28
18
25
10
29
2
2
2
1
1
1
2
2
1
1
1
2
35 >110 53
56
17
30
25
12
24
12
21
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Soils depth (cm)
30
50
50
46
13 >55
10
10
41
4
33
67
60
56
75
62
20
35
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
FLa
aF
aF
a
aF
La
La
Fa
aF
a
A
FLA
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
Fa
F
La
FL
a
Fla
aL
FaL FaL aL
<1
<1
Fa
a
<1
<1
Fa
<1
<1
<1
aL
<1
a
aL
FaL
<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
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
5
5
15
30
10
5
10
5
10
20
20
40
39
20
30
20
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
5
5
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
RUILOPEZIO LOPEZ-PALACII
-
Order
<1
a
35
45
<1
a
% Cov. Small shrubs < 60 cm
35
<1
aF
% Cov. Shrubs & dwarf trees >60 cm
10
<1
a
40
% outcrops and/or bare soil
15
<1
FL
53 >80 45
1
35
Slope shape
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE
Alliance
1. Ruilopezio - Neurolepidetum glomeratae
Association
2. Disterigmo acuminatum - Arcytophylletum nitidum
2.1. pentacalietosum cachacoensis
Subasociacion
2.2. typicum
Variant
1. Ruilopezio paltonioides - Neurolepidetum glomeratae
. . 4 3 4
1 . 3 2 2
1 . 1 . .
. . 2 1 .
. . 2 . .
2. Disterigmo acuminatae - Arcytophylletum nitidum
Disterigma acuminatum
1 . . . .
Gaultheria hapalotricha
1 . . . .
Arcytophyllum nitidum
1 . . . .
Ageratina theifolia
. . . . .
Galium hypocarpium
. . . . .
Polypodium funckii
. . . . .
Eriosorus flexuosus
. . . . .
Hymenophyllum myriocarpum
. . . . .
2.1. pentacalietosum cachacoensis
Pentacalia cachacoensis
. . . . .
Vaccinium corymbodendron
. 4 3 . .
Melpomene moniliformis
. . . . .
Gaultheria anastomosans
. . . . .
Themistoclesia dependens
. . . . .
Hesperomeles sp.
. . . . .
2.2. typicum
Ugni myricoides
. . . . 1
Rubus acanthophyllos
. . . . .
Ilex guaramacalensis
. . . . .
Valeriana quirorana
. . . . .
Ruilopezia paltonioides
Disterigma alaternoides
Nertera granadensis
Pentacalia greenmaniana
Sphyrospermum buxifolium
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.
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE
Blechnum schomburgkii
1 3 3 3 4 . 1 2 2 2
Hypericum paramitanum
1 . 3 4 2 1 2 3 3 2
Neurolepis glomerata
5 5 5 . 5 5 5 3 . 2
Cybianthus marginatus
. . . . 1 . 1 . . 5
Hesperomeles obtusifolia
4 3 . 4 . . . . 1 .
Sphagnum meridense
4 3 2 . . . 2 3 . .
Libanothamnus griffinii
1 . . . . 2 2 . . .
Elaphoglossum cf. lingua
1 . . . . . . . . .
Puya sp.
. . . 2 1 . . . 4 2
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Paepalanthus pilosus
. . . .
4. Puyo aristeguietae - Ruilopezietum lopez-palacii
Puya aristeguietae
. . . .
Chusquea tessellata
. . . .
Castilleja fissifolia
. . . .
Festuca guaramacalana
. . . .
Monnina sp.
. . . .
Bejaria aestuans
. . . .
Rhynchospora lechleri
. . . .
Oreobolus venezuelensis
. . . .
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Miconia tinifolia
Muehlenbeckia tamnifolia
Epidendrum frutex
Myrsine dependens
Diplostephium obtusum
Rhynchospora sp.
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3.Hypericetum juniperinum
Hypericum juniperinum
Orthrosanthus acorifolius
Calamagrostis sp. A
74
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
2
2
3
2
3
3
3
3
3
3
3
3
2
3
3
2
2
2
2
2
2
2
3
3
2
2
2
3
0
0
0
9
0
0
0
0
0
8
9
9
0
8
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9
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3
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4
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6
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2
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5
0
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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
21
11 26 26
11
18
12
12
11
11
13 28 19
19
13
24 23
16
1
2
1
2
1
1
1
1
2
1
2
65 20 56 40 73
10
2
3
3
3
3
3
5
5
0
0
0
0
S
S
W NE SE SE E
S
S NW S
8
S W NE SE SE SW SW SE SE N
2
2
3
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
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15 15
11
21
2
3
1
1
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2
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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
1
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2
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1
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2
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3
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2
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2
15 115 20 32 120 55 106 80 26
2
3
2
1
2
1
2
3
2
1
2
2
48 60 28
5
5
2
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
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
aL aL
11 10 9
9
7
<1
aL aL aL aL La La aL a
10
5
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2
5 20 5
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5
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25
5
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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
10
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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
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5 20 10 15
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45
1
15 40 10
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10 10 30 10 20 20 50 15 40 45 20 50 20 20 30 3 30 10
5
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10
15 35 40 20 15 15
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L LA aL aL AF Fa Fa La aL aL aF aF La FLA FL aL aL LF LF aL aL aL LF Aa
5 20 3
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30 30 20 <1 10
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15 20 10
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5
0
- CHUSQUEETALIA ANGUSTIFOLIAE
HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
3. Cortaderio hapalotrichae - Hypericetum juniperinum
3.1. typicum
. . 1
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. 1 .
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4. Puyo aristeguietae - Ruilopezietum lopez-palacii
5. R. gollmeri - Ruilopezietum jabonensis
3.2. disterigmatosum acuminatum
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75
Flora, vegetation and ecology in the Venezuelan Andes
Releve number
1
Releve (field number)
2
3
4
5
6
7
8
9 10 11 12 13 14 15 16 17 18 19
47a 47b 48b 32b 48a 39a 39b 11a 32a 3a 12a 12b
2
2
2
32
33
3
3
2
3
3
2
2
2
2
36 37
38
17a 17b 37b 7b
2
3
3
2
3
2
3
3
3
2
3
3
2
3
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
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
NW NW
N
SE
N
NW NW NW
SE
SE
S
S
W
30
30
30
17
20
25
30
1
2
2
2
1
1
1
1
1
95 106 45
38
60 >90 25
34
40
18
2
34 35
7a
0
18
2
30 31
3
19
2
29
3
30
2
28
0
45
2
26 27
3
45
3
24 25
T
Slope angle (degrees)
2
23
L
Slope exposition
3
22
19a 19b 2b 46a 46b 3b 45a 45b 29b 37a 29a 18b 34a 34b 43a 43b 18a 31a 31b 49b
A
(m)
3
20 21
2a
3
3
4
0
0
0
0
0
0
5
5
5
0
0
0
0
0
0
0
0
0
0
0
0
5
0
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
13
37
22
22
22
20
35
36
10
29
24
25
25
37
23
28
18
25
10
29
2
2
2
1
1
1
2
2
1
1
1
2
35 >110 53
56
17
30
25
12
24
12
21
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
Soils depth (cm)
30
50
50
46
13 >55
10
10
41
4
33
67
60
56
75
62
20
35
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
FLa
aF
aF
a
aF
La
La
Fa
aF
a
A
FLA
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
Fa
F
La
FL
a
Fla
aL
FaL FaL aL
<1
<1
Fa
a
<1
<1
Fa
<1
<1
<1
aL
<1
a
aL
FaL
<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
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
5
5
15
30
10
5
10
5
10
20
20
40
39
20
30
20
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
5
5
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
RUILOPEZIO LOPEZ-PALACII
-
Order
<1
a
35
45
<1
a
% Cov. Small shrubs < 60 cm
35
<1
aF
% Cov. Shrubs & dwarf trees >60 cm
10
<1
a
40
% outcrops and/or bare soil
15
<1
FL
53 >80 45
1
35
Slope shape
HYPERICO PARAMITANUM - HESPEROMELETION OBTUSIFOLIAE
Alliance
1. Ruilopezio - Neurolepidetum glomeratae
Association
2. Disterigmo acuminatum - Arcytophylletum nitidum
2.1. pentacalietosum cachacoensis
Subasociacion
2.2. typicum
Variant
5. R. gollmeri - Ruilopezietum jabonensis
Ruilopezia jabonensis
. . . . . .
Rhynchospora gollmeri
. . . . . .
Isidrogalvia robustior
. . . . . .
Gentianella nevadensis
. . . . . .
HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
Xyris subulata var. acutifolia
. . . . . .
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4
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2
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3
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1
3
4
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2
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2
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2
3
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4
.
1
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1
2
2
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4
1
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2
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3
.
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2
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6
1
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2
4
1
4
.
2
3
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2
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3
4
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4
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3
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2
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4
4
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2
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2
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3
4
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1
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.
2
2
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4
2
2
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2
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1
.
1
1
2
3
.
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2
2
2
4
.
3
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2
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2
4
2
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2
.
3
1
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2
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5
3
1
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4
.
3
3
.
2
6
.
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5
2
4
2
1
.
3
2
3
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5
.
.
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3
3
3
4
4
1
.
1
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1
5
.
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4
4
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2
1
3
.
2
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.
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4
.
1
.
4
5
2
2
3
1
.
3
.
4
.
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.
.
.
4
5
3
3
2
1
.
1
.
.
2
6
.
.
.
0
1
0
0
0
0
1
0
0
0
0
1
0
0
0
0
1
0
0
1
0
1
0
0
1
0
1
0
0
1
0
1
0
0
1
0
1
0
0
1
0
1
0
1
0
0
1
0
1
0
0
1
0
1
0
0
1
0
1
1
0
1
0
1
1
0
1
1
0
0
0
1
1
0
1
0
1
1
0
1
0
1
1
0
1
0
1
1
0
1
0
1
1
0
1
0
1
1
1
0
0
1
1
1
0
0
1
1
1
0
0
1
1
1
0
0
1
1
1
0
0
1
1
1
1
0
1
1
1
1
0
1
1
1
1
0
1
1
1
1
Hypericum cardonae
. . . . . . . . 1 .
Carex bonplandii
. . . . . . . . . .
Ruilopezia viridis
. . . . . . . . . .
Calamagrostis planifolia
. . . . . . . . . .
RUILOPEZIO LOPEZ-PALACII - CHUSQUEETALIA ANGUSTIFOLIAE
Cortaderia hapalotricha
. . . 4 . 3 4 5 3 2
Chusquea angustifolia
3 4 1 4 5 . . 3 4 2
Lycopodium clavatum subsp. contiguum 1 . . 2 . 4 4 4 3 4
Ruilopezia lopez-palacii
Geranium stoloniferum
Pernettya prostrata
Rhynchospora guaramacalensis
Rhynchospora macrochaeta
Jamesonia imbricata
Chaetolepis lindeniana
Daucus montanus
Sphagnum sparsum
Hieracium avilae
Hymenophyllum trichomanoides
Hypericum sp.
.
.
1
2
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2
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1
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2
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1
1
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1
2
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1
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2
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2
1
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1
3
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2
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4
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1
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3
2
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5
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3
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1
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4
1
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6
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1
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2
2
1
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.
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3
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1
3
.
2
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.
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1
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1
3
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.
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.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
1
0
0
1
0
0
0
1
0
0
0
1
1
0
0
1
1
0
0
1
1
Cybianthus laurifolius? 3a(1)
Gaultheria erecta 34b(1)
Greigia sp. 44a(1)
76
Huperzia amentacea 3b(1)
Hymenophyllum sp. 34a(1)
Melpomene flabelliformis 17a(1)
Melpomene xiphopteroides 17b(1)
Polypodium sp. 34b(1)
Utricularia alpina 21b(1)
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
2
2
3
2
3
3
3
3
3
3
3
3
2
3
3
2
2
2
2
2
2
2
3
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
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
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
21
11 26 26
11
18
12
12
11
11
13 28 19
19
13
24 23
16
1
2
1
2
1
1
1
1
2
1
2
65 20 56 40 73
10
2
3
3
3
3
3
5
5
0
0
0
0
S
S
W NE SE SE E
S
S NW S
8
S W NE SE SE SW SW SE SE N
2
2
3
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
2
3
1
1
1
1
2
2
1
1
2
2
1
1
1
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
1
1
1
2
2
2
1
1
1
1
1
1
1
1
1
2
1
1
1
3
2
2
2
2
2
2
15 115 20 32 120 55 106 80 26
2
3
2
1
2
1
2
3
2
1
2
2
48 60 28
5
5
2
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
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
aL aL
11 10 9
9
7
<1
aL aL aL aL La La aL a
10
5
<1
2
5 20 5
<1
1
15 <1 <1
5
10
15
<1
<1
<1
<1
5
25
5
20
10
5
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
5
1
<1
<1
15
5
5
10
10
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
10
10
5
10
5
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
1
5
1
<1
5 20 10 15
1
5
1
20
1
5
5
<1
45
1
15 40 10
5
15
15
20
15
30
15
15
15
L
10 10 30 10 20 20 50 15 40 45 20 50 20 20 30 3 30 10
5
1
aL
10
15 35 40 20 15 15
<1
aL FaL aL aL
5
5
<1
a
5
5
2
a
2
1
1
aL
15
5 15 5
5
L FaL aL
5
5
1
a
<1
5
1
L LA aL aL AF Fa Fa La aL aL aF aF La FLA FL aL aL LF LF aL aL aL LF Aa
5 20 3
10
<1
a
<1
5
5
aF
30 30 20 <1 10
5
10
a
15 20 10
10
10
5
0
- CHUSQUEETALIA ANGUSTIFOLIAE
HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
3. Cortaderio hapalotrichae - Hypericetum juniperinum
3.1. typicum
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4. Puyo aristeguietae - Ruilopezietum lopez-palacii
5. R. gollmeri - Ruilopezietum jabonensis
3.2. disterigmatosum acuminatum
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2
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3
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. .
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. .
5 4 3 3 1 4
1 1 2 2 1 1
3 1 . . . .
. . . . 4 1
. . . . . .
1
1
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. 2 . . 1 .
. . . . 1 1
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. . . . . .
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3
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4
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3
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3
2
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1
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2
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3
1
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2
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5
. .
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3
4
2
3
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2
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3
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3
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2
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4
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77
Flora, vegetation and ecology in the Venezuelan Andes
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, Chusquea angustifolia, Cortaderia hapalotricha, Daucus montanus, Geranium stoloniferum, Hymenophyllum trichomanoides, Jamesonia imbricata, Lycopodium contiguum, 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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
Dwarf tree species of high Andean forest (or subalpine rain forest, SARF) are
common, such as: Cybianthus laurifolius, C. marginatus, Gaultheria erecta, Hesperomeles 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, Gaultheria anastomosans, Diplostephium obtusum, Pentacalia greenmaniana, Hypericum juniperinum x cardonae. Small ericaceous prostrate shrubs including:
Disterigma acuminatum, Pernettya prostrata, Gaultheria hapalotricha, Themistoclesia 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 guaramacalensis 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 flabellaformis, M. moniliformis and M. xiphopteroides. The trailings Rubus acanthophyllos 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 marginatus, 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 common 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
Flora, vegetation and ecology in the Venezuelan Andes
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 / Pajonalarbustal 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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
Flora, vegetation and ecology in the Venezuelan Andes
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: Hypericum paramitanum; Mp: Muehlenbeckia tamnifolia; Ng: Neurolepis glomerata; Ngr: Nertera granadensis; Pg: Pentacalia greenmanniana; Pp:
Pernettya prostrata; Rgu: Rhynchospora guaramacalensis; Rp: Ruilopezia
paltonioides; Sb: Sphyrospermum buxifolium; Vc: Vaccinium corymbodendron.
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
83
Flora, vegetation and ecology in the Venezuelan Andes
flabelliformis, M. xiphopteroides and Polypodium funckii, are also present, among
others.
Syntaxonomy: This is a highly diverse association represented by 28 line-intersect
surveys with 61 species of vascular plants.
Diagnostic species are Arcytophyllum nitidum, Ageratina theifolia, Disterigma
acuminatum and Gaultheria hapalotricha. Two subassociations are distinguished,
pentacalietosum cachacoensis and the typicum one.
Ecology and distribution: This subpáramo bamboo shrub is generally found
surrounding areas of dwarf forests (SARF), at edges of slopes or hill tops, and in
contact with communities of Ruilopezia paltonioides and Neurolepis glomerata. It
represents humid shrub páramo, transitional between forest and páramo.
Figure 3.3. Physiognomy of the vegetation of the association Disterigmo acuminatum Arcytophylletum nitidum subass. Typicum (L31a 2960 m). An: Arcytophyllum
nitidum; Bs: Blechnum schomburgkii; Cha: Chusquea angustifolia; Cm:
Cybianthus marginatus; Da: Disterigma acuminatum; Dm: Daucus montanus;
Gh: Gaultheria hapalotricha; Hp: Hypericum paramitanum; Ji: Jamesonia
imbricata; Lc: Lycopodium clavatum subsp. contiguum; Ng: Neurolepis
glomerata; Pp: Pernettya prostrata; Ra: Rubus acanthophyllos; Rl: Ruilopezia
lopez-palacii; Rm: Rhynchospora macrochaeta; Um: Ugni myricoides; V:
Valeriana quirorana.
Disterigmo acuminatum – Arcytophylletum nitidum
2.1. subassociation pentacalietosum cachacoensis Cuello & Cleef 2009
Typus: Rel. No. 17 (Cuello L46a). Table 3.1. Figure 3.4
Pentacalia cachacoensis subassociation / Subasociación de Pentacalia cachacoensis
Physiognomy: Dense shrubby vegetation in a matrix of tussock grasses of
Cortaderia hapalotricha and bamboos of Chusquea angustifolia and Neurolepis
glomerata; shrubs, dwarf trees (1-1.5 (3) m) and prostrate shrubs are present at
high density. There is a carpet of species of Sphagnum, together with other mosses,
84
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
as well as the presence of liverworts, such as Scapania portoricensis and species of
Plagiochila.
Composition and syntaxonomy: This subassociation is represented in 13 lineintersect 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 480 cm, with mixed textures predominantly sandy (sand-muddy to sand-silty or siltsand-loam), with pH 3.3-4.1 and dark colors in the superficial layers, varying in
colour until reddish and grayish with a high clay content at increased depth.
Disterigmo acuminatum – Arcytophylletum nitidum
2. 2. subassociation typicum Cuello & Cleef 2009
Typus: Rel. No. 31 (Cuello L31a). Table 3.1. Figure 3.3
Subassociation of Arcytophyllum nitidum / Subasociación de Arcytophyllum nitidum
Physiognomy: Shrubs and dwarfed trees dominate (up to 2 m, 20-40% cover);
with a presence of tall stem rosettes of up to 3.5 m.
Composition and syntaxonomy: The subassociation is represented in 15 lineintersect 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
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
Composition and syntaxonomy: This alliance is defined on the basis of 53 lineintersect 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 lopezpalacii; 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
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
89
Flora, vegetation and ecology in the Venezuelan Andes
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.
90
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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 lineintersect 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
93
Flora, vegetation and ecology in the Venezuelan Andes
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 lopezpalacii; 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
ASTERACEAE
ERICACEAE
POACEAE
CYPERACEAE
CLUSIACEAE
MELASTOMATACEAE
MYRSINACEAE
ROSACEAE
RUBIACEAE
GRAMMITIDACEAE
BROMELIACEAE
94
#
GENERA
6
7
4
3
1
3
2
2
3
2
2
# SPP
13
10
7 (+3 indets)
6
4
3
3
3
3
3 (+2 indets)
3
GENERA
Hypericum
Rhynchospora
Ruilopezia
Melpomene
Gaultheria
Hymenophyllum
Pentacalia
# SPP
4
4
4
3 (+2 indets)
3
3
3
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
Association
1. Ruilopezio paltonioides Neurolepidetum glomeratae
2. Disterigmo acuminatum Arcytophylletum nitidum
3.Cortaderio hapalotrichae Hypericetum juniperinum
4. Puyo aristeguietae Ruilopezietum lopez-palacii
5. R. gollmeri - Ruilopezietum
jabonensis
#
Families
# spp
20
41
27
61
22
50
22
45
13
22
Most diverse families
Ericaceae (7), Asteraceae (5), Clusiaceae,
Cyperaceae, Myrsinaceae and Poaceae (3)
Ericaceae (8), Asteraceae (6), Clusiaceae
(4)
Asteraceae (8), Cyperaceae, Ericaceae,
Poaceae (5), Clusiaceae (4)
Asteraceae, Ericaceae and Poaceae (6),
Cyperaceae (5)
Cyperaceae (4), Asteraceae and
Clusiaceae (3)
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 microphanerophytic 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.
95
Flora, vegetation and ecology in the Venezuelan Andes
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
phanerophyte
microphanerophyte
nanophanerophyte
phanerophytic lignified grass
rosullate phanerophyte
hemicryptophyte
caespitose hemicryptophyte
climbing hemicryptophyte
chamaephyte
frutescent chamephyte
reptant chamaephyte
Epiphyte
Total phanerophytes
Total hemicriptophytes
Total chamaephytes
Total spp
Total life forms
(1)
6
6
2
1
5
4
6
1
7
20
11
10
41
10
(2)
5
11
4
1
5
9
7
3
6
3
26
19
15
61
12
Number of species by vegetation type
(3)
(4)
(5)
3
3
1
8
9
1
4
2
2
1
2
1
7
5
2
5
5
3
12
12
8
1
5
5
2
9
4
2
1
23
21
7
17
17
11
10
7
4
50
45
22
10
9
9
Total spp
8
15
4
2
8
13
16
3
2
8
2
2
37
32
20
91
13
(b)
Number of species by vegetation type
Growth forms*
(1)
(2)
(3)
(4)
(5)
upright shrubs
9
12
10
9
2
tussocks
7
8
13
13
8
erect herbs
6
9
5
7
3
dwarf trees
6
9
6
5
2
prostrate herbs
2
8
4
2
2
ground rosettes
2
3
5
3
2
prostrate shrubs
5
5
3
2
1
cushions
1
1
1
stem rosettes
3
3
3
3
1
trailing herbs
1
3
epiphitic herbs
1
Total spp
41
61
50
45
22
* Adapted from Ramsay & Oxley, 1997 ad Hedberg & Hedberg, 1979
Total
spp
18
17
14
10
9
6
6
3
3
3
2
91
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
Table 3.5. Standard canonical coefficients and interset variables of CCA ordination axis for
páramo vegetation of Ramal de Guaramacal.
Variable
1 Alt
2 Slope angle
3 Soils depth
4 pH
5 Humus depth
Canonical Coefficients
Standardized
Original Units
Axis 1 Axis 2 Axis 3 Axis 1 Axis 2 Axis 3
0.135
0.389 -0.124
0.002
0.005 -0.002
-0.495
0.018 -0.059 -0.053
0.002 -0.006
-0.054 -0.012 -0.326 -0.002
0.000 -0.011
0.075 -0.100 -0.035
0.269 -0.361 -0.125
-0.054 -0.031
0.181 -0.006 -0.003
0.019
S.Dev
0.774E+02
0.940E+01
0.302E+02
0.278E+00
0.952E+01
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. Cortaderio 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
97
Flora, vegetation and ecology in the Venezuelan Andes
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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
Number of relevés
10
1. Ruilopezio paltonioides - Neurolepidetum
glomeratae
Ruilopezia paltonioides
IV
Disterigma alaternoides
II
Nertera granadensis
II
Pentacalia greenmaniana
I
Sphyrospermum buxifolium
I
Cybianthus laurifolius
I
2. Disterigmo acuminatum - Arcytophylletum
nitidum
Disterigma acuminatum
II
Gaultheria hapalotricha
I
Arcytophyllum nitidum
I
Ageratina theifolia
.
Galium hypocarpium
.
Polypodium funckii
.
Eriosorus flexuosus
.
Hymenophyllum myriocarpum
.
2.1. pentacalietosum cachacoensis
Pentacalia cachacoensis
.
Vaccinium corymbodendron
I
Melpomene moniliformis
.
Gaultheria anastomosans
.
Themistoclesia dependens
.
Hesperomeles sp.
.
Huperzia amentacea
.
2.2. typicum
Ugni myricoides
II
Rubus acanthophyllos
.
Ilex guaramacalensis
.
Valeriana quirorana
.
Gaultheria erecta
.
Hymenophyllum sp.
.
Melpomene flabeliformis
.
Melpomene xiphopteroides
.
Polypodium sp.
.
HYPERICO PARAMITANUM –
HESPEROMELETION OBTUSIFOLIAE
2
28
2.1
13
2.2
15
3
25
3.1
12
3.2
13
4
17
5
11
II
II
.
.
.
.
II
I
.
.
.
.
I
.
.
.
.
.
I
.
I
.
.
.
I
.
I
.
.
.
I
.
II
.
.
.
I
I
.
.
.
.
.
.
.
.
.
.
V
III
III
II
I
I
I
I
V
IV
II
II
I
I
I
I
V
III
IV
I
I
.
I
I
III
I
II
I
.
.
.
.
.
I
.
II
.
.
.
.
V
I
III
.
.
.
.
.
II
.
.
I
.
.
.
.
.
.
I
.
.
.
.
.
II
II
II
II
I
I
I
IV
III
III
III
I
I
I
I
I
.
I
I
.
.
.
II
I
I
.
I
.
.
II
I
II
.
I
.
.
II
.
I
.
.
.
.
I
.
I
.
.
.
.
.
.
.
.
.
.
II
II
I
I
I
I
I
I
I
I
.
.
.
.
.
.
.
.
III
II
I
I
I
I
I
I
I
I
.
.
.
.
.
.
.
.
.
.
I
.
.
I
.
.
.
.
.
I
.
.
I
.
.
.
.
.
.
.
.
.
.
.
.
.
.
I
.
.
.
.
.
99
Flora, vegetation and ecology in the Venezuelan Andes
Community group
1
2
2.1
2.2
3
3.1
Blechnum schomburgkii
.
.
V
IV
IV
IV
Hypericum paramitanum
I
II
V
V
V
IV
Neurolepis glomerata
II
II
I
.
IV
III
Cybianthus marginatus
II
II
I
I
I
IV
Hesperomeles obtusifolia
II
II
I
III IV
III
Libanothamnus griffinii
II
I
II
I
I
.
Elaphoglossum cf. lingua
I
II
I
II
.
.
Puya sp.
II
I
I
I
.
.
Miconia tinifolia
I
I
I
.
.
.
Muehlenbeckia tamnifolia
I
I
I
.
.
.
Epidendrum frutex
I
I
I
.
.
.
Myrsine dependens
.
I
II
I
.
.
Diplostephium obtusum
I
I
I
I
I
II
Rhynchospora sp.
I
I
I
.
I
I
3.Cortaderio hapalotrichae- Hypericetum
juniperinum
Hypericum juniperinum
.
II
II
I
V
V
Orthrosanthus acorifolius
.
I
I
I
II
III
Calamagrostis sp. A
.
.
.
.
I
II
Paepalanthus pilosus
.
.
.
.
I
I
Greigia sp.
.
.
.
.
I
.
4. Puyo aristeguietae - Ruilopezietum lopezpalacii
Puya aristeguietae
I
I
.
I
I
.
Chusquea tessellata
.
.
.
.
.
.
Castilleja fissifolia
.
.
.
.
.
.
Festuca guaramacalana
.
.
.
.
.
.
Monnina sp.
.
.
.
.
.
Bejaria aestuans
.
.
.
.
.
.
Rhynchospora lechleri
.
.
.
.
.
.
Oreobolus venezuelensis
.
.
.
.
.
.
Festuca sp.
.
.
.
.
.
.
Utricularia alpina
.
.
.
.
.
5. R. gollmeri - Ruilopezietum jabonensis
Ruilopezia jabonensis
.
.
.
.
I
I
Rhynchospora gollmeri
.
.
.
.
I
.
Isidrogalvia robustior
.
.
.
.
I
.
Gentianella nevadensis
.
.
.
.
.
.
Calamagrostis planifolia
.
.
.
.
.
.
HYPERICO CARDONAE - XYRIDION ACUTIFOLIAE
Xyris subulata var. acutifolia
.
I
.
I
III
V
Hypericum cardonae
I
I
I
.
III
V
Carex bonplandii
.
.
.
.
I
III
Ruilopezia viridis
.
.
.
.
I
I
RUILOPEZIO LOPEZ-PALACII - CHUSQUEETALIA ANGUSTIFOLIAE
Cortaderia hapalotricha
III
V
V
V
V
V
Chusquea angustifolia
IV
IV III
V
IV
IV
Lycopodium contiguum
IV
IV
IV
V
V
V
Ruilopezia lopez-palacii
III
IV III
IV
III
IV
Geranium stoloniferum
.
III III
III
IV
IV
Pernettya prostrata
V
V
IV
V
V
V
Rhynchospora guaramacalensis
II
II
I
.
III
III
Rhynchospora macrochaeta
I
II
I
III
III
IV
Jamesonia imbricata
II
I
II
I
III
III
Chaetolepis lindeniana
II
IV
IV
III
III
III
Daucus montanus
I
II
II
II
III
III
Hieracium avilae
.
I
I
I
I
I
Hymenophyllum trichomanoides
.
I
I
I
I
I
Hypericum sp.
I
I
I
.
I
I
100
3.2
II
I
I
I
IV
II
.
.
.
.
.
.
I
I
4
II
II
I
I
.
I
I
.
.
.
I
I
.
I
5
.
I
I
.
.
.
.
.
.
.
.
.
.
.
V
I
.
I
I
II
.
.
.
.
II
.
.
.
.
I
.
.
.
.
.
.
.
.
.
IV
II
II
I
I
I
I
I
I
I
.
.
.
.
.
.
.
II
.
.
I
.
I
.
.
.
.
I
.
.
V
IV
I
I
I
II
II
I
I
IV
I
.
I
V
III
.
.
V
IV
V
III
V
V
II
II
III
II
II
II
I
.
V
IV
V
V
II
V
III
II
IV
I
I
II
.
.
V
V
V
I
IV
II
.
V
II
.
.
I
.
.
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
Cord. Oriental
Association
1
2
3
4
5
Colombia
Ageratina
.
II
I
I
.
.
Aragoa
.
.
.
.
.
I
Arcytophyllum
I
II
.
I
III
III
Azorella
.
.
.
.
.
I
Bartsia
.
.
.
.
.
V
Bejaria
.
.
.
I
.
.
Blechnum
.
II
.
I
V
IV
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
I
.
.
IV
III
Chusquea
IV
IV
IV
V
V
V
Cortaderia
II
III
V
V
V
V
Cybianthus
II
II
I
I
.
.
Cyperus
.
.
.
.
.
I
Daucus
I
II
I
.
.
III
Diplostephium
I
I
I
.
.
I
Disterigma
II
.
II
III
V
III
Elaphoglossum
I
II
.
I
.
.
Epidendrum
I
I
.
I
.
.
Eriosorus
.
I
.
.
.
.
Eryngium
.
.
.
.
.
I
Espeletia
.
.
.
.
.
IV
Festuca
.
.
.
I
.
II
Galium
.
I
.
.
.
.
Gaultheria
I
I
.I
.
III
Gentiana
.
.
.
.
.
II
Gentianella
.
.
.
.
I
IV
Geranium
.
II
I
III
IV
IV
Greigia
.
.
I
.
.
.
Halenia
.
.
.
.
.
I
Hesperomeles
II
.
.
.
III
III
Hieracium
.
I
I
II
I
I
Huperzia
.
I
.
.
.
I
Hydrocotyle
.
.
.
.
.
I
Hymenophyllum
.
I
I
.
.
.
Hypericum
I
V
V
V
III
V
Hypochaeris
.
.
.
.
.
III
Ilex
.
I
.
.
.
.
Isidrogalvia
.
.
I
I
I
.
101
Flora, vegetation and ecology in the Venezuelan Andes
Number of relevés
10
28
25
17
11
Association
1
2
3
4
5
III
.
II
IV
.
I
I
.
I
.
II
IV
.
.
.
.
.
I
V
.
.
II
.
V
.
V
.
.
III
I
.
II
.
I
.
.
.
II
.
I
IV
.
II
I
.
I
I
.
I
.
.
.
I
.
II
V
.
I
I
.
IV
II
IV
.
.
III
.
I
II
.
II
I
.
I
II
.
I
V
.
I
.
.
.
.
I
I
.
.
.
II
I
.
V
.
.
I
.
V
.
V
.
.
I
.
.
I
.
II
I
.
III
IV
.
I
V
.
.
.
I
.
I
.
I
.
I
.
.
.
.
V
.
.
IV
.
V
.
V
.
.
I
.
.
I
.
I
I
.
IV
II
.
.
V
.
.
.
.
.
.
.
I
.
II
.
.
.
.
II
.
.
.
.
V
.
V
.
.
Jamesonia
Laestadia
Libanothamnus
Lycopodium
Lysipomia
Melpomene
Miconia
Monnina
Muehlenbeckia
Myrsine
Nertera
Neurolepis
Niphogeton.
Oreobolus
Oritrophium
Orthrosanthus
Paepalanthus
Pentacalia
Pernettya
Plantago
Polypodium
Puya
Rhacocarpus
Rhynchospora
Rubus
Ruilopezia
Scirpus
Sisyrinchium
Sphagnum
Sphyrospermum
Themistoclesia
Ugni
Utricularia
Vaccinium
Valeriana
Xenphyllum
Xyris
.
.
.
.
.
.
.
V
25
Cord. Oriental
Colombia
I
I
.
II
I
.
.
.
.
.
II
.
I
III
III
.
I
III
I
I
.
I
II
I
.
.
I
I
III
.
.
.
I
I
I
.
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 km 2. 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
The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
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Flora, vegetation and ecology in the Venezuelan Andes
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 lopezpalazii. 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.
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The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
Figure 3.10. Gradient SARF – Zonal páramo, 3000-3050 m, North of „Las Antenas‟, Ramal
de Guaramacal, Andes, Venezuela 1. Ruilopezio paltonioides - Neurolepidetum 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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Flora, vegetation and ecology in the Venezuelan Andes
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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The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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
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Flora, vegetation and ecology in the Venezuelan Andes
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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The páramo vegetation of Ramal de Guaramacal: 1. Zonal communities
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.
109
Variety of Sphagnum species found in azonal páramo vegetation of Ramal de Guaramacal: (a)
Sphagnum recurvum covering the wet shore of Laguna El Pumar; (b, c) S. recurvum (detail); (d) S.
recurvum covering the north-west side of Laguna EL Pumar, S. cuspidatum submerged in the water; (e)
S. cuspidatum (detail); (f) S. meridense forming the ground cover in shrubparamo ; (g) S. recurvum and
S. magellanicum (darker); (h) S. sparsum with Campylopus cuspidatus.
Chapter 4
The páramo vegetation of Ramal de Guaramacal, Trujillo State,
Venezuela. 2. Azonal vegetation
Nidia L. Cuello A. and Antoine M. Cleef
PHYTOCOENOLOGIA, 39 (4), 389–409. 2009
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
4.1 INTRODUCTION
The azonal páramo vegetation in Guaramacal was studied between 2870 and 3050
m, mainly in two peat bog areas of the Sector Páramo El Pumar (Laguna El Pumar
and Laguna Seca). Azonal patches are also present in the small valleys or
depressions where water collects in Páramo de Guaramacal, near the „Las Antenas‟
area.
Peat bogs associated with glacial and seasonal lakes or fluvio-glacial valleys are
common features in Andean and Costa Rican páramos. A great variety of azonal
bog vegetation communities associated with glacial lakes and terrain depressions
have been described and named from the Colombian (Cleef 1981, Sánchez &
Rangel 1990, Cleef et al. 2005, 2008, Rangel et al. 2006 among others) and Costa
Rican páramos (Brak et al. 2005). A low number of diverse aquatic and peat bog
vegetation communities have been reported for Venezuelan páramos (Vareschi
1955, 1980; Monasterio 1980a, Bono 1996, Berg 1998, Berg & Suchi 2001) with
only a few of them treated in a syntaxonomic context of the upper páramo
vegetation of Sierra Nevada de Mérida (Berg, 1998). Vareschi (1955) described an
association („Sphagnetum maghellanici‟) from Naiguatá between 2500-2700 m.
As mentioned before Sphagnum bogs are present in the equatorial Andes and the
Central American Talamancas; up to date, they are not classified at the level of
order and class. Other bogs and mires which have been described for the northern
Andes concern vascular cushion bogs (Plantagini rigidae-Distichietea muscoides
Rivas Martínez & Tovar 1982) and cyperaceous reedswamps (Galio canescentisGratiolion bogotensis Cleef 1981), grass mires (Calamagrostion ligulatae Cleef
1981), both belonging to the order Marchantio plicatae-Epilobietalia denticulatae
Cleef 1981. Other azonal aquatic vegetation includes flush communities
(Xenophyllion crassae-Wernerion pygmaeae Cleef 1981), the vegetatation of
glacial lake bottoms (Ditricho submersi-Isoëtion karstenii Cleef 1981) and ponds
(Limoselletea australis Cleef 1981). Sphagnum bogs have not been classified in the
absence of comprehensive synthetic presence tables thus far. They are found in
valleys in the uppermost forests and the lower páramo, where conditions allow for
Sphagnum growth. Eutrophic to mesotrophic conditions allow for mires, which are
characterized by active mineroptrophic input from surrounding zonal vegetation on
slopes. The highest bogs in páramos are the vascular cushion bogs consisting of
Plantago rigida. Distichia muscoides, Oreobolus cleefii and the flat cushions of
Xyris subulata var. breviscapa (Bosman et al. 1993, Cleef 1981, Cleef et al. 2005,
2008, Moscol Olivera & Cleef 2009, Ramsay 1992, Coombes & Ramsay 2001,
Rangel Ch. & Ariza-N. 2000, Salamanca et al. 2003). In the Colombian Eastern
Cordillera páramos vascular cushion bogs replace altitudinally the Sphagnum bogs
at 3800-3900 m, as probably also in the Sierra Nevada de Mérida.
For Chusquea-Sphagnum bogs reference can be made to Cleef (1981), Sánchez &
Rangel Ch. (1990), Rangel Ch. & Franco (1985) and Cleef et al. (2006). They also
have been observed in the páramos of Costa Rica (Chaverri & Cleef 1992, Brak et
al. 2005).
The main goal of the present study is to identify, define and characterize the azonal
vegetation of two páramo areas of Ramal de Guaramacal (Páramo de Guaramacal
and Páramo El Pumar) aiming at the establishment of a syntaxonomic scheme
113
Flora, vegetation and ecology in the Venezuelan Andes
based on the analysis of the physiognomy, floristic composition and ecological
relations of the different vegetation communities.
This work was carried out within the framework of a larger project aimed at the
study of the diversity of the flora and vegetation of the Guaramacal Nacional Park
(Cuello, 1999; 2000, 2002; 2004). The classification of the vegetation of forests
and zonal páramo of Guaramacal range are described separately (Cuello & Cleef,
2009a; b; Chapters 2 and 3).
4.2 STUDY AREA
The azonal páramo communities have been studied in two páramo areas at the top
of Ramal de Guaramacal, between ca. 2900 and 3100 m in the area surrounding
the „Las Antenas‟ site in the Páramo de Guaramacal, and along the road crossing
the Ramal and the „Lagunas del Pumar‟ zone in Páramo El Pumar at 2.5 km to the
Southwest from „Las Antenas‟ (Fig 4.1).
Figure 4.1. Geographic location of Páramo de Guaramacal in the Venezuelan Andes.
The area studied in Páramo de Guaramacal concerns a small pond located at 9 o 14‟
1.02” N; 70o 11‟ 6.47” W with surrounding bamboo páramo vegetation present at
the bottom of a small valley where water collects at ~3080-3100 m (Photo 4.1).
This pond seems to be a remnant of small lake that existed in the past, according to
observations of 1960‟s aerial photographs from Páramo de Guaramacal.
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The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
Photo 4.1. Azonal vegetation associated to a pond at 3080 m in Páramo de Guaramacal (9o
14‟ 1.02” N; 70o 11‟ 6.47” W), Andes, Venezuela.
In the Páramo El Pumar, the associated vegetation of two contiguous glacial lakes
was studied. The Laguna El Pumar, ~2880 m located at 9 o 12‟ 52.36” N; 70o 12‟
8.04” W, which is covered mostly by water and bordered by Sphagnum bogs
(Photo 4.2). The second lake at around 2890 m is located 110 m to the South of
Laguna El Pumar. This is an evaporated lake called „Laguna Seca‟ at 9 o 12‟ 47.7”
N; 70o 12‟ 7.27” W, which is totally covered by Sphagnum bog and surrounded by
bamboo vegetation or „chuscales‟.
Ramal de Guaramacal is an outlier of the Venezuelan Andes, located South of the
town of Boconó in Trujillo state, about 120 km Northeast of the city of Mérida in
the centre of the Sierra Nevada de Mérida. For a more complete description of the
study area the reader is referred to Cuello (1999) and Chapter 2.
4.3 METHODS
Field Sampling
Azonal vegetation was studied by means of observations, plant collections and
surveys of small plots of between 0.25 to 6 m2 according to minimum area and the
extent of the homogenous and representative patches under consideration
(Westhoff & Van der Maarel 1973, Cleef 1981). In each plot, and per vegetation
layer, the percentage of periphery cover for each plant species was estimated. A
total of 71 relevés (approx. 100 m2) were surveyed. Eight line intercept transects of
5 m in length (as used in Cuello & Cleef, 2009b, Chapter 3) were surveyed in
bamboo páramo vegetation and included in the vegetation analysis. Azonal
vegetation associated with slopes trail borders and areas of disturbance were not
115
Flora, vegetation and ecology in the Venezuelan Andes
included in the phytosociological analysis which was based solely on observations
and collections of species composition. Field surveys were carried out only during
the dry season. In the „Las Antenas‟ area of Páramo de Guaramacal, sampling was
conducted by both authors in February 15, 2006, whilst in the remote area of
Páramo El Pumar sampling was completed by the first author and coworkers
during two different visits: one on March 1 st 2006, the other on February 18-19,
2007.
Botanical vouchers of all recorded species were collected. Photographs were taken
where possible. The collected botanical material was processed, identified and
deposited at Herbario Universitario PORT of the Universidad Nacional
Experimental de los Llanos “Ezequiel Zamora” (UNELLEZ) in Guanare,
Venezuela. For vascular plants the nomenclature follows Dorr et al. (2000),
complemented by Luteyn (1999) for other plant groups. Drs J. Hickey (Isoëtes
karstenii), G. Davidse (MO) and S. Laegaard (AAU) were helpful with the
identification of some selected grasses. Duplicates of vascular plants are deposited
in MER, VEN and US. Duplicates of the bryophytes were sent to Dr. D. Griffin III
(FLAS) with lichens sent to Dr. H.J.M. Sipman (B) for identification. Additional
duplicates of bryophytes and lichens were also deposited in L and MERC. The
record of bryophyte and lichen species has not been completed by the first author.
Only the most prominent and conspicuous species were collected.
Processing and data analysis
The data from each survey were stored and processed in Microsoft Excel. For each
species at each plot of azonal vegetation we used the percentage of cover estimated
in the field.
The data matrix of percentage cover for 53 species and 79 surveys of azonal
vegetation was processed with TWINSPAN (Hill 1979) using the PC-Ord 4
program (McCune & Mefford 1999). Data were then interpreted in terms of
community delimitation, the syntaxonomical vegetation classification based on
cover and floristic affinities following the Zürich-Montpellier approach (BraunBlanquet 1979). The names of the syntaxa are according to the International Code
of Phytosociological Nomenclature (Weber et al. 2000). The original cover values
of the relevés taken in percentages are available from the first author by request.
The diverse subunits, recognized in a progressive way by the TWINSPAN
procedure, were hierarchized in associations and higher (alliances, order) and
lower (subassociations) syntaxa and variants. The associations represent the basic
unit of description of the vegetation and are defined on the basis of floristic
composition (diagnostic, character species), particular appearance (growth form)
and habitat conditions. Two or more associations that share diagnostic species are
combined into an alliance. Two or more alliances combine to form an order.
Associations with some marked differences, or only variations in their floristic
composition, are subdivided into subassociations and eventually variants,
respectively.
In order to elucidate floristic relationship with Sphagnum dominated páramo
communities elsewhere in Colombia and Venezuela a Bray-Curtis similarity
cluster analysis (Bray & Curtis 1957) has been used. The „Spagnetum
116
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
maghellanici‟ Vareschi 1955 has been left out; only one species, Sphagnum
magellanicum, in common with Guaramacal bogs (Table 4.1).
Photo 2. Laguna El Pumar, 2880 m, Ramal de Guaramacal, Andes, Venezuela.
4.4 RESULTS
Flora diversity
A total of 53 morphospecies, belonging to 30 species of vascular plants, 20 species
of cryptogams and 3 undetermined species of algae have been recorded from a
total of 79 plots of the azonal vegetation in Páramo de Guaramacal and Páramo El
Pumar, Ramal de Guaramacal, Venezuela. The vascular plants include: 13 species,
belonging to 11 genera and 8 families of dicots; 15 species, 12 genera and 5
families of monocots and 2 species, 2 genera and 2 families of ferns. The identified
cryptogams include 14 species, 5 genera and 5 families of mosses, 2 species, 2
genera and 2 families of liverworts and 4 species, 3 genera and 3 familes of
lichens.
Azonal páramo plant communities
The interpretation of the TWINSPAN table, based on floristic composition,
affinities and species cover, allowed the recognition of six vegetation communities
at association level grouped into two alliances and one order (Table 4.1 and 4.2).
The azonal vegetation communities recognized in Ramal de Guaramacal are
summarized as follows:
117
118
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CH3 Hypericum juniperinum x cardonae
CH3 Hypericum cardonae
NP1 Hypericum juniperinum
Sphagnum sp.(orange)
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CARICETALIA BONPLANDII
1 0.8 0.5 0.3 0.3
SPHAGNO RECURVI - PAEPALANTHION PILOSI
GERANIO STOLONIFERUM
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2 2 1 3 . 3 1 5 5 5 . 1 5 4 3 5 5 2 1 . .
1.2. typicum
1 1 1 1
1 3 1 2 . 1 2 5 5 3 2 1 5 5 4 5 4 1 1 2 2 4 1
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Campylopus albidovirens
CH2 Arenaria venezuelana
CH2 Lachemila verticillata
Breutelia rhythidoides
DS 1.1. ortachnetosum erectifoliae
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Breutelia squarrosa
Polytrichum commune
Cladonia dydima
Cladonia andesita
H2 Rhynchospora gollmeri
Cladia aggregata
DS 1.2. typicum
Polytrichum juniperinum
H3 Sisyrhinchium sp.
1.1. ortachnetosum erectifoliae
1.Paepalantho pilosi - Agrostietum basalis
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Sphagnum sparsum
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NP1 Diplostephium obtusum
H3 Agrostis sp. B
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CARICI - CHUSQUEION
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3. Sphagno sparsi - Caricetum bonplandii
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LS L L SF S L LS L L L L L L L L L L L L L L LS LS SF8 SF8 PL S L L PF PF PF PF S SL LP LP7 L LS L L LS LS L PF SF8 SF SF SF SF P PL PL PF PL LP1 LP2 LP LP LP5 G L2 L2 L2 L2 L3 L3 L3 L3
S1 18 S2 S2 29 4 S5 43 5* 88 91 92 94 L 7 8 42 2 *
7
1 L6 S3 S3 36 40 44 45 F9 9* 6
11 S1 S1 96 L1 S7 10 S S8 S9 S S S S S2 S2 S S S S6 S1 19 3 6
3 4
C 6b 6a 7a 7b 6a 6b 3a 3b
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DS 1.Paepalantho pilosi - Agrostietum basalis
Variant
Subassociation
Association
Alliance
Area (m2)
Order
Releve number
Releve (field number)
Flora, vegetation and ecology in the Venezuelan Andes
1
2
3
4
5
Sphagnum recurvum
1
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Hemicryptophyte / caespitose ( > 30 cm)
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(10 - 30 cm)
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(3 - 10 cm)
Chamephyte / frutescent (10 - 30/50 cm)
"
(3 - 10 cm)
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4. Carici - Chusqueetum
CARICI - CHUSQUEION
4 0.35 0.5 0.6 0.5 0.35 6
3.2. Pernettya prostrata
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3.1.Diplostephium obtusum
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NP1 Nanophanerophyte (30/50 - 100 cm)
PLG Phanerophytic lignified grass (30 - 100 cm)
RP1 Rosullate phanerophyte (10 - 30 cm)
RP2
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2. Sphagno recurvi-Caricetum bonplandii
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CARICETALIA BONPLANDII
1 0.8 0.5 0.3 0.3
SPHAGNO RECURVI - PAEPALANTHION PILOSI
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LP, PL, PF Laguna El Pumar, 2880 m. Páramo El Pumar
LS, SL, SF, L36 Laguna Seca, 2890 m. Páramo El Pumar
GCC, L26, L27, L33 Páramo de Guaramacal, 3030 m.
* Representative relevé
DS Diagnostic Species
4
GERANIO STOLONIFERUM
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Rhacocarpus purpurascens
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H3 Agrostis perennans
DS GERANIO STOLONIFERUM - CARICETALIA BONPLANDII
H3 Carex bonplandii
. 1 . . . . 1 1 .
4 5 5 5 4 5 . . .
CH2 Geranium stoloniferum
.
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5 5 5 5 5 5 5 5 5 5 5 5 4 4 4 2 5 2 4 5 1
1 . 4 . 4 3
1
. 1 1 2 1 2 . . . . . 2 5 5 5 5 5
1.1. ortachnetosum erectifoliae
1.Paepalantho pilosi - Agrostietum basalis
1
H3 Calamagrostis sp. A
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DS 4. Carici bonplandii - Chusqueetum angustifoliae
PLG Chusquea angustifolia
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Sphagnum sancto-josephense
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H3 Xyris subulata
Campylopus richardii
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RP1 Ruilopezia jabonensis
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H4 Cortaderia hapalotricha
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H3 Gentianella nevadensis
Peltigera neopolydactyla
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H2 Oreobolus venezuelensis
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H3 Hieracium avilae
Jamesoniella rubricaulis
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CH2 Nertera granadensis
H2 Ophioglossum crotalophorioides
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Plagiochila sp.
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CH2 Paepalanthus pilosus
6
LS L L SF S L LS L L L L L L L L L L L L L L LS LS SF8 SF8 PL S L L PF PF PF PF S SL LP LP7 L LS L L LS LS L PF SF8 SF SF SF SF P PL PL PF PL LP1 LP2 LP LP LP5 G L2 L2 L2 L2 L3 L3 L3 L3
S1 18 S2 S2 29 4 S5 43 5* 88 91 92 94 L 7 8 42 2 *
7
1 L6 S3 S3 36 40 44 45 F9 9* 6
11 S1 S1 96 L1 S7 10 S S8 S9 S S S S S2 S2 S S S S6 S1 19 3 6
3 4
C 6b 6a 7a 7b 6a 6b 3a 3b
3
1
7
0
0 1
12 15 25 22 3 4 26 27 28
1
0
*
3 4
C
*
6*
3
DS SPHAGNO RECURVI - PAEPALANTHION PILOSI
Variant
Subassociation
Association
Alliance
Area (m2)
Order
Releve number
Releve (field number)
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
119
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
GERANIO STOLONIFERUM – CARICETALIA BONPLANDII Cuello & Cleef
2009
I. Sphagno recurvi – Paepalanthion pilosi Cuello & Cleef 2009
1. Paepalantho pilosi – Agrostietum basalis Cuello & Cleef 2009
1.1 subassociatio ortachnetosum erectifoliae Cuello & Cleef 2009
1.2. subassociation typicum Cuello & Cleef 2009
2. Sphagno recurvi – Caricetum bonplandii Cuello & Cleef 2009
3. Sphagno sparsi – Caricetum bonplandii Cuello & Cleef 2009
3.1. Variant with Diplostephium obtusum
3.2. Variant with Pernettya prostrata
II. CARICI BONPLANDII – CHUSQUEION ANGUSTIFOLIA Cuello & Cleef 2009
4. Carici bonplandii – Chusqueetum angustifoliae Cuello & Cleef 2009
III. DISTRICHO SUBMERSI – ISOETION Cleef 1981
5. Community of Sphagnum cuspidatum
6. Isoëtetum karstenii Cleef 1981
GERANIO STOLONIFERUM – CARICETALIA BONPLANDII Cuello & Cleef
2009
Representative alliance: Sphagno recurvi–Paepalanthion pilosi
Azonal páramo vegetation of the Geranium stoloniferum - Carex bonplandii order /
Vegetación de páramo azonal del orden de Geranium stoloniferum y Carex bonplandii
Physiognomy: The order Geranio stoloniferum - Caricetalia bonplandii concerns
the azonal páramo peat bog vegetation along the shore of lakes, and is represented
by Sphagnum peat bogs predominantly covered by Carex bonplandii together with
open bunchgrass patches dominated by Agrostis basalis and Ortachne erectifolia.
The order also includes the bamboo páramo „chuscales‟ of Sphagnum-Chusquea
angustifolia growing close to the lake shores or at the bottom of small valleys.
Composition and syntaxonomy: The order is defined on the basis of 69 relevés
with 28 vascular species and 19 cryptogams. The most species diverse vascular
families are Poaceae, Cyperaceae, Asteraceae and Clusiaceae. Sphagnaceae is the
most speciose and dominant bryophyte family in the ground layer.
This order is composed of two alliances: Sphagno recurvi - Paepalanthion pilosi
and Carici bonplandii - Chusqueion angustifoliae. Diagnostic species are Carex
bonplandii and Geranium stoloniferum. Sphagnum recurvum is another important
species and is present in both alliances.
120
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
Ecology and distribution: Azonal páramo vegetation of the Geranium stoloniferum - Carex bonplandii order is found in Páramo El Pumar (2870-2990 m) along
and close to the shores of both lakes Laguna El Pumar and Laguna Seca as well as
in a small wet valley South of „Las Antenas‟ in Páramo de Guaramacal.
SPHAGNO RECURVI – PAEPALANTHION PILOSI Cuello & Cleef 2009
Typus: Paepalantho pilosi – Agrostietum basalis
Azonal páramo vegetation of the Agrostis basalis - Paepalanthus pilosus alliance /
Vegetación de páramo azonal de la alianza de Agrostis basalis y Paepalanthus pilosus
Physiognomy and composition: Sphagnum bogs characterized by a ground layer
formed by dense cushions of Sphagnum spp. and Paepalanthus pilosus with the
occasional presence of a layer of variable cover of small tussock grasses, 10-25 cm
tall, which may be composed by Agrostis basalis, Carex bonplandii,
Rhynchospora golmerii, Xyris subulata var. acutifolia, and a layer of grasses 30-50
cm tall, formed of Agrostis perennans, Calamagrostis bogotensis, Cortaderia
hapalotricha, Ortachne erectifolia, and a species of Sisyrhinchium. A layer of little
shrubs may also be present, composed of: Diplostephium obtusum, Hesperomeles
obtusifolia, Hypericum cardonae, H. juniperinum, H. juniperinum x cardonae and
Pernettya prostrata.
Syntaxonomy: Sixty relevés are recognized as belonging to this alliance,
comprising a total of 21 vascular species and 13 species of cryptogams accounting
for the total species richness. Diagnostic of the alliance are: Agrostis basalis,
Sphagnum recurvum and Paepalanthus pilosus. This alliance contains three
associations: Paepalantho pilosi–Agrostietum basalis, Sphagno recurvi–Caricetum
bonplandii and Sphagno sparsi–Caricetum bonplandii.
Ecology and distribution: Vegetation belonging to this alliance may be found all
over the evaporated lake “Laguna Seca” and on the humid shore of Laguna El
Pumar in Páramo El Pumar (~2870-2890 m), as well as in wet areas around a pond
to the South of „Las Antenas‟ in Páramo de Guaramacal at ~3080 m.
1. Paepalantho pilosi – Agrostietum basalis Cuello & Cleef 2009
Typus: Rel. No. 17 (Cuello LS26). Table 4.1, Fig. 4.2. Photo 4.3 (center to right)
Peat bog with cushions of Paepalanthus pilosus and Agrostis basalis groundrosette bunchgrass vegetation / Vegetación de turbera con cojines de Paepalanthus pilosus y
pajonal de Agrostis basalis
Physiognomy and composition: The association is made up of small patches of
bunchgrass vegetation on top of a former peat bog. There is a ground layer formed
by dense cushions of Paepalanthus pilosus, Arenaria venezuelana and Lachemilla
verticillata. Over and among the cushions of Paepalanthus pilosus there is a
diversity of bryophytes (and lichens), with the moss Campylopus albidovirens
forming a dense and cespitose mat that, together with the other ground layer
species constitutes a substrate for the establishment of the bunchgrasses Agrostis
121
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
basalis and Ortachne erectifolia. In the ground layer, the most common
cryptogamic species are: Breutelia rhythidoides, B. squarrosa, Polytrichum
commune, P. juniperinum, Sphagnum magellanicum, S. recurvum and the lichens
Cladonia andesita, C. dydima and Cladia aggregata.
Syntaxonomy: The association is defined on the basis of 22 relevés with 10
vascular species and 11 species of cryptogams. The diagnostic species are:
Agrostis basalis, Arenaria venezuelana, Lachemilla verticillata, Paepalanthus
pilosus and the mosses Campylopus albidovirens, Breutelia squarrosa, B.
rhythidoides and Polytrichum commune.
A subassociation ortachnetosum erectifoliae and a subassociation typicum have
been recognized for this association.
Ecology and distribution: The vegetation of the association Paepalantho pilosiAgrostietum basalis has to date been found established solely to the North and the
South side of an area of central drainage (small channel with water) to the western
border of the peat bog (dry lake) of the Páramo El Pumar ~2870 m. This area
seems to be a site where wild fauna (probably the „Puma‟ Puma concolor and
other mammals), that go to the site for water, are concentrated. The frequent
animal footsteps seem to have caused a fragmentation and decomposition of the
Sphagnum layer, thereby favoring the establishment of other plant species.
1. Paepalantho pilosi – Agrostietum basalis
1.1 subassociation ortachnetosum erectifoliae Cuello & Cleef 2009
Typus: Rel. No. 8 (Cuello LS16). Table 4.1, Fig. 4.2, Photo 4.3
Peat bog with cushions and Ortachne erectifolia bunchgrass vegetation / Turbera con
vegetación de cojines con pajonal de Ortachne erectifolia
Physiognomy and composition: The vegetation is made up of a grass layer
dominated by tussocks of Ortachne erectifolia (height 30-45 cm and 30-70%
cover), small tussocks of Agrostis basalis [height 15-25 cm and 20-30% of cover]
and other herbs (1-5% cover) such as Carex bonplandii and Rhynchospora
gollmeri. The ground layer is composed of dense cushions of Paepalanthus pilosus
(10-80% cover) and Lachemilla verticillata (3-30% cover) with a mat of
Campylopus albidovirens (2-20 (40) % cover) growing in between. Other species
with variable densities and cover are Arenaria venezuelana and Geranium
stoloniferum, the bryophytes Breutelia rhythidoides, B. squarrosa, Campylopus
cuspidatus var. dicnemioides, Polytrichum commune, P. juniperinum, Sphagnum
magellanicum and S. recurvum as well as the lichens Cladonia andesita and C.
dydima.
Syntaxonomy: This subassociation is represented by 13 relevés, with 8 vascular
and 11 species of cryptogams. Paepalanthus pilosus (only by maximum cover),
Ortachne erectifolia, Breutelia squarrosa and Polytrichum commune are
diagnostic; as is the lichen Cladonia dydima.
122
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
Ecology and distribution: The vegetation of the subassociation ortachnetosum
erectifoliae covers a small area (approx 30-50 m2) to the northwestern edge of the
Laguna Seca in Páramo El Pumar at ~2870 m. This patch is surrounded by peat of
the association Sphagno recurvi-Caricetum bonplandii.
Figure 4.2. Physiognomy of the vegetation of the association of Paepalantho pilosiAgrostietum basalis. Páramo El Pumar, 2870 m. Lev. LS16. Ab: Agrostis
basalis; Br: Breutelia rythidioides; Bs: Breutelia squarrosa; Ca: Campylo-pus
albidovirens; Cd: Cladonia dydima; Cla: Cladonia andesita; Oe: Ortachne
erectifolia; Pc: Polytrichum commune; Pp: Paepalanthus pilosus; Sm:
Sphagnum magellanicum.
1. Paepalantho pilosi – Agrostietum basalis
1.2. subassociation typicum Cuello & Cleef 2009
Typus: Rel. No. 17 (Cuello LS26). Table 4.1
Open and low Agrostis basalis bunchgrass vegetation on peat bog with dominance of
Polytrichum juniperinum / Pajonal ralo y bajo de Agrostis basalis sobre turbera con
dominancia de Polytrichum juniperinum
Physiognomy and composition: Open and low vegetation with an herbaceous
layer (15-25 cm height) dominated by small tussocks of Agrostis basalis (1-20%
cover) and discrete individuals of Carex bonplandii (1-10% cover). The ground
layer is dominated by cushions of Arenaria venezuelana (12-40% cover),
Lachemilla verticillata (35-85% cover) and Polytrichum juniperinum (15-60%
cover). Sphagnum magellanicum and S. recurvum are also present in the ground
layer.
123
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Syntaxonomy: The subassociation typicum of the Paepalantho pilosi–Agrostietum
basalis is represented by 9 relevés, with 9 vascular species and 7 species of
cryptogams. Polytrichum juniperinum is diagnostic. This subassociation typicum is
separated from the previous subassociation by the absence of Ortachne erectifolia,
a very low presence of Breutelia squarrosa and Polytrichum commune, and by a
greater density and cover of Polytrichum juniperinum.
Ecology and distribution: The vegetation of the subassociation typicum covers a
patch at the southwestern end of the dry lake peat bog of Páramo El Pumar at
~2870 m. This side of the dry lake is lower and more humid than the northern side.
Near the higher and drier southern border of the dry lake, the vegetation of this
association is in contact with that of the Sphagno recurvi–Caricetum bonplandii
association.
Photo 4.3. Vegetation association on the northwestern edge of the Laguna Seca in Páramo
El Pumar at ~2870 m. Center-right: Paepalantho pilosi - Agrostietum basalis
subassociation ortachnetosum erectifoliae. Left: Sphagno recurvi - Caricetum
bonplandii.
2. Sphagno recurvi – Caricetum bonplandii Cuello & Cleef 2009
Typus: Rel. No. 35 (Cuello LS9). Table 4.1, Fig. 4.3
Sphagnum recurvum - Carex bonplandii peat bog / Turbera de Sphagnum recurvum y
Carex bonplandii
Physiognomy: Peat bog dominated by a dense green carpet of Sphagnum
recurvum with 100% cover.
124
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
Composition and syntaxonomy: This association is represented by 24 relevés,
with 13 vascular species and 8 species of bryophyte. The diagnostic species are
Sphagnum recurvum and Carex bonplandii.
Two provisional variants of this association are distinguished. The vegetation
of variant typicum has an open aspect. This common peat bog variant is
represented by 15 relevés and accounts for a total of 12 vascular species with very
low cover. Carex bonplandii (height 15-25 cm, cover 35-70%), growing on a
green carpet of Sphagnum recurvum (60-100% cover) is especially prominent. The
other vascular species present (in the association but) with very low cover include:
Agrostis sp. B, Arenaria venezuelana, Calamagrostis sp. Diplostephium obtusum,
Gentianella nevadensis, Hypericum cardonae, Lachemilla verticillata, Nertera
granadensis, Paepalanthus pilosus, Sisyrinchium sp., Xyris subulata var.
acutifolia.
The variant typicum lacks proper diagnostic species. The variant of Diplostephium
obtusum includes only 7 vascular species. The vegetation of this variant occurs
near the eastern dry edges of the evaporated lake, Laguna Seca. The presence of
Diplostephium obtusum is diagnostic (5-40% cover) with variable densities of
Carex bonplandii [10-50% (90%)], as is the presence of Agrostis sp. B.
Ecology and distribution: In its typical form, the vegetation of this association is
found on the humid shore of Laguna El Pumar, as well as in the central humid or
semi-humid areas of the West shore of Laguna Seca in Páramo El Pumar (~28702890 m), and in wet areas around a pond of water in a little valley South of „Las
Antenas‟ in Páramo de Guaramacal at ~3080 m.
3. Sphagno sparsi – Caricetum bonplandii Cuello & Cleef 2009
Typus: Rel. No. 56 (Cuello LP1). Table 4.1, Fig. 4.3, 4.4, Photo 4.3 (left to
bottom)
Sphagnum sparsum – Carex bonplandii peat bog / Turbera de Sphagnum sparsum y Carex
bonplandii
Physiognomy and composition: Peat bog that consists of an herb layer (15-25 cm
in height), covering between 10-80%, and dominated by Carex bonplandii. A
ground layer with 100% cover, formed by a continuous carpet of several
Sphagnum species, among which, S. sparsum dominates, followed by S. recurvum
and S. magellanicum. Also common are compact cushions of Paepalanthus
pilosus, and a variable cover of Campylopus cuspidatum.
In this association, a shrub layer made up of Diplostephium obtusum may be
present, or a layer of very low shrubs of Hypericum juniperinum, H. cardonae, H.
juniperinum x cardonae, and Pernettya prostrata.
Syntaxonomy: The association of Sphagno sparsi-Caricetum bonplandii is
represented by 16 relevés with 13 vascular species and 5 species of moss. The high
presence and cover of Sphagnum sparsum and S. magellanicum is diagnostic.
125
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Two variants are distinguished: one with Diplostephium obtusum, the other with
Pernettya prostrata.
Ecology and distribution: The peat bog of this association is located at the dry
northeastern shore at 1-6 m from the edge of Laguna Seca, and also on hummocks
and the non-flooded edges of Laguna El Pumar in Páramo El Pumar at 2880-2890
m.
Figure 4.3. Physiognomy of the vegetation of a hummock-hollow páramo peat bog of the
association of (1) Sphagno sparsi-Caricetum bonplandii var. Pernettya
prostrata and (2) Sphagno recurvi-Caricetum bonplandii at Laguna El Pumar,
Páramo El Pumar, 2880 m. Cb: Carex bonplandii; Gm: Geranium
stoloniferum; Hc: Hypericum cardonae; Hjxc: Hypericum juniperinum x
cardonae; Pp: Paepalanthus pilosus; Ppr: Pernettya prostrata; Sm:
Sphagnum magellanicum; Sr: Sphagnum recurvum. Ss: Sphagnum sparsum.
3.1. Variant with Diplostephium obtusum
Representative rel.: No. 46 (Cuello LSF85). Table 4.1, Fig. 4.4
Variante con Diplostephium obtusum
Physiognomy and composition: Peat bog of Sphagnum magellanicum and Carex
bonplandii with a shrub layer of Diplostephium obtusum (height 30-120 cm, cover
15-45%).
Syntaxonomy: The variant is represented by 9 relevés with a total of 7 vascular
species. The diagnostic species is Diplostephium obtusum, together with an
absence of Pernettya prostrata and associated species.
Ecology and distribution: The vegetation of this variant is present on the higher
and drier edges of the NE-SE part of the peat bog of Laguna Seca in Páramo El
126
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
Pumar. This community can also be found in small peaty valleys with drainage
embedded in azonal bamboo páramo of the association Carici bonplandii–
Chusqueetum angustifoliae.
Figure 4.4. Physiognomy of the bog of the association Sphagno sparsi-Caricetum
bonplandii var. Diplostephium obtusum. Páramo El Pumar. Laguna Seca.
2890 m. Lev. SF85. Do: Diplostephium obtusum; Cb: Carex bonplandii;
Gm: Geranium stoloniferum Pp: Paepalanthus pilosus; Sm: Sphagnum
magellanicum; Sr: Sphagnum recurvum Ss: Sphagnum sparsum.
3.2. Variant with Pernettya prostrata
Representative rel. Cuello LP1. Table 4.1, Fig. 4.3
Variante con Pernettya prostrata
Physiognomy and composition: Vegetation on hummocks near the edges of peat
bog dominated by a layer of Carex bonplandii (height 15-25 cm, cover 30-80%)
with a layer of a few low shrubs (height 5-40 cm, cover 1-40%) consisting of
Pernettya prostrata, Hypericum cardonae and H. juniperinum x cardonae. Scarse
young individuals of Hypericum juniperinum and Hesperomeles obtusifolia may
also be present among the shrubs.
A bryophytic ground layer is dominated by Sphagnum sparsum (30-100% cover),
S. recurvum (10-40% cover) and Campylopus cuspidatum.
Syntaxonomy: The variant is represented by 7 relevés with 9 vascular species and
5 moss species. Diagnostic species are: Pernettya prostrata, Hypericum cardonae
and Campylopus cuspidatum. Sphagnum magellanicum has a low presence and
cover when contrasted with the variant with Diplostephium obtusum.
Ecology and distribution: Vegetation on hummocks in the non-flooded areas
around of Laguna El Pumar at 2880 m, Páramo El Pumar.
127
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
CARICI BONPLANDII – CHUSQUEION ANGUSTIFOLIAE Cuello & Cleef 2009
Typus: Carici bonplandii – Chusqueetum angustifoliae
Azonal Carex bonplandii - Chusquea angustifolia bunchgrass-bamboo páramo alliance /
Páramo azonal de pajonal-bambusal (chuscales) de la alianza de Carex bonplandii y
Chusquea angustifolia
Physiognomy and composition: This alliance groups azonal bamboo páramo
(„chuscales‟) growing in humid level areas of low inclination dominated by
Chusquea angustifolia.
Syntaxonomy: This alliance is defined on the basis of 9 relevés with 14 vascular
species and 5 moss species. Carex bonplandii, Chusquea angustifolia, Sphagnum
sancto-josephense and Xyris subulata are the diagnostic species. The alliance
contains one association so far, Carici bonplandii–Chusqueetum angustifoliae.
Ecology and distribution: The vegetation of the alliance Carici bonplandii–
Chusqueion angustifoliae is found growing close to lakes shores in Páramo El
Pumar (2870-2890 m) and on small wet valleys in both Páramo El Pumar and
Páramo de Guaramacal (~2900-3100 m).
4. Carici bonplandii – Chusqueetum angustifoliae Cuello & Cleef 2009
Typus: Rel. No. 64 (Cuello L27a). Table 4.1, Fig. 4.5, Photo 4.3 (top)
Carex bonplandii - Chusquea angustifolia bunchgrass-bamboo páramo / Páramo de
pajonal-bambusal de Chusquea angustifolia con Carex bonplandii
Physiognomy: Dense bamboo páramo, or “chuscal”, with a bamboo layer of
Chusquea angustifolia (height 1-1.5 m, cover 30-70%), a herbaceous layer, 20-30
cm in height dominated by Carex bonplandii, and a ground layer dominated by
cushions of Sphagnum sancto-josephense and S. sparsum together with other
bryophytes and some lichens.
Composition and syntaxonomy: The association of Carici bonplandii - Chusqueetum angustifoliae is represented by 9 relevés with 14 vascular species and 9
bryophytes.
Chusquea angustifolia (dominant), Carex bonplandii and Sphagnum sanctojosephense are diagnostic of the assemblage. Agrostis perennans, Daucus montanus and Xyris subulata var. acutifolia are present in the herb layer. Paepalanthus
pilosus, Arenaria venezuelana, the bryophytes Breutelia squarrosa, Campylopus
subjugorum, C. pilifer, C. nivalis, Sphagnum sancto-josephense, S. sparsum, the
liverworts Jamesoniella rubricaulis, Lepidozia cf. macrocolea (3034), and Plagiochila sp., and the lichens Cladia aggregata and Peltigera neopolydactyla have also
been observed in the ground layer. On the canes of Chusquea angustifolia the
epiphytic moss Campylopus trichophorus can be found. In this association a provisional subassociation of Xyris subulata is distinguished by the presence of Xyris
subulata var. acutifolia (rel.nr. 61-65) together with a few other common species
of the zonal páramo association of Rhynchospora gollmeri - Ruilopezia jabonensis
128
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
which is in contact in some locations. More relevés are needed for the formal
description of this Xyris subulata subassociation.
Ecology and distribution: The bamboo vegetation of the association of Carici
bonplandii - Chusqueetum angustifoliae is found in level or concave areas of low
slope (1 to 8 degrees) with a southwestern-west exposure at altitudes between
~2880 - 3100 m. They are positioned adjacent to lake margins or covering small
wet valleys. This bamboo vegetation grows on relatively deep soils (50-120 cm),
with gray colors (dry) and very dark (humid), sandy-loamy to loamy textures. The
pH of the upper layer ranges from 3.6 to 4.2.
Figure 4.5. Physiognomy of the association of Carici bonplandii-Chusqueetum angustifoliae (L27, 3030 m). Cb: Carex bonplandii; Ch: Cortaderia hapalotricha;
Cha: Chusquea angustifolia; Rg: Rhynchospora gollmeri; Rj: Ruilopezia
jabonensis; Ss: Sphagnum sparsum; Ssj: Sphagnum sancto-josephense; Xs:
Xyris subulata var. acutifolia.
DISTRICHO SUBMERSI – ISOETION Cleef 1981
Table 4.2 rel. nrs 1-10
Alliance of submerged bryophytic-isoetid communities in páramo lakes described
from the Cordillera Oriental of Colombia (Cleef 1981).
5. Community of Sphagnum cuspidatum
Representative rel.: No. 4 (Cuello PL4). Table 4.2, rel. 1-5, Photo 4.4
Submerged aquatic community with Sphagnum cuspidatum present at great
density close the peaty shores of Laguna El Pumar. Water depth ranges between 30
and 120 cm. The community is also found in a small peaty depression with flushes
of water in bamboo páramo near the „Las Antenas‟.
6. Isoëtetum karstenii Cleef 1981
Table 4.2, rel. 6-8
129
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Flora, vegetation and ecology in the Venezuelan Andes
Submerged aquatic community of Isoëtes karstenii, associated with Eleocharis
acicularis (sterile) and filamentous algae thriving at a depth of between 0.6-1 m in
Laguna El Pumar (2890 m). In deeper areas of the lake (1-1.5 m) black
filamentous algae (cf. Microspora sp.) are also present within this community.
More relevés are needed in order to more clearly define a possible further
subdivision of the Isoëtetum karstenii.
Table 4.2. Phytosociological table of aquatic communities of Ramal de Guaramacal, Andes,
Venezuela.
Rel. Num.
Releves (Field Number)
HD
HD
HD
HD
HD
HD
1
2
3
4
5
6
7
8
9
10
GCC2 PL5 LS20 PL4* LP12 LP13 LP8 LP9 LP10 LP11
6. Isoëtetum
Community/Association 5. Sphagnum cuspidatm
karstenii
5. Community of Sphagnum cuspidatum
Sphagnum cuspidatum
5
5
5
5
5
1
.
.
.
.
Eleocharis acicularis
.
.
.
5
3
5
5
4
5
.
black filamentous Algae
.
.
.
.
5
.
.
.
4
5
purple filamentous Algae
1
.
.
.
.
.
.
.
.
.
6. Isoëtetum karstenii
gelatinous Algae
.
.
.
.
.
5
5
5
5
.
Isoëtes karstenii
.
.
.
.
.
4
3
5
.
.
LP, PL Laguna El Pumar, 2880 m. Páramo El Pumar
LS Laguna Seca, 2890 m. Páramo El Pumar
GCC Páramo de Guaramacal, 3030 m.
HD Hydrophyte
* Type relevé
Photo 4.4. Submerged aquatic community of Sphagnum cuspidatum in Laguna El Pumar.
130
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
______________________________________________________
4.5 DISCUSSION
Phytosociological classification and methodological limitations
The classification of azonal páramo vegetation of Guaramacal resulted in one new
order, two new alliances, one earlier described alliance and six associations. Four
of them are described as new syntaxa, one as a provisional community whilst one
association (Isoëtetum karstenii Cleef 1981) was previously known from
Colombia. A summary (presence) table of azonal páramo vegetation communities
of Ramal de Guaramacal is shown in Table 4.3. The vegetation has been described
on the basis of a relatively limited number of relevés from only two peat bogs and
a small pond from two main páramo areas of Ramal de Guaramacal (Páramo de
Guaramacal and Páramo El Pumar). Other azonal vegetation communities may be
present in the páramos of Ramal de Guaramacal as other peat bogs are known to
exist in the area but have not yet been reached and remain as yet unexplored. The
limited accessibility of the area throughout most of the year, together with high
precipitation levels and the frequency of mist, made the exploration of peat bogs
areas of Ramal de Guaramacal extremely difficult, hence limiting the study of the
vegetation to only the drier climatic conditions at only the most accessible sites.
During these drier spells, some annual species were found in a senescent condition
making taxonomic identification difficult. Some plants could even be ignored in
the survey as they could already only persist as seeds in the seed bank.
However, the low floristic diversity observed in azonal communities of Ramal de
Guaramacal can be mainly attributed to the stress caused by extreme humidity with
a subsequent dominancy of only a few well-adapted species. Also the relative
isolation from the main system of the Cordillera de Mérida is probably a factor. In
the Laguna Seca the substrate of the lake bottom remains humid, even in the dry
season, sometimes with a small pond.
An important issue is the (almost) absence of proper diagnostic species in the
Sphagno recurvi-Caricetum bonplandii of the Agrostio-Paepalanthion. This
phenomenon corresponds to the „central syntaxon concept‟ of Dierschke (1981,
1994). The almost absence of diagnostic species is differential against both other
associations.
Azonal bunchgrass patches
Azonal bunchgrass páramo is represented by two small patches of vegetation
belonging to associations of the new alliance Paepalantho pilosi - Sphagnion
recurvi; both of which grow on top of a former peat bog. These bunchgrass
communities are very restricted in both surface area covered and spatial location in
Guaramacal, thus comparison (in ecology and floristic composition) with other
communities elsewhere is limited. As far as we are aware, no similar communities
have been reported from páramos. The presence of these communities, just on the
border of the evaporated lake in Páramo El Pumar and on both sides of a remnant
pond, suggests a relationship with wildlife in the origin of these communities.
Páramo El Pumar got its name by the apparent abundance of the „Puma‟ Puma
concolor, as indicated by the observed large quantities of vestiges, such as paw
prints and the remains of digested prey. The evaporated lake „Laguna Seca‟ is
131
______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
surrounded by patches of dwarf high Andean forest which offer shelter to these
animals which appear to walk across the peat bog to drink or to hunt prey.
The bunchgrass Ortachne erectifolia was previously described by the second
author in 1981 as a being a species with a wide ecological range occurring between
about 3500 and 4300 m. A bunchgrass community of Ortachne erectifolia
(Lorenzochloetum erectifoliae Cleef 1981) is known from the dry zonal
bunchgrass páramos at 3550-3650 m in the Colombian Cordillera Oriental (CLEEF
1981). That community, however, differs greatly in both ecology and floristic
composition and limits comparisons with the subassociation ortachnetosum
erectifoliae of the azonal Paepalantho pilosi–Agrostietum basalis from
Guaramacal. The bunchgrass Ortachne erectifolia is also a common species in the
zonal widespread grass páramo community of Espeletia schultzii–Aciachne
acicularis in the Sierra Nevada de Mérida (Fariñas 1980, Berg 1998, Berg & Suchi,
2001). This characteristic, medium-sized bunchgrass species with stiff blades is
also present in páramos of Costa Rica, Ecuador and Peru (Luteyn 1999, Briceño &
Morillo 2006). The original Ortachnetum erectifolii is considered secondary
vegetation having developed after severe disturbance, probably by fire (Cleef
1981).
The other lax and low bunchgrass vegetation growing on former peat bog,
characterized by the presence of Agrostis basalis and Polytrichum juniperinum, the
subassociation typicum, is known only from this site to date. Agrostis basalis is an
endemic species described from the Sierra Nevada de Mérida páramos (Laguna
Negra) (Luces 1953) and has also been reported from Distrito Federal, Mérida,
Miranda and Táchira states, where it is found growing between 2100 and 4150 m
(Briceño & Morillo 2006; Hokche et al. 2008).
Sphagnum bogs
A regional study on the Sphagnum bogs of the northern Andes is still lacking as
most studies report only on local peat bog types. Some of the azonal vegetation
communities reported for the páramos of the Colombian Cordillera Oriental (Cleef
1981; Franco et al. 1986; Sanchez & Rangel 1990; Rangel 2000a) are also found in
the lowermost superpáramos of Sierra Nevada de Mérida, such as: the
Aciachnetum pulvinatae, the Wernerion community (Wernerietalia), communities
with Carex bonplandii and communities with Gentiana sedifolia (Berg 1998; Berg
& Suchi 2001). However, Sphagnum bog communities have not yet been formally
reported despite being present in the páramos of the Sierra Nevada de Mérida.
With regards to the Chusquea angustifolia páramos of Ramal de Guaramacal,
affinities to other páramo communities and comparisons are limited. There are few
species common to some of the vegetation types described for the Colombian
Cordilleras (e.g. Cleef 1981; Cleef et al. 2005; 2008; Restrepo & Duque 1992,
Franco et al. 1986; Sánchez & Rangel 1990). Curiously, Gentiana sedifolia,
present in páramo and puna bogs in the tropical Andes, is lacking in Guaramacal
páramo bogs. Isolation, low altitude and a deficit of phytosociological studies
account for the presence of the assemblage of species observed in Ramal de
Guaramacal which remain undescribed for other páramo areas to date.
132
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
_______________________________________________________
Table 4.3. Presence table of azonal páramo vegetation communities of Ramal de
Guaramacal, Andes, Venezuela. Associations: 1. Paepalantho pilosi Agrostietum basalis; 2. Sphagno recurvi - Caricetum bonplandii; 3. Sphagno
sparsi - Caricetum bonplandii; 4. Carici bonplandii - Chusqueetum
angustifoliae; 5. Community of Sphagnum cuspidatum; 6. Isoetetum karstenii.
Presence classes: I (0–20%), II (21–40%), III (41–60%), IV (61–80%) and V
(81–100%).
Number of relevés
22 22 16
9
5 3
Number of relevés
22 22 16
9
5 3
Association
1
2
3
4
5 6
Association
1
2
3
4
5 6
Agrostis basalis
V
I
.
.
.
.
Calamagrostis sp. A
.
.
.
I
.
.
Campylopus albidovirens
IV
.
.
.
.
.
.
.
.
V
.
.
Arenaria venezuelana
III
I
.
I
.
.
Lachemila verticillata
Chusquea angustifolia
Sphagnum sanctojosephense
.
.
.
IV .
.
III
.
.
.
.
.
Breutelia rhythidoides
Xyris subulata
.
.
I
III .
.
I
.
.
.
.
.
Ortachne erectifolia
Campylopus richardii
.
.
.
II
.
.
IV
.
.
.
.
.
Breutelia squarrosa
Polytrichum commune
Ruilopezia jabonensis
.
.
.
II
.
.
III
.
.
II
.
.
III
I
.
.
.
.
Cortaderia hapalotricha
.
.
I
II
.
.
Cladonia dydima
II
.
.
.
.
.
Gentianella nevadensis
.
.
I
II
.
.
Cladonia andesita
I
.
.
.
.
.
Peltigera neopolydactyla
.
.
.
II
.
.
Rhynchospora gollmeri
I
.
.
II
.
.
Oreobolus venezuelensis
.
.
.
I
.
.
Cladia aggregata
I
.
.
.
.
.
Hieracium avilae
.
.
.
I
.
.
III
II
.
.
.
.
Jamesoniella rubricaulis
.
.
.
I
.
.
Sisyrinchium sp.
I
I
.
.
.
.
.
.
.
I
.
.
Calamagrostis bogotensis
I
.
.
.
.
.
Nertera granadensis
Ophioglossum
crotalophorioides
.
.
.
I
.
.
.
.
.
I
.
.
Polytrichum juniperinum
Sphagnum sparsum
.
I
V
II
.
.
Plagiochila sp.
Sphagnum magellanicum
II
I
IV
.
.
.
Rhacocarpus purpurascens
.
.
.
I
.
.
.
Agrostis perennans
.
.
I
.
.
.
III
V
V
V
.
.
.
Diplostephium obtusum
.
II
II
.
.
Agrostis sp. B
.
I
.
.
.
.
Carex bonplandii
Pernettya prostrata
.
.
III
.
.
.
Geranium stoloniferum
III
I
III
II
.
Sphagnum cuspidatum
.
.
.
.
V 1
Eleocharis acicularis
.
.
.
.
II 5
black filamentous Algae
.
.
.
.
I .
purple filamentous Algae
.
.
.
.
I .
gelatinous Algae
.
.
.
.
. 5
Isoëtes karstenii
.
.
.
.
. 5
Campylopus cuspidatus
Hypericum juniperinum x
cardonae
I
.
II
.
.
.
.
.
I
.
.
.
Hypericum cardonae
.
I
I
.
.
.
Hypericum juniperinum
.
.
I
.
.
.
Sphagnum sp.(orange)
.
.
I
.
.
.
Hesperomeles obtusifolia
.
.
I
.
.
.
Sphagnum recurvum
III
V
V
Paepalanthus pilosus
V
II
II
I
I
.
.
133
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Flora, vegetation and ecology in the Venezuelan Andes
Sphagnum bogs in the páramos of Ramal de Guaramacal are represented by two
new associations belonging to the new alliance of Carici bonplandii - Sphagnion
recurvi. Sphagnum revurvum, S. sparsum, S. magellanicum, and S. sanctojosephense are the most characteristic species of the Sphagnum bogs of
Guaramacal. Sphagnum cuspidatum is also common mostly in submerged
conditions, whilst S. meridense is present with large cover in humid shrub páramo
and adjacent dwarf forest edges. Sphagnum recurvum is the most dominant species
in the peat bogs of Guaramacal. The moss cover of both associations of Sphagno
recurvi - Caricetum bonplandii and Sphagno sparsi - Caricetum bonplandii grow
together in the same peat bogs of Páramo El Pumar. The variants of each
association correspond to different succesional stages related decreasing humidity
(see below).
A Bray-Curtis cluster similarity analysis comparing species composition of azonal
Guaramacal Carex bonplandii associations with those of the Carex bonplandii
communities described from Colombian cordilleras and the Sierra Nevada de
Mérida, Venezuela, is shown in Fig. 4.8. The presence of Carex bonplandii and
Sphagnum spp. has been reported in peat bog vegetation in Tatamá Park in the
Colombian Western Cordillera (Cleef et al. 2005). An association of Caricetum
bonplandii has been described from Laguna Chingaza (Franco et al. 1986, Rangel
2000c) and a Sphagnum sancto-josephense - Carex bonplandii community from
Páramo de Monserrate (Vargas & Zuluaga 1985), both sites being near Bogotá in
the Colombian Oriental Cordillera. Despite the common presence of Carex
bonplandii, Sphagnum magellanicum and S. sancto-josephense in Sphagnum
peatbog communities in Colombia, there are no other common species which
allow establishment of relationships to the Sphagnum bog communities of
Guaramacal. It is evident that the Guaramacal Sphagnum recurvum communities
are most related to each other (Fig. 4.8). Similarities to other Sphagnum
communities collected hap-hazardly in literature deal with different habitats (with
different ecology): Sphagnum bog in morrainic valleys, more minerotrophic
conditions with Sphagnum cover, Sphagnum fringes along glacial lake shores, and
Sphagnum cover on different geological substrates.
A coherent and representative body of relevés is lacking for a safe approach to
classify the Sphagnum bogs of the northern Andes, as outlined above. The second
author has some 60 unpublished relevés of Sphagnum bogs, mainly of the páramos
of the Eastern Cordillera of Colombia (Cleef 1981). However, it was not the aim of
the present study to develop a rather complete syntaxonomic scheme of páramo
Sphagnum bogs. This is a task for the future. This is also the reason that we did not
like to produce presence tables in our study, because the material published so far
is too scanty, making the effort not meaningful.
Aquatic communities
Two submerged aquatic communities were recognized in páramos of Ramal de
Guaramacal: (1) the association Isoëtetum karstenii and (2) the community of
Sphagnum cuspidatum (Table 4.2). The The presence of Isoëtes karstenii of
Laguna El Pumar shows a relationship of this low altitude páramo vegetation with
other proper upper páramo aquatic communities observed in páramo lakes of the
134
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
_______________________________________________________
Sierra Nevada de Mérida and described from such lakes in the Colombian
Cordillera Oriental. The aquatic association Isoëtetum karstenii was documented
from cold lakes, mostly with mineral bottoms, in the grassparamo (3500-3700 m)
up to the superpáramo at 4425 m of the Sierra Nevada del Cocuy, and up to 4100
m in the Sumapaz páramo of the Colombian Cordillera Oriental (Cleef 1981). The
association has also been found at 4300 m on the volcano S. Isabel in the
Colombian Cordillera Central (Salamanca et al. 2003) and further south to Nariño,
southern Colombia. The association Isoëtetum karstenii belongs to the alliance of
Ditricho submersi-Isoëtion (Cleef 1981). In the Sierra Nevada de Mérida in
Venezuela, Isoëtes karstenii has been collected between 3430 and 4250 m (Cleef
1981, unpubl.; Small & Hickey 2001). One relevé (Cleef 552A) of Isoëtetum
karstenii typicum has been made by the second author at 4250 m in the lower
superpáramo of La Culata (see Table 4.4). Isoëtes karstenii grows submerged in
permanent lakes and ponds (occasionally streams) between ca. 3300-4600 m. The
occurance of Isoëtes karstenii in Guaramacal is the lowest recorded and could be a
relict from Glacial times. Its habitat generally corresponds to the upper páramo
proper and the superpáramo. During Glacial times, it is most likely that these lakes
on the top of the Ramal de Guaramacal range were part of the superpáramo. Shifts
upslope under Holocene conditions was impossible because the present lake is on
top of the ridge of Guaramacal. With increasing temperature and humidity (now a
bamboo páramo in nature) the Isoëtes karstenii plants survived, growing on an
organic lake bottom, and became associated with other plant species of peaty lake
bottoms, such as: Eleocharis acicularis, Sphagnum cuspidatum and diverse algae.
Table 4.4. Table of presence of Isoetetum karstenii in páramo areas of Colombia and
Venezuela. Presence classes: I (0–20%), II (21–40%), III (41–60%), IV (61–
80%) and V (81–100%). *cover values in percentage. Sites: (1). Páramos
Cocuy, Sumapaz, Colombian Cordillera Oriental (Cleef 1981); (2). Lev. A.M.
Cleef & S. Salamanca # 622A and #584. Laguna de Silencio, Base de S. Isabel.
Alt. 4170-4315 m. (Parque Los Nevados), Colombian Cordillera Central
(Salamanca et al. 2003); (3). Lev. Cleef 552A (with A. Chaverri & O. Rangel).
Venezuela, Páramo La Culata, superpáramo bajo. Lagunita glaciar a 4.250 m.;
(4). Laguna El Pumar, 2880 m. Ramal de Guaramacal, Andes, Venezuela.
Isoëtetum karstenii
Number of releves
Altitude (m)
Site number
Isoëtes karstenii
Blindia magellanica
Ditrichum submersum
Eleocharis acicularis
Isotachis serrulata s.l.
Sphagnum cuspidatum
Algae
Cord. Oriental
Colombia
Cord. Central
Colombia
8
3500-3700
1
V
I
II
.
II
.
V
2
4170-4350
2
5
.
3
.
.
.
.
Páramo La
Culata, Mérida,
Venezuela
1
4250
3*
(80)
.
.
.
.
.
(1)
Guaramacal
Venezuela
(this study)
3
2880
4
5
.
.
5
.
1
5
135
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Under these conditions we expected the presence of Isoëtes palmeri, known from
lower páramo lakes with gyttja bottom in the Sierra Nevada de Mérida and the
Eastern Cordillera of Colombia (Cleef 1981; Small & Hickey 2001). It would
appear that, to date, the latter mentioned species has not yet arrived in Guaramacal.
Table 4.5. Table of presence of the Sphagnum cuspidatum community from Guaramacal
combined to those S. cuspidatum communities from páramo areas near Bogota,
Colombia. Presence classes: I (0–20%), II (21–40%), III (41–60%), IV (61–
80%) and V (81–100%).* cover values in percentage. (1 & 2) Cordillera
Oriental (Cleef 1981); (3) Norte de Pantano Largo, Flanco Noroeste en el
Páramo de Guargua. Colombia (Sánchez & Rangel 1990); (4) Lev. Cleef 54.
Páramo de Palacio, Laguna Buitrago, 3.620 m. Cundinamarca. Colombia; (5)
Laguna El Pumar, 2880 m. Ramal de Guaramacal, Andes, Venezuela (this
study).
Num. relevés
Alt. (m)
Site
Sphagnum cuspidatum
Eleocharis acicularis
Eleocharis stenocarpa
‘black/purple filamentous Algae’
‘gelatinous Algae’
Bartsia sp.
Carex bonplandii
Carex aff.pygmaea
Carex pichinchensis
Juncus stipulatus
Juncus breviculmis
Lysipomia sphagnophila ssp. minor
Nertera granadensis
Gentianella corymbosa
Halenia gentianoides
Lepidozia macrocolea
Riccardia smaragdina
Breutelia chrysea
Riccardia hansmeyeri
Sphagnum magellanicum
Campylopus cuspidatus var.
dicnemoides
Pleurozium schreberi
Chisaca
Neusa
Guargua
Palacio
1
3625
1
3
5
.
5
5
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
3690
2
2
30
.
1
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
3480-3730
3
V
1
3620
4*
95
III
.
.
.
II
.
IV
.
I
.
III
.
.
.
.
.
.
.
II
.
.
.
<1
<1
10
<1
.
<1
<1
30
5
2
1
1
<1
Guaramaca
l
5
2890
5
V
II
.
I
I
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
<1
.
<1
Also only in site 3:
Campylopus pittieri (III), Peltigera sp. (III), Agrostis sp. (II), Blechnum loxense (II), Calamagrostis
effusa (II), Hydrocotyle bonplandii (II), Hypericum myricariifolium (II), Lachemilla fulvescens (II),
Paspalum bonplandianum (II), Pernettya prostrata (II), Plagiocheilus solivaeformis (II), Riccardia sp.
(II), Rubus acanthophyllos (II), Valeriana longifolia (II), Agrostis tolucensis (I), Brachythecium sp. (I),
Callitriche nubigena (I), Cortaderia bifida (I), Festuca sp. (I), Grammitis moniliformis (I), Hieracium
avilae (I), Hypotrachyna sp. (I), Laestadia muscicola (I), Niphogeton ternata (I), Pentacalia abietina
(I), Pentacalia nitida (I), Puya santosii (I), Rhynchospora macrochaeta (I).
136
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
_______________________________________________________
Sphagnum cuspidatum aquatic communities of Laguna El Pumar show some
relationships to other communities with S. cuspidatum described by Cleef (1981)
from páramo areas near Bogotá in the Colombian Cordillera Oriental due to the
common presence of Sphagnum cuspidatum, Eleocharis acicularis and undetermined filamentose algae. Another vegetation community with Sphagnum cuspidatum
was also previously described near Bogotá by Sánchez & Rangel (1990), such as
the Carici-Sphagnetum cuspidati (Sánchez & Rangel l.c.). However, this is a peat
bog vegetation community with only Sphagnum cuspidatum in common with the
aquatic community from Guaramacal. One relevé (Cleef 54) has apparently been
taken under very similar ecological conditions as in Guaramacal; however, no
other species besides S. cuspidatum are common to it (see Table 4.5).
In Sphagnum peat bog zones of Europe, the association Sphagnetum cuspidatoobesi Tüxen & von Hübschmann 1958 em. Schaminée et al. has been recognized.
The stages of succession are as those contained in the association Sphagno
cuspidate - Rhynchosporetum albae Osvald 1923 with subassociations marking
hydroseral succession: sphagnetosum cuspidati and sphagnetosum recurvi. The
peat of Caricetum limosae Osvald 1923 em. Dierssen 1982 is slightly richer in
nutrients; Carex bonplandii could be a vicariant sedge species. All these European
Sphagnum peatbog communities belong to the class of Scheuchzerietea Den Held,
Barkman & Westhoff 1969. Some prominent bryophyte species are common to the
Sphagnum cuspidatum community from the Venezuelan Andes.
The Guaramacal relevés from the Eleocharis acicularis community compare easily
to the community of the same species from the Colombian Eastern Cordillera
páramos at between 3550-3850 m (Cleef 1981, Table 4.9). In the Holarctic zone of
the northern hemisphere, these communities have been described under the class
Littorelletea Br.-Blanquet & Tüxen. The temperate association Littorrello uniflorae
- Eleocharitetum acicularis Malcuit 1929 and the alliance Eleocharition acicularis
Pietsch 1966 em. Dierssen 1975 are distributed from Iceland to NW Europe.
Diagnostic species for the association and alliance are: Eleocharis acicularis,
Elatine hexandra and Echinodorus repens. Dierssen (1975) ranks E. acicularis as
a species with a wide ecology, mostly forming monospecific communities with
regional companions (‘races’). This habit is apparently also shared by the
neotropical plants of E. acicularis.
Aquatic/bog vegetation, dominated by Eleocharis acicularis and Sphagnum
recurvum, has been documented for the peat bog area of Laguna La Chonta at
2310 m in the Costa Rican Cordillera de Talamanca (Brak et al. 2005). Ruthsatz
(1977) also refers to Eleocharis acicularis growth in shallow puna lakes in
northern Argentina (3500-3800 m).
Deil (2005) reviewed the worldwide ephemeral vegetation inclusive of the
amphibic communities described thus far. With relevance to our case Sphagnum
cuspidatum and Eleocharis acicularis communities have been discussed, as has the
Ditricho-Isoëtion karstenii Cleef 1981 alliance which also includes the Guaramacal
Eleocharis acicularis - Isoëtes karstenii lake bottom vegetation.
137
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Spatial distribution and succession of vegetation communities
A detailed map of spatial distribution of vegetation types associated with the lakes
of Páramo El Pumar are shown in Figure 4.6. A schematic showing the hydroseral
sequence of vegetation communities from open water in the central part of the
Laguna El Pumar towards the shore is presented in Fig. 4.7.
Figure 4.6. Vegetation map of Laguna Seca, Páramo El Pumar, 2890 m. Ramal de
Guaramacal, Andes. Venezuela.
The submerged community of Isoëtetum karstenii is followed or surrounded by
Eleocharis acicularis. Next, there are dense masses of submerged Sphagnum
cuspidatum with E. acicularis. Towards the marshy shore there is the typicum
variant of the Sphagno recurvi - Caricetum association. The Sphagno recurvi –
138
The páramo vegetation of Ramal de Guaramacal: 2. Azonal vegetation
_______________________________________________________
Caricetum bonplandii is associated with the more humid areas of the peat bog,
representing an earlier succesional stage, which is fully dominated by Sphagnum
recurvum in the wet shallow areas and on hummocks with the shores being first
colonized by Carex bonplandii. The Sphagno sparsii - Caricetum bonplandii
corresponds to a later succesional stage and is present on drier areas as shown also
in Figure 4.3. The drier hummocks with the association of Sphagno sparsi Caricetum bonplandii near the shores are further colonized by small prostrate
dwarfshrub species which form the variant of Pernettya prostrata.
Figure 4.7. Physiognomy and hydroseral sequence of the vegetation associations of Laguna El
Pumar: (1) Association of Sphagno recurvi - Caricetum bonplandii. (2). Community
of Sphagnum cuspidatum. (3) Isoetetum karstenii. Cb: Carex bonplandii; Ea:
Eleocharis acicularis; Ip: Isoëtes karstenii; Sc: Sphagnum cuspidatum; Sr:
Sphagnum recurvum. Ss: Sphagnum sparsum.
Bamboo páramo
As discussed in Cuello & Cleef (2009b, Chapter 3), Chusquea angustifolia
bamboo páramos had not previously been studied in Venezuela. Bamboo páramo
communities of Chusquea angustifolia or Chusquea spencei have been reported
for the wet páramos of Táchira state (Bono, 1996). Several azonal bamboo páramo
communities (‘chuscales’), dominated by the bamboo species Chusquea tessellata
growing in wet páramo areas, have been widely documented from Andean
páramos along the Colombian cordilleras (e.g., Cleef 1981, Rangel 2000, Rangel et
al. 2006, Cleef et al. 2006, 2008)
139
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Figure 4.8. Cluster analysis comparing species presence values of azonal Guaramacal Carex
bonplandi páramo associations with those azonal Carex bonplandii páramo communities of the Colombian Cordilleras and Sierra Nevada de Mérida, Venezuela.
Sites: (1) Caricetum bonplandii, Laguna de Chingaza, Cordillera Oriental, Colombia
(Franco et al. 1986); (2) Caricetum bonplandii, Tatamá massif, Cordillera Occidental, Colombia (Cleef et al. 2005); (3) Junco effuse - Caricetum bonplandii, Páramo Frontino, Cordillera Occidental, Colombia (Rangel et al. 2005); (4) Swamp
with Carex, Llano de Paletara, Cordillera Central, Colombia (Restrepo & Duque,
1992); (5) Peat bog S of Bogota, Chisacá, Cordillera Oriental, Colombia (Sánchez &
Rangel, 1990); (6) Sphagno-Caricetum bonplandii, Páramo de Monserrate, Colombia
(Vargas & Zuluoaga, 1985); (7) Comunity of Carex bonplandii-Lachemilla sprucei,
Sierra Nevada de Mérida, Andes, Venezuela (Berg, 1998); (8) Sphagno sparsiCaricetum bonplandii, Guaramacal, Andes, Venezuela (this study); (9) Sphagno
recurvi-Caricetum bonplandii, Guaramacal, Andes, Venezuela (this study); (10)
Carici bonplandii - Chusqueetum angustifoliae, Guaramacal, Andes, Venezuela (this
study).
140
Chapter 5
Phytogeography of the vascular páramo flora of Ramal de
Guaramacal (Andes, Venezuela) and its ties to other páramo floras
Nidia L. Cuello A., Antoine M. Cleef and Gerardo A. Aymard C.
The text of Chapter 5 has been submitted to FLORA (general part; to be accepted after
review) and to ANALES DEL JARDÍN BOTÁNICO DE MADRID (Venezuelan part, under review)
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
5.1 INTRODUCTION
Páramo is the open equatorial alpine vegetation located above the upper forest line
(UFL) and below the permanent snow line from the northern Andes to Panamá and
Costa Rica. Páramo flora is considered the high-mountain flora most rich in
species of the world (Smith & Cleef 1988). Phytogeographical studies at the
generic level have shown that páramo flora has evolved mainly by immigration of
cool-adapted plants from temperate regions (temperate elements) and, in relatively
lower proportion, by adaptation of lower-elevation plants (tropical elements) to
high-altitude environments and by speciation through repeated isolation in situ
(Van der Hammen & Cleef 1986, Smith & Cleef 1988; Cleef & Chaverri 1992;
Ramsay 1992; Ricardi et al. 1997; Sklenář & Balslev 2007).
Páramo areas in Venezuela exhibit great environmental variability in climate at
regional and local scales. Through the about 400 km southwest to northeast
extension of the main Venezuelan Andean mountain chain, the Cordillera de
Mérida, there is a wide range of páramo hydrological conditions, from dry
páramos with 650 mm/year in a single rainy season, to permanently humid
páramos with over 3000 mm distributed throughout the year (Monasterio & Reyes
1980). The latter conditions characterize the páramo areas of Ramal de
Guaramacal, an outlier and comparatively low elevation (3130 m) range located at
the northeastern end of the Venezuelan Andes (Fig. 5.1).
North Andean páramo vegetation has been divided into several altitudinal zones
(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; Luteyn 1999; Hooghiemstra et al. 2006; Rangel-Ch. 2000a). For Venezuelan
páramos, Monasterio (1980a) 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).
Studies of phytogeography of the Venezuelan páramo flora started with a first
approach of the worldwide distribution of Venezuelan páramo flora presented by
Faría (1978) after the publication of the 'Flora de los Páramos de Venezuela' by
Vareschi (1970). This very first flora of the páramos was not complete, but
anyway representative.
Local floristic listings and phytogeographical analyses that include páramo areas
such as those from Táchira and Trujillo states have appeared (Bono 1996; Ortega
et al. 1985; Rivero & Ortega 1989; Dorr et al. 2000; Aymard 1999). Bono (1996)
also included a phytogeographical breakdown into geographic flora elements of
the páramo flora of Táchira State, Venezuela.
More recent phytogeographical analyses of the Venezuelan páramo flora have
been published by Ricardi et al. (1997, 2000). The first study deals with the
phytogeography of the Mérida superpáramo; the second study highlights the Sierra
Nevada de Mérida as a new phytogeographical subprovince of the northern Andes.
Briceño & Morillo (2002, 2006) recently published a list of the flowering species
143
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
of the Venezuelan Andean páramos, first the dicots, later followed by the
monocots.
The aim of this study is to analyse the phytogeographical affinities of the low
altitude and wet páramo of Ramal de Guaramacal in order to contribute to a better
understanding of the distribution, origin, and diversity of its flora. Particular
emphasis is given to the analysis of the floristic connections of the Guaramacal
páramo flora with the neighboring dry páramos of the Sierra Nevada de Mérida
and other páramo floras of the northern Andes and Central America.
One of our main objectives was to determine whether the phytogeographical
analysis and patterns of the páramo flora of Ramal de Guaramacal are determined
by temperature (a function of altitude) as has been established in previous studies
(e.g. Cleef 1979; Mérida Andes, Ricardi et al. 1997, 2000) or more by the overall
humidity, which characterizes the Guaramacal bamboo páramo. We have some
indications that ambient humidity may play a role, e.g. in the case of the bamboo
páramo of Tatamá (Cleef 2005), the páramos of Podocarpus National Park (PNP)
in southern Ecuador (Lozano et al., 2009) and also in the Talamancas of Costa
Rica (Cleef & Chaverri 1992).
5.2 STUDY AREA
Ramal de Guaramacal is located south of the town of Boconó, Trujillo state,
approximately 120 km Northeast of Mérida, in the centre of the Sierra Nevada de
Mérida (Fig. 5.1). Páramo areas of the summit of Ramal de Guaramacal are found
between 2800-3100 m, in the surroundings and between of 'Las Antenas' area (9 o
14’ 1.02” N; 70o 11’ 6.47” W) and Páramo El Pumar (9o 12’ 45.6” N; 70o 12’
5.55” W), 2.5 km Southwest of 'Las Antenas'.
The climate is very humid. According the first climatic records of the Davis Pro 2
climate station installed near the summit of Guaramacal (3100 m) by the first
author beginning in December 2006, there are over 290 days/year of rain.
Maximum precipitation occurs during April - July. Yearly precipitation is high,
reaching over 3200 mm/year and relative humidity attains 100% most of the year.
Temperatures remain low throughout the year with a diurnal temperature variation
from 4-6 oC to 14-16 oC; mean minimum temperature of 5.3 oC and mean maximum of 12.3o C; the lowest temperatures recorded being between -0.1-1.3 oC in the
month of January; the highest between 17.8-18.3 oC in the month of March, with
mean yearly temperature of 8.1-8.6 oC for the period from December 2006 - July
2009. Dominant wind directions are of ESE, SE and WNW, with a registered
average speed of 3.9-5.8 km/h. Maximum wind speed registered has been of 77.2
km/h, SE in the month of July 2008.
The vegetation of the Páramo of Guaramacal characterized by a mosaic of
subpáramo formations (shrub páramo, bunchgrass páramo, most common bamboo
páramo), intermingled with patches of dwarf forests (Subalpine Rain Forest or
SARF sensu Grubb 1977), distributed between 2800 and 3130 m (Cuello & Cleef
144
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
2009a, b). For detailed information on forest and páramo vegetation of Guaramacal we refer to Chapters 2-4 and Cuello and Cleef (2009a, b, c)
The study and full inventory of the flora of the whole Ramal de Guaramacal range
is still ongoing. Preliminary accounts of the vascular flora were first presented by
Ortega et al. (1988) and later by Dorr et al. (2000). After that, several new records
for the flora as well as new species to science have been documented for
Guaramacal (Taylor 2002; Stergios & Dorr 2003; Stančík 2004; Niño et al. 2005;
Cuello & Aymard 2008). A species inventory from páramo areas, including,
páramo and subpáramo-connected dwarf forest vegetation islands is presented in
this study (Appendix 5).
Figure 5.1. The location of Guaramacal páramo study site (G) and the other páramo areas in
northern South America and Central America which floristic comparison are
made: Sierra Nevada de Mérida (SNM) in Venezuela, Talamancas páramos
(PT) in Costa Rica – Panamá; Sierra Nevada del Cocuy (SNC), Serranía de
Perijá (P), Tatamá massif (T) and Sumapáz páramo (S) in Colombia; and
Podocarpus National Park (PNP) in southern Ecuador.
145
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
5.3 METHODS
Páramo data were collected from phytosociological studies (Cuello & Cleef 2009b,
c) and 585 numbers of general plant collections made by the first author from
páramo areas of Ramal de Guaramacal. Additional information was obtained from
herbarium collections and database of Herbario Universitario PORT, UNELLEZ in
Guanare. The data set includes a total of 251 vascular plant taxa belonging to 153
genera and 69 families that are listed in Appendix 5.
For each vascular genus listed the present geographical distribution has been
determined on basis of Mabberley (2008); occasionally also recent phylogenetic
studies (e.g. Chacón et al. 2006: Oreobolus; von Hagen & Kadereit 2003: Halenia;
Meudt & Simpson 2007: Ourisia, etc). Species distribution was also determined by
literature and by the W3Tropicos database. Plant genera have been grouped into
different phytogeographical elements belonging to three mayor components
according to Cleef (1979, 1981, and 2005) and Cleef & Chaverri (1992).
1) THE TROPICAL COMPONENT is made up of four flora elements:
(a) Wide tropical (WTR) taxa;
(b) Andean alpine (NT-AA) taxa;
(c) Páramo endemics (P);
(d) Neotropical montane elements (NT-M )
Thus, the former ‘Other Neotropical elements’ (Cleef 1979), viz. ‘Neotropicalmontane element’ (Cleef & Chaverri 1992) is subdivided into the Andean alpine
element (NT-AA) and Neotropical montane element (NT-M) following Simpson
& Todzia (1990) and Sklenář & Balslev (2007).
2) TEMPERATE COMPONENT contains three flora elements:
(a) Widely distributed temperate (WTE) taxa;
(b) Holarctic (HO) groups;
(c) Austral-Antarctic (AA) taxa.
3) COSMOPOLITAN COMPONENT consists of only the Cosmopolitan taxa (CO).
For a biogeographical analysis into species level, overall species distribution was
grouped into ten different geographic elements, adapting from previous
phytogeographical studies in the Andean region such as those used by Kelly et al.
(1994) and Schneider (2001). From the total 251 taxa recorded for the Guaramacal
summit area, for the specific biogeographical analysis we used only 224 species
with a defined distribution (those which were determined to species and/or
146
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
infraespecific level), including all open páramo and dwarf forest islands (of
Subalpine Rain Forest or SARF sensu Grubb 1977) vegetation species.
Floristic relationships of Guaramacal páramo generic flora to other páramo floras
of the northern Andes and Central America were assessed using ordination
(Detrended Correspondance Analysis – DCA, Principal Component Analysis PCA) and classification (Cluster analysis) methods for seven additional available
different páramo flora datasets. Floristic lists from each páramo site were obtained
from literature or unpublished data from authors (see Table 5.1).
The accounts on the different páramo floras were carefully screened by the authors
for taxonomic update and true forest taxa were deleted. Two dataset were
considered for these analyses, A) one which included the Guaramacal list of total
genera of 150 from páramo & SARF combined, and B) the other that includes
Guaramacal list of 108 genera from open páramo only. For these analyses, both
data matrices A (404 genera x 8 sites) and B (347 genera x 8 sites) of
presence/absence of genera in the eight páramo floras were analyzed using
program PC-Ord 4 (McCune & Mefford 1999). Cluster analyses of shared genera
used Sørensen (Bray-Curtis) as distance measure method and Group Average as
group linkage method.
Table 5.1. Reference information for the eight páramo flora dataset used for comparative
multivariate analysis.
PARAMO
Max.
Aprox.
Area
Number of Source of floristic
Elev.
Prec.
(ha)
genera
data
(m)
(mm/year)
considered
Sierra Nevada del 5330
1300 112,418
213
Cleef, unpubl. data
Cocuy, Colombia
ca.3000
Sierra Nevada de
4980
813 69100
149
Ricardi et al. 1997,
Mérida,
1811
Berg & Suchi, 2001
Venezuela
Sumapaz,
4250
~1200102,945
211
Cleef 1979, Franco
Cordillera
3000
& Betancur 1999,
Oriental,
Pedraza-Peñaloza et
Colombia
al. 2004, RangelCh. 2000c
Tatamá massif,
4100
>3000
5,000
114
Cleef et al. 2005,
Cordillera
Cleef 2005
Occidental,
Colombia
Serranía de
4100
~2000
4,560
137
Rivera-Díaz 2007
Perijá, Colombia
Talamancas,
3850
200015,205
177
Barrington 2005,
Costa
4000
Vargas & Sánchez
Rica/Panamá
2005
South Ecuador:
3695
~5000
14,169
201
Lozano et al. 2009,
Podocarpus
mm
Bussmann 2002,
National Park
Keating 1999
(PNP)
Guaramacal,
3130
>3200
~400
150/108
This study
Venezuela
mm
147
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
5.4 RESULTS
Flora characteristics
To date, the vascular flora of summit areas of Ramal de Guaramacal is composed
of a total of 251 taxa; 17 families, 28 genera, and 65 species of ferns, and 52
families, 124 genera and 186 species of angiosperms. In general, the most species
rich families are Asteraceae, Poaceae, Ericaceae and Orchidaceae, followed by the
ferns families Grammitidaceae and Lycopodiacae.
The most diverse genera are the ferns Elaphoglossum, Huperzia and
Hymenophyllum. Of the total 251 taxa, only 169 species belonging to 108 genera
have been registered for proper subpáramo-páramo vegetation, excluding the
SARF vegetation (Table 5.2).
Geographical composition of genera
The composition of genera of phytogeographic elements in páramo areas of Ramal
de Guaramacal is presented in Table 5.3. A total of 150 genera is contained in
Table 5.3, including 41 genera of woody, herbaceous and epiphytic plant species
found inside the forest islands (of SARF vegetation) surrounded by páramo
vegetation, and 27 genera present in azonal páramo vegetation. Exotic weedy
genera such as Polypogon, Rumex and Sonchus among others, present in disturbed
areas, are excluded. Proportions of phytogeographic elements and components of
the studied data set are shown in Figure 5.2 as well as in Table 5.5.
Table 5.2. Most diverse families and genera from the vascular flora of summit areas
(including SARF) of Ramal de Guaramacal, Andes, Venezuela. For only proper
páramo flora numbers of taxa are indicated in parenthesis.
Num
Num
Num
FAMILIA
Gen
spp.
Genus
spp.
ASTERACEAE
14(10)
24 (17)
Elaphoglossum
10 (5)
POACEAE
10
21(20)
Huperzia
8 (6)
ERICACEAE
10(8)
15(13)
Hymenophyllum
7 (2)
ORCHIDACEAE
9(4)
14(7)
Chusquea
7 (6)
GRAMMITIDACEAE
6(3)
13(6)
Rhynchospora
6
LYCOPODIACEAE
3
12(10)
Gaultheria
5
CYPERACEAE
4
10
Hypericum
4
DRYOPTERIDACEAE
2(1)
11(5)
Blechnum
4 (2)
RUBIACEAE
6(5)
7(6)
Melpomene
4
HYMENOPHYLLACEAE
1
7(2)
Miconia
4 (1)
MELASTOMATACEAE
3
6(3)
Pentacalia
4 (2)
BROMELIACEAE
4
5
Ruilopezia
4
MYRSINACEAE
3(2)
5(3)
Weinmannia
4 (0)
CLUSIACEAE
1
4
ROSACEAE
3
4
BLECHNACEAE
1
4(2)
CUNONIACEAE
1(0)
4(0)
Totals 69 (53) families
148
150
(108)
genera
251
(169)
species
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
Table 5.3. Composition of genera of phytogeographic elements in páramo areas of Ramal de
Guaramacal in the Venezuelan Andes. Asterisk* represents genera recorded from
SARF vegetation.
Element
Tropical
(TRO)
Páramo endemics (P)
Andean alpine
(NT-AA)
Neotropical montane
(NT-M)
Wide tropical
(WTR)
Genus
Libanothamnus Ernst, Paragynoxys* (Cuatrec.) Cuatrec., Ruilopezia Cuatrec.
Lachemilla (Focke) Rydb.
Ageratina Spach, Arcytophyllum Willd. ex Schult. & Schult. f., Aulonemia
Goudot, Baccharis* (Less.) DC., Bejaria Mutis ex L., Bomarea Mirb.,
Brachionidium* Lindl., Campyloneurum C. Presl., Cavendishia Lindl.,
Centropogon C. Presl., Ceradenia L. E. Bishop, Cestrum* L., Chusquea
Kunth, Cochlidium* Kaulf., Corynaea* Hook. f., Cranichis* Sw., Cybianthus
Mart., Dendrophtora Eichler, Deprea Raf., Diplostephium Kunth, Disterigma
Sleumer, Elleanthus C. Presl., Epidendrum L., Eriosorus* Fée, Excremis
Willd., Freziera* Willd., Gaiadendron* G. Don, Gamochaeta Wedd.,
Geissanthus* Hook. f., Glossoloma* Hanst., Gomphichis* Lindl., Greigia
Regel, Guzmania Ruíz & Pavón, Hesperomeles Lindl., Huperzia Bernh.,
Isidrogalvia Ruíz & Pavón, Jamesonia Hook. & Grev., Lellingeria* A.R. Sm.
& R.C. Moran, Macrocarpea* (Griseb.) Gilg, Manettia Mutis ex L., Miconia
Ruíz & Pavón, Monnina Ruíz & Pavón, Monochaetum (DC.) Naud.,
Munnozia Ruíz & Pavón, Myrcianthes* O. Berg, Odontoglossum Kunth,
Oreopanax* Decne. & Planch., Pachyphyllum* Kunth, Paepalanthus Kunth,
Palicourea Aubl., Pentacalia Cass., Phoradendron* Nutt., Pleurothallis* R.
Br., Psammisia* Klotzsch, Pterichis Lindl., Puya Molina, Siphocampylus
Pohl, Sphyrospermum Poepp. & Endl., Terpsichore* A.R. Sm.,
Themistoclesia Klotzsch, Thibaudia* Ruíz & Pavón, Tillandsia L.,
Tropaeolum L., Ugni Turcz.
Achyrocline (Less.) DC., Begonia* L., Chaetolepis (DC.) Miq., Clethra* L.,
Culcita* C. Presl., Cyathea* Sm., Elaphoglossum Schott ex J. Sm.,
Grammitis Sw., Hedyosmum* Sw., Histiopteris (J. Agardh) J. Sm.,
Hymenophyllum Sm., Ilex L., Melpomene A.R. Sm. & R.C. Moran, Mikania*
Willd., Myrsine L., Paesia J. St.-Hil., Peperomia* Ruíz & Pavón, Phytolacca
L., Pilea* Lindl., Plagiogyria* (Kunze) Mett., Psychotria* L., Sticherus C.
Presl., Symplocos* Jacq., Xyris L.,
Temperate
Austral-Antarctic
(AA)
Calceolaria L., Cortaderia Stapf., Cotula L., Drimys* J.R. Forst. & G. Forst.,
Fuchsia* L., Gaultheria L., Hypoxis L., Muehlenbeckia Meisn., Nertera
Banks ex Gaertn., Oreobolus R. Br., Ortachne Nees ex Steud, Orthrosanthus
Sweet, Pernettya Gaudich., Sisyrhynchium L., Weinmannia* L.
Holarctic
(HO)
Castilleja Mutis ex L. f., Diplazium* Sw., Gentianella Moench, Halenia
Borkh, Sibthorpia* L., Vaccinium L.
Wide temperate
Agrostis L., Arenaria L., Calamagrostis Adans., Carex L., Danthonia DC.,
Daucus L., Epilobium L., Festuca L., Galium L., Geranium L., Hieracium L.,
Hypericum L., Isoëtes L., Juncus L., Luzula DC., Plantago L., Poa L.,
Polypogon Desf., Stellaria* L., Valeriana L., Viola L.
(WTE)
Cosmopolitan
(CO)
Asplenium* L., Blechnum L., Cynoglossum L., Eleocharis R. Br., Equisetum
L., Gnaphalium L., Hydrocotyle L., Lycopodiella Holub., Lycopodium L.,
Ophioglossum L., Oxalis L., Polypodium L., Rhynchospora Vahl, Rubus L.,
Solanum L., Thelypteris Schmidel, Utricularia L.
149
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
1. Tropical component
On the basis of 150 vascular plant genera more than half 61.3% (92 genera) are
tropical. Neotropical montane element genera are those that range from montane
forest into the supraforest zone. This element is represented by 64 genera (42.7%).
Twenty one of them (including 10 herbaceous genera) correspond to SARF
vegetation (Table 5.3). When considering only the genera recorded from páramo
vegetation, the Neotropical montane element is represented by forty two genera
(38.9%), four of them are found in azonal páramo (Fig. 5.2).
Wide tropical element genera are widely distributed in the tropics, including those
exclusively African-American and Asian-American. This element is represented
by 24 genera (16%). Ten of them (including five herbaceous genera) were found in
SARF islands (Table 5.3). When considering only páramo vegetation genera, the
wide tropical element accounts for twelve genera (11.1%) and only one of them
(Xyris) is found in azonal páramo.
Páramo endemic element genera are those confined to páramo (and sometimes also
in the downslope Andean forests) and represented in the study area by 3 genera
(2%), two of them small trees: Libanothamnus at the UFL and Paragynoxys, a
species from SARF. Most spectacular are the 4 species of Ruilopezia
(Espeletiinae), endemic for Venezuela. Only one Páramo endemic genus
(Ruilopezia) is found in azonal páramo.
The Andean alpine element is represented by only one herbaceous genus (0.7%):
Lachemilla, which is found mainly in azonal páramo.
2. Temperate component
Forty two genera are of temperate distribution (28%), including six genera from
SARF. When considering only páramo vegetation genera, the temperate
component is represented by 36 genera or 33.3%. These include 31 herbaceous
genera, 16 of them counted from azonal páramo.
Widespread temperate element genera are distributed in temperate and cool
regions from both hemispheres. This element is represented in the study area by
twenty one genera (14%). The genus Stellaria was recorded from borders of SARF
vegetation. When excluding this genus, the wide temperate element is represented
by 18.5% for twenty páramo genera, eight of them counted from azonal páramo.
Austral-Antarctic element genera have southern temperate distribution. This
element is represented by fifteen genera (10%). Among them, three genera were
registered from SARF (Table 5.3). Twelve Austral-Antarctic element genera
(including 8 herbaceous) of only páramo vegetation account for 11.1%. Eight
genera are counted from azonal páramo.
Holarctic element genera have northern temperate including Mediterranean climate
distribution. Only six genera with Holarctic distribution (4%) were found in the
study area. The genus Sibthorpia, which corresponds to a small herb species and
the fern Diplazium have been found in borders of SARF vegetation or in the upper
forest line. Excluding the SARF genera, the Holarctic element is represented by
150
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
four genera (three of them herbaceous) or 3.7%. Gentianella was the only
Holarctic genus counted from azonal páramo.
(a)
100
100
Phytogeographic elements
Páramo & SARF
%
%
10.7
28.0
42.7
50
50
16.0
2.0
0.7
P
NT-AA
10.0
14.0
4.0
10.7
61.3
CO
TEMP
TROP
0
NT-M
WTR
AA
HO
WTE
CO
0
N=150
(b)
Phytogeographic
components
100
100
%
Phytogeographic elements
Páramo (zonal & azonal)
%
50
33.3
50
38.9
1.9
0.9
P
NT-AA
13.9
11.1
11.1
WTR
AA
18.5
13.9
52.8
3.7
CO
TEMP
TROP
0
NT-M
HO
WTE
CO
0
N=108
(c)
Phytogeographic
components
100
100
%
Phytogeographic elements
Azonal páramo
%
50
50
29.6
3.7
P
NT - AA NT - M
CO
TEMP
TROP
14.8
3.7
3.7
25.9
0
N=27
59.3
25.9
14.8
3.7
14.8
WTR
AA
HO
WTE
CO
0
Phytogeographic
components
Figure 5.2. Proportions (%) of phytogeographic components and elements of (a) genera of
páramo and SARF, (b) of all páramo genera, and (c) the genera from azonal
communities from Ramal de Guaramacal, Andes, Venezuela.
151
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
3. Cosmopolitan component
Cosmopolitan element genera are those with worldwide, or nearly so, distribution.
The Cosmopolitan element is represented in the study area by sixteen genera
(10.7%). The fern genus Asplenium, represented by the species A. serra, is found
in the understory of SARF vegetation. The Cosmopolitan component for only
fifteen páramo genera (13 of them herbaceous) is represented by 13.9%. In the
azonal páramo the Cosmopolitan component is represented by four genera.
Table 5.4. Analysis of the geographic range of the páramo flora based on 224 taxa with a
defined geographical range in Appendix 5. F: Ferns and fern allies, A:
Angiosperms; %: percentage of total vascular species. Numbers in parentheses
are percentages of total Venezuelan endemics.
Number of páramo species
Group
Description
1
2
3
4
Number of páramo & SARF
species combined
F
A
Total %
F
A
Total
%
Widespread in the Neotropics and
also occurring elsewhere
Widespread in the Neotropics
3
9
12
7.8
4
9
13
5.8
4
10
14
9.2
9
13
22
9.8
Widespread in Tropical South
America
Widespread in Central America,
northern (western) South America
and the West Indies
0
3
3
2.0
0
3
3
1.3
11
3
14
9.2
12
4
16
7.1
5
Central America, northern and
western South America, including
the Guyana highlands
2
3
5
3.3
3
5
8
3.6
6
Widespread from Costa Rica to
Bolivia
Widespread in the Andes from Col
to Bolivia
Confined to Venezuela, Colombia
and Ecuador
Confined to Venezuela and
Colombia
4
24
28
18.3
14
31
45
20.1
5
16
21
13.7
12
25
37
16.5
1
9
10
6.5
1
12
13
5.8
3
13
16
10.5
3
17
22
9.8
0
2
2
0
4
4
1
2
3
3
2
0
15
15
0
24
0
10
10
0
12
Total Venezuelan endemics
1
29
30
3
32
Species Totals
34
119
153
1.3
(6.7)
2.0
(10)
9.8
(50)
6.5
(33.3)
19.6
(100)
100
61
163
7
8
9
10. Endemic to Venezuela:
10.1
Andean region and Coastal
cordillera
10.2
Andean region and Venezuelan
Guayana (highlands)
10.3
Endemic to Andean region of
Venezuela
10.4
Endemic to Guaramacal
152
1.8
(8.9)
5
2.2
(11.1)
24 10.7
(53.3)
12
5.4
(26.7)
45 20.1
(100)
224 100.0
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
Species geographical range
The geographical range of the vascular species present in páramo areas of Ramal
de Guaramacal, grouped in ten major groups (or distribution types), is shown in
Table 5.4. Neotropical widespread distributed species all over in the whole
Neotropics or in a wide range from Central America to Bolivia are broken down
into five (1-5) groups. Andean distributed species are split into groups 6 to 9.
Venezuelan endemic species (group 10) is divided into four subgroups. The
number of vascular species, by taxonomic groups (ferns and Angiosperms) and
percentages of the total are presented for each distribution category. From the total
224 taxa determined to species, only 153 species belong to proper
páramo/subpáramo vegetation.
Páramo flora relationship
Figure 5.3 shows the dendrograms of generic similarity among páramo sites
resulting from the cluster analyses. In both graphs, over fifty percent of similarity,
four main groups can be recognized. The closest relationships (about 90%) among
páramos is observed between the generic páramo floras of the Colombian
Cordillera Oriental of each Sumapaz and Sierra Nevada del Cocuy, which are both
closely related to Sierra Nevada de Mérida in Venezuela. The generic páramo flora
of Ramal de Guaramacal shows the closest relationship to southern Ecuador
páramo flora of Podocarpus National Park, with more than 50% similarity, when
considering Guaramacal generic flora from páramo and SARF combined (Fig.
5.3a), however no relationship of Guaramacal to any other páramo flora is
observed when taking into account only the open generic páramo flora of
Guaramacal.
Figure 5.4 shows the resulting DCA (a, c) and PCA (b, d) ordination diagrams for
both A (a, b) and B (c, d) datasets of presence/absence of genera and 8 páramo
floras analyzed. An altitudinal gradient may be represented on first axis of DCA
(a) and second axis of PCA (d), while a humidity gradient is mainly captured by
second axis of PCA (b).
The results of ordination also show that for dataset A (that includes the páramo and
SARF genera from Guaramacal) páramos with greatest values of humidity and
rainfall according to Table 5.1 are grouped in line to the lower right corner on both
DCA(a) and PCA(b) diagrams (e.g. Tatamá massif, 4100 m, ~2000-3000 mm/year
(Cleef et al. 2005); South Ecuador, PBR, 3695 m, ~5000 mm/year (Lozano et al.
2009); and Guaramacal, 3100 m, > 3200 mm/year and relative humidity of 100%
during most part of the year), while drier and higher elevation páramos are
grouped to the lower left corner of DCA(a) and upper left corner of PCA(b).
However, that humidity relationship is not obvious for dataset B (that with
Guaramacal only open páramo genera), where páramo sites seem to be arranged
mainly in relation to an altitudinal gradient in axis 2 of PCA(d).
Compared to other generic páramo floras (Table 5.5), Guaramacal shows the
greatest proportion of Neotropical montane element genera and the lowest
proportion of Andean-Alpine element genera. The proportion of the Holarctic
153
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
element is the lowest of all páramo floras compared, but the Cosmopolitan element
is the highest.
a)
b)
Figure 5.3. Sørensen (Bray-Curtis) cluster analysis dendrogram of floristic similarity among
8 páramo sites based on (a) the presence/absence of 404 genera (including
páramo & SARF genera from Guaramacal), (b) the presence/absence of only
347 genera (including only proper páramo genera from Guaramacal).
Table 5.5. Proportions (%) of phytogeographic elements of páramo genera for seven
additional páramo floras compared to Guaramacal. (a) SARF and páramo
genera combined, (b) páramo genera only.
Phytogeographic
element
P
NT-AA
NT-M
WTR
AA
HO
WTE
CO
Total %
Total
genera
154
Guaramacal
South
Ecuador
Perijá
S. N.
Cocuy
S. N.
Mérida
Sumapaz
Tatamá
Costa
Rica
(a)
2.0
0.7
42.7
16
10
4.0
14.0
10.7
100
(b)
2.8
0.9
37.6
11.0
12.8
3.7
20.2
11.0
100
4
5.5
32.5
12
13
10.5
13.5
9
100
5.8
3.6
27.0
12.4
10.2
13.9
20.4
6.6
100
6.5
8.4
27.6
7.9
10.7
12.1
18.2
8.4
100
5.4
8.1
22.3
8.1
10.8
14.2
23.0
8.1
100
4.8
7.1
25.7
8.6
12.4
11.9
18.6
11.0
100
1.8
8.0
25.7
9.7
14.2
8.8
22.1
9.7
100
1.7
3.4
27.7
8.5
12.4
16.4
19.2
10.7
100
150
108
200
137
214
148
210
113
177
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
Páramos of Colombian Cordillera Oriental (S.N. Cocuy and Sumapáz) and Sierra
Nevada de Mérida show the most similar proportions of phytogeographic elements
among them. Páramos of Costa Rica/Panama and Tatamá show the lowest
proportion of Páramo endemic genera. Páramos of South Ecuador and Guaramacal
show both more similar (the highest) proportions of Neotropical genera and also
the lowest proportions of Holarctic genera.
a)
b)
c)
d)
Figure 5.4. DCA (a, c) and PCA (b,d) Ordination diagrams of 404 (a, b: including páramo &
SARF genera from Guaramacal) and 347 (c, d: including only proper páramo
genera from Guaramacal) genera for 8 páramo floras datasets. a) DCA Axis 1
Eig=0.422; Axis 2 Eig=0.321; (b) PCA Axis 1 Eig=473.827; Axis 2
Eig=121.424; (c) DCA Axis 1 Eig=0.223; Axis 2 Eig=0.189; (d) PCA Axis 1
Eig=803.377; Axis 2 Eig= 87.539.
155
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
5.5 DISCUSSION
Floristic features
As in almost all páramo and other alpine floras (Rangel-Ch. 2000c; Vargas &
Sánchez 2005; Rivera-Diaz 2007; Rangel-Ch. et al. 2008; Briceño & Morillo 2002,
2006; Lozano et al. 2009), Asteraceae and Poaceae rank as most dominant in terms
of genera and species (Table 5.2). Remarkable for the páramo of the study area is
the third position of Ericaceae with 10 (8) genera and 15 (13) species. Orchids and
Grammitidaceae take the 4th and 5th position respectively in the general flora list,
but for proper páramo flora only, Lycopodiaceae is more diverse. The relative
importance of Pteridophytes under wet climate is also supported by Dryopteridaceae with Elaphoglossum displaying 10 (5) species and Hymenophyllaceae
with Hymenophyllum containing 7 (2) species.
In terms of number of species (Table 5.2) Elaphoglossum, Huperzia and
Hymenophyllum and Chusquea take the first four positions in the general flora list.
For páramo flora only, Chusquea, Huperzia and Rhynchospora with six species are
the most diverse genera. The high diversity of Rhynchospora is remarkable.
Rhynchospora sect. Paniculatae is supposed to be derived from lowland savanna
stock (Wayt Thomas, pers. comm.). Earlier it was supposed that the ascent to the
Andean páramos from savanna flora was most likely from the lower ranges of the
eastern extreme/end of the Andes of Venezuela (Cleef et al. 1993). Rhynchospora
oreoboloidea Gómez-Laur. of the Holarctic sect. Oreoboloides, a common species
of the lower páramos in the northern Andes and in the Talamancas, is absent in the
Guaramacal páramo. In Colombian páramos hardly there are found 6 different
species of Rhynchospora in one study site.
Chusquea is considered here including three species formerly belonged to
Neurolepis (Fisher et al. 2009). One páramo species, Chusquea steyermarkii, has
vicariant bamboo communities on the tepuies.
In conclusion, the taxa listed in Table 5.2 are almost all indicative of wet páramo
climate. Hypericum and Pentacalia contain species thriving both under wet and
drier páramo climate.
Phytogeographical composition at genus level
Based on the studies of the Tatamá páramo flora (Cleef 2005) or that of the
Talamancas in Costa Rica (Cleef & Chaverri 1992) we expected that humidity
would play a role in determining the floristic composition of the Guaramacal
range. In fact values for the Neotropical montane element (38.9%) are high in the
Guaramacal páramo, as well as for the Austral-Antarctic element (11.1%).
Increased values for the Austral-Antarctic element also have been observed in the
Podocarpus National Park, Tatamá and Talamanca páramos. However the
substantial proportion of the Neotropical montane element may also be related to
the low altitude of the Guaramacal range, 3000 m more or less, and one summit at
3130 m. Páramo endemic genera rank low (2%), probably also because of the
general low altitude and one predominant humid climate type. There are also fewer
156
Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
distinct habitats in the Guaramacal páramo, as caused by the limited altitudinal
amplitude of maximally about 200 m, but most of the range even less. It is striking
that the Andean-alpine element is represented only by one genus (Lachemilla) and
that the Holarctic element only accounts for 3.6%. Genera belonging to both these
elements are mostly herbaceous and favoured by higher altitude. Further they are
well adapted to periodical stress by dryness (Gutte 1992). We suppose that
bamboo páramo has been present in the summit area of Ramal de Guaramacal
since Holocene times and that the prevailing wet climate served as a kind of filter
preventing the arrival or survival of dry páramo species from the Mérida páramos.
Another interesting feature is the relative isolation of the Guaramacal páramo from
the main cordillera of the Sierra Nevada de Mérida. A small connection is found
on the northern side at about 2200 m. During glacial times the summit areas of
Guaramacal range were glaciated; remnants of former glacial lakes with terminal
moraines are still present at different sites in the páramo belt as well as at lower
altitude of about 2000 m near the Park headquarters. Páramo vegetation actually
occurred during glacial times at lower altitude along the very steep slopes. In the
uppermost part of Guaramacal range with a type of superpáramo, which is
completely absent today. Isoëtes karstenii, a submerged species found from grass
páramo up to the highest lakes in the superpáramo in Colombia (Cleef 1981,
Salamanca et al. 2003) and Venezuela (Fuchs-Eckert 1982; Small & Hickey 2001)
has been found in a small lake in the Guaramacal páramo. Its presence in a glacial
lake in the modern páramo of Ramal de Guaramacal can probably be considered as
a ‘glacial relict’. The Temperate component is best represented in azonal páramo
vegetation (Sphagnum bogs) on top of Ramal de Guaramacal (Fig. 5.2c).
When the genera of the SARF vegetation in the Guaramacal bamboo páramo are
taken into account the overall proportion of the Tropical component rises from
53.6% to 61.6% , mainly because of more Neotropical montane and Wide tropical
genera. For comparison with other páramo floras (Table 5.5), the taxa from SARF
vegetation (column a) have not to be considered, though, sometimes this is
difficult to do as well. Looking at the case of the extremely humid páramos of
Podocarpus National Park in southern Ecuador (Lozano et al. 2009), with a gradual
transition of SARF into shrub páramo, it is noticeable that even the trees adapt to
the general structure of shrub páramo vegetation (Bussmann 2002; Richter &
Moreira-Muñoz 2005; Peters 2009, Lozano et al. 2009; Cleef pers. obs.).
Species geographic range
The tropical American part of the vascular flora of Páramo de Guaramacal is
largely composed of (1) Neotropical widespread distributed species all over the
Neotropics or in a wide range from Central America to Bolivia, (2) a group of
Andean distributed species, part of them confined to the northern Andes and part
widespread in the Andes from Colombia to Bolivia, and (3) a group of Venezuelan
endemics (Table 5.4).
There is quite a difference between the 153 species with defined geographical
distribution range reported for the Guaramacal páramo and the 224 species for the
páramo including the SARF islands of Guaramacal. However, the phytogeo157
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
graphical proportions change slightly between both data bases: they maintain
rather the same percentages. Looking more closely at the three main distribution
types of the Guaramacal páramo flora (sensu strictu, without the SARF islands) we
can state that there are 48 species, i.e. ca. 31%, for the groups or distribution types
1-5 (Table 5.4), these species displaying a more wide Neotropical distribution. The
second and largest species group includes the distribution types 6-9 and is
basically tropical Andean in distribution and accounts for seventy five species or
about 49%. Group 10 contains with thirty species (almost 20%) endemic to
Venezuela. Ten species (6.5%) are narrow endemics of the Guaramacal páramo.
They include 3 species of Espeletiinae stem rosettes: two species of Ruilopezia,
and one species of Libanothamnus. Also, two species of Miconia, one species each
of Bomarea, Epidendrum, Festuca, Ilex and Rhynchospora.
About 69 species or about 30% of the Guaramacal páramo species are shared with
Central America – surprising given the distance and remoteness of Ramal de
Guaramacal, although 29 of them correspond to ferns (Table 5.4). In contrast, only
3 species (2%) are shared with the Guayana Highlands which are at much closer
distance indicating lack of exchange between these two areas. Most remarkable is
the northernmost extension of the bamboo species Chusquea steyermarkii.
Páramo flora relations
We found a strong floristic similarity and similar phytogeographical composition
among the páramo floras of Sierra Nevada del Cocuy, Sumapáz and Mérida
páramos (Fig. 5.3, 5.4, Table 5.5). These mountain chains are contiguous in
geographical position and display similar climatic characteristics with regard to the
exposition of the ascending trade winds loaded with atmospheric water and the
drier wind shadow areas. The Central American páramos of Panamá and Costa
Rica, which are more humid, present about 75% similarity of páramo flora with
those of the Mérida and Colombian Eastern Cordillera páramos (Fig. 5.3a). The
Colombian Perijá páramo (drier side) ranks with about 40% similarity versus the
wet páramo cluster of Guaramacal and PNP in S. Ecuador. Both remote páramo
floras are similar at about a 60% value, which is most remarkable, because of the
large distance between both areas. The similarity between the páramo floras of
Guaramacal and PNP of South Ecuador is observed only when considered the
páramo and SARF genera of Guaramacal (Fig.5.3a). When considered only open
páramo genera of Guaramacal (Fig 5.3b), the páramo flora of Guaramacal is not
related to any other of the paramo floras analyzed, and in this case PNP (South
Ecuador) flora appears to be rather related with the group formed by the páramo of
Perijá and the group of drier and higher paramos of S. Cocuy, Sumapáz and S.N.
Mérida, conversely, in this case, the páramo flora of Costa Rica/Panama has little
relationship with this group. On the other hand, in the DCA and PCA ordinations,
when SARF genera of Guaramacal are not included (Fig 4c, d), the relationship to
a humidity gradient is not so obvious, and an altitudinal gradient seem to prevail in
PCA (Fig. 5.4d), while in the DCA (Fig. 5.4c) the relationship to those
environmental variables is not so clear, and instead of them a latitudinal gradient
may be detected.
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Phytogeography of the vascular páramo flora of Ramal de Guaramacal
_______________________________________________________
Judging from the results it is most clear that the wet páramos floras are more
similar to each other than to seasonally dry páramos (containing both dry
bunchgrass páramo and bamboo páramo). In the case of the exclusively wet
páramos it appears that humidity is more important than a temperature gradient. In
fact the Ecuadorian Podocarpus National Park and Guaramacal páramos are
similar in that both are relatively low in altitude with a maximum of about 200 m
altitudinal amplitude in Guaramacal and about 400-500 m in the Podocarpus
National Park although the highest core area of the latter reaches ~3700 m in
elevation. That the ambient humidity gradient apparently overrules that of
temperature (viz. altitude), seems also confirmed by the DCA en PCA ordination
diagrams of Fig. 5.4(a,b), which are based on a comparison of eight páramo floras.
159
Venezuela endemic species of the Espeletiinae, found in Páramo de Guaramacal: (a-c) Ruilopezia
jabonensis; (d, e) Ruilopezia lopez-palacii; (f-h) Ruilopezia paltonioides; (i-l) Ruilopezia viridis.
Chapter 6
Functional diversity of Andean forests in Venezuela changes
with altitude
Joost F. Duivenvoorden and Nidia L. Cuello A.
submitted to GLOBAL ECOLOGY AND BIOGEOGRAPHY
Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
6.1 INTRODUCTION
Tropical Andean forests are one of the world's biodiversity hot-spots (Myers et al.
2000). These forests have a rich biodiversity (Gentry 1995) and are highly
threatened because of increasing deforestation (Etter & Wijngaarden 2000;
Armenteras et al. 2003). The rising temperature in the past decades and associated
upslope shifts in species distribution may impose extra threats to Andean forests
(Colwell et al. 2008; Svenning & Condit 2008). Potential threats due to losses in
forest cover and biotic attrition might be exacerbated by degradation in functional
diversity, i.e. the variety of life-history traits presented by an assemblage of
organisms (Mayfield et al. 2005; Girao et al. 2007). Decreasing functional
diversity is generally seen as indication of degradation and a hazard for ecosystem
resilience (Tilman et al. 1997). For example, the lower diversity in reproductive
traits in forest fragments in lowland Amazonia of Brazil may have detrimental
consequences for the population size of pollinators and the trophic structure (Girao
et al. 2007). The principal aim of our study is to examine if functional diversity
changes with altitude in undisturbed Andean forests, to contribute as reference
information for studies of degraded Andean systems.
Along mountain slopes temperature change strongly defines the rate of
photosynthesis (Rada et al. 1992; Cabrera et al. 1998), physiological and metabolic
processes (Lambers et al. 2008), growth (Grubb 1977; Medina & Klinge 1983;
Ashton 2003; Leuschner & Moser 2008), nutrient uptake (Bruijnzeel 1991; Gerold
2008; Leuschner & Moser 2008) and decomposition (Illig et al. 2008), and is
therefore the principal driver of ecosystem functioning (Chapin & Körner 1998;
Colwell et al. 2008; Svenning & Condit 2008). In general nutrient availability and
decomposition rates decrease at higher elevations in tropical wet montane forests
(Cavelier 1996). Above 1500 m, chances on occasional frost increase at higher
elevations. Yet, because of the strong insolation, the maximum daily temperature
remains quite similar to lowland values, resulting in a larger diurnal temperature
range upslope (Hansen et al. 2002). In upper montane and subalpine rain forest
(SARF), canopy trees receive a large proportion of ultraviolet light, which
potentially affects growth (Flenley 1992). Lastly, terrain conditions (more
summits) and the proximity of the upper forest line dictate that less space becomes
available for continuous forests at higher elevations, which makes fragmentation
by natural causes, in principal, more frequent. Most of these factors contribute to
stronger upslope levels of ecological filtering (Keddy 1992; Weiher & Keddy
1995; Ackerly 2003) acting upon montane forest plants, reducing the number of
traits relative to species (underdispersion). Alternatively, increased competition for
more limiting resources at higher altitudes (for example due to the lower
decomposition rates) might invoke ecological differentiation leading to higher trait
diversity relative to species diversity (overdispersion) (Weiher & Keddy 1995;
Mayfield et al. 2005).
Temperature-constrained processes likely become manifest in plants through
variation in response traits (Gitay & Noble 1997; Naeem & Wright 2003; Violle et
al. 2007) related to the energy balance (growth form, leaf shape and leaf size)
(Cornelissen et al. 2003). Fragmentation hampers dispersal and cross-pollination,
affecting the distribution of regenerative response traits like dispersal mode and
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Flora, vegetation and ecology in the Venezuelan Andes
pollination mode, fruit type, and flower and seed size (Cornelissen et al. 2003;
Girao et al. 2007). We studied the vascular plant composition of 44 small plots
located between 1330 m and 3060 m altitude in a well-protected forest reserve in
the Andes of Venezuela (Cuello & Cleef 2009a). We linked each species to the
above-mentioned functional traits by means of literature and herbarium studies.
Randomizing the species assemblages in our relevés (Legendre et al. 1997; Dray &
Legendre 2008) we tested if the composition and diversity of energy balance
related traits and fragmentation related traits changed with elevation.
6.2 METHODS
Study area
Ramal de Guaramacal is an outlier of the Venezuelan Andes, which lies to the
southeast of Boconó, Trujillo State, approximately 120 km northeast of Mérida, in
the centre of the Sierra Nevada de Mérida (9° 05–21' N and 70° 00–20' W). This
mountain range reaches up to about 3100 m, and most pertains to a National Park,
which includes an approximate surface area of 21,466 ha. The average yearly
rainfall measured over 2002-2008 at a climate station in the study area (Laguna de
los Cedros at 1980 m; 9° 15' 55'' N; 70° 13' 13'' S) was 2106 mm, and showed a
unimodal pattern with February as driest month and June as wettest. Temperature
average around 18 to 20° C between 1000 and 1500 m, and 9 to 12° C above 2500
m (Cuello & Barbera 1999). Above 2500 m seasonal frost may occur (Urriola
1999). The high precipitation in the area favors intense lixiviation and
acidification, and acid soils predominate (Marvez & Schargel 1999).
The vegetation of Guaramacal Park area is predominantly represented by montane
rain forests with height and density decreasing with altitude. These forests have
been described (Cuello & Cleef 2009a) into discrete zones corresponding to lower
montane, upper montane, and SARF, following Grubb (1977). The montane
forests can be found from 1350 m to about 2800 m. Between 2800 m and 3130 m
SARF is found intermingled in a mosaic with subpáramo formations (Cuello &
Cleef 2009b).
Ramal de Guaramacal has received the status of National Park since 1988, keeping
most human activities and impacts outside the park borders. Fires have occurred in
the past, especially in páramo areas. Some selective timber extraction is known to
occur at low intensity and generally takes place in close proximity to the park
limits. In the surroundings of Ramal de Guaramacal there has been a long history
of agricultural activity mainly for coffee plantation, slash and burn cultivation and
extensive cattle ranging, among other land uses (Barbera 1999). However, the high
ridges and steep slopes of Guaramacal have kept most of the montane forest areas
with minimum disturbance.
Field methods
The fieldwork was carried out in 1995, 1996, 1999, 2003, 2005 and 2006. Montane
forests were studied along the altitudinal gradient on both sides of the range with
164
Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
different slope expositions (Cuello & Cleef 2009a, b). Thirty five 0.1 ha (20 x 50
m) plots were surveyed, positioned a distances of 30 to 150 m between 1350 m and
2890 m altitude, and nine plots of variable size (50 m2 to 400 m2) were surveyed
in SARF between 2800-3050 m. Within each plot, all rooted individuals – trees,
shrubs, lianas, tall and thick-stem or climbing terrestrial herbs and hemi-epiphytes
≥ 2.5 cm DBH (diameter at breast height, taken at 1.3 m from the base of the
trunk, or lower for shrubs and thick-stemmed herbs) were recorded, labeled with
numbered aluminum tags and their DBH and height recorded.
A total of 2082 numbers botanical specimens of vascular plants were collected.
Botanical material was processed and identified at Herbario Universitario PORT of
UNELLEZ. Other herbaria, such as MO and US, were also consulted. Some
specimens were sent to specialists at other institutions to confirm identification.
All specimens collected have been deposited at PORT, some duplicates have been
sent to VEN, MER, MERF, MO and US.
From a total of 388 morphospecies recorded from all surveys, 357 were identified
to species or genus level. These we used to compiled trait state data on energy
balance and fragmentation related traits, on the basis of literature, floras and
botanical monographs, web searches, and herbarium voucher information (Table
6.1).
Table 6.1 Plant response traits and their respective categories or trait states considered in
this study.
Trait
Energy balance related traits
Growth form
Leaf type
Leaf size
Fragmentation related traits
Dispersal
Pollination
Sexual system
Fruit type
Fruit size
Flower size
Trait states or categories
bamboo (39), climbing herb (41), erect herb (45), hemiepiphytic tree (46), liana (47), palm (48), stem rosette (49), tree
(52), tree fern (53), upright shrub (55)
simple (56), dissected (57), compound (58)
leptophyll (<0.25 cm2)(59), nanophyll (0.25-2.25 cm2)(60),
microphyll (2.25-20.25 cm2)(61), notophyll (20.25-45 cm2)(62),
mesophyll (45-182.25 cm2)(63), macrophyll (182-1640.25
cm2)(64), megaphyll (>1640.25 cm2)(65)
autochory (1), anemochory (2), hydrochory (3), zoochory (4)
insect (5), bat (6), bird (7), self (8), water (9), wind (10), none
(11)
dichogamy (12), dioecious (13), monoecious (14), polygamous
(15), hermaphrodite (16), none (17)
achene (18), berry (19), capsule (20), drupe (21), fleshy
capsule/pome (22), follicle (23), legume (24), naked seed (25),
syncarp (26), none (27)
tiny (<2 mm2) (28), small (2–5 mm2)(29), medium (6–15 mm2
long)(30), large (16–25 mm long)(31), ex-large (36–100 mm
long)(32), huge (>100 mm long)(33)
Inconspicuous (<4 mm)(34), small (4-10 mm)(35), medium (1020 mm)(36), large (20-30 mm)(37), very large (>30 mm)(38)
165
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Flora, vegetation and ecology in the Venezuelan Andes
Data analysis
Detrended correspondence analysis (DCA) was done on the basis of the species-torelevee matrix, in which the species abundances were log-transformed, and on the
basis of the species-to-trait state matrix, in which all trait states were entered as
dummy variables. Variance partitioning was done using Canonical Correspondence Analysis (CCA; scaling focus of inter-species-distances and biplot scaling
type) after Borcard et al. (1992). In this, the environmental information was
represented by the elevation of the relevés in m above sea level. The spatial
variables were selected by means of a forward selection procedure (using a
probability level of 0.05) in CCA of the log species abundances against all terms
of the third-degree polynomial of the centered UTM X and Y coordinates
(Legendre & Legendre, 1998). The pure effects of the explanatory variables on the
species patterns were tested by means of Monte-Carlo permutation tests under
reduced model, applying 499 permutations. All DCA and CCA analyses were done
in CANOCO for Windows 4.5.
The relevé scores to visually show the variation in trait composition against
elevation were calculated as the weighted average of the species scores of the
DCA of the species-to-trait state matrix, with for each relevee the number of plants
per species as weight. Fourth-corner analysis was done applying 999 permutations
under models 1 and 3 (Legendre et al. 1997; Dray & Legendre 2008) with the
fourthcorner function implemented by S. Dray in the ade4 package (Dray &
Dufour 2007) in r 2.10.
Trait state diversity was quantified by the Shannon and the Simpson (1-D) indices.
Because of few aberrant relevee sizes, also Fisher's alpha was used to reduce
possible effects of variable sampling sizes. All indices were calculated (applying
Vegan 1.17-2 in r 2.10) on the basis of the number of species or the number of
individuals per trait state. Thus, in analogy to the calculations of species diversity,
trait states were used as equivalent of species and the numbers of species or the
numbers of individuals per trait state were used as equivalent of the number of
individuals (Girao et al. 2007). The Pearson coefficient of the correlation of the
diversity indices with elevation was tested by means of 999 permutations of the
species-to-relevee matrix according to the same permutation models and the same
number of permutations applied in the fourthcorner tests, after adding the reference
value (Pearson correlation coefficient from the unpermuted matrix) to the
distribution of the null model (Legendre et al. 1997).
6.3 RESULTS
Trait composition against elevation
In the 44 relevés a total of 357 species were recorded (see Appendix 6). Most
species (85%) were fully identified. The relevee scores of the first DCA axis of the
species-to-relevee matrix were highly correlated with altitude (Fig. 6.1A). The
gradient length of this axis was 9.1, indicating a substantial degree of species
turnover between the relevés (Hill & Gauch 1980). In space, the relevés were
clustered in about five groups, the largest of which consisted of the relevés made
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Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
most upslope (Fig. 6.1B). The forward selection procedure of the CCA of the logtransformed species information against the nine spatial variables produced five
significant terms (X, Y, X2, Y2, and X*Y). Of these X*Y was skipped from the
final analysis because of its high correlation with both X2 and Y2 terms (the
variable inflation factors of this final CCA were below 5). In the variance
partitioning, the pure elevation effect explained 6.2% of the species variation, and
the pure spatial effect 12.9%. Both these pure effects were significant (MonteCarlo permutation tests p=0.002). Elevation and space combined explained 2.8%
and the fraction of unexplained variation was 78%.
Figure 6.1 (left). Sources of variation in vascular plant species composition in the forests of
Ramal de Guaramacal in the Venezuelan Andes. A: Altitude: the association
between the first DCA axis of the species-to-relevé matrix and the altitude of
the relevés. B: Space: the spatial configuration of the relevés. Wider circles
were made at higher elevations.
Figure 6.2 (right). The principal variation in energy balance related traits (A) and
fragmentation related traits (B) extracted by means of the DCA of the speciesto-trait state matrix. Most trait states are simply abbreviated; leaf1-7, fr1-6, and
fl1-5 means leaf size, fruit size, and flower size in increasing order (compare
Table 6.1).
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Flora, vegetation and ecology in the Venezuelan Andes
In the DCA analysis of the species-to trait state matrix, the main variation (13.7%)
in energy balance related traits (Table 6.1) was clearly related to leaf size (Fig.
6.2A). Species with large leaves scored low scores along the first DCA axis and
fine-leaved species high scores. The variation between palms and tree ferns
(showing compound leaves), and herbs (either climbing or erect with dissected
leaves) mainly determined the variation along the second DCA axis. The
ordination of the fragmentation related traits was mainly driven by fruit and flower
size (associated with fleshy fruits, legumes, bat pollinated flowers, autochorous
dispersal and self-pollinated) with highest scores along DCA axes 1 and 2,
whereas small sized flowers and fruits (associated with dispersal and pollination
by wind, achenous fruits and fern reproduction) were situated towards the lower
left side of the ordination diagram (Fig. 6.2B).
Table 6.2 Association between elevation and the DCA axes of energy balance related traits
and fragmentation related traits, as given by squared Pearson correlation
coefficients (r) and their probabilities (p) obtained by means of the fourth-corner
analysis applying two permutation models.
DCA axis 1
model 1 model 3
Energy balance
related traits
Fragmentation
related traits
r
0.35
p
0.001
p
0.001
r
-0.11
-0.16
0.001
0.007
-0.25
DCA axis 2
model
model 3
1
P
p
0.064
0.028
0.001
0.001
Table 6.3. Association between elevation and three diversity indices of energy balance
related traits and fragmentation related traits, as given by Pearson correlation
coefficients (r) and their probabilities (p) obtained by applying two permutation
models.
Shannon
Simpson (1-D)
Fisher's alpha
model
1
mode
l3
model
1
model
3
r
p
p
r
p
p
0.001
0.41
0.001
0.001
0.74
0.001
0.001
0.001
0.022
-0.10
>0.2
>0.2
0.31
0.016
>0.2
-0.24
0.044
>0.2
-0.34
0.008
>0.2
0.34
0.012
0.001
-0.66
0.001
0.002
-0.62
0.001
0.003
-0.56
0.001
>0.2
r
Species-based trait
diversity
0.52
Energy balance
related traits
-0.58
Fragmentation
related traits
model
1
model
3
p
p
0.001
Individual-based trait
diversity
Energy balance
related traits
Fragmentation
related traits
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Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
Visually, elevation correlated well to the first DCA axis of energy balance related
traits (Fig. 6.3A) but less convincingly so to the second DCA axis. Both first and
second DCA axes of the fragmentation related traits seemed associated to
elevation (Fig. 6.3B). Fourth-corner analysis allows the selection of the most
appropriate null model to test for association between functional trait composition
of species in relevés and environmental properties of these relevés (Legendre et al.,
1997). Because of the significant effect of altitude on the species composition in
the relevés and the continuous species turnover along with altitude (Fig. 6.1A), it
seemed likely the individual ecological and physiological species responses to
elevation ruled the co-occurrences of species in the relevés. For this reason we
selected permutation model 1 (for each species separately the abundance values
are randomly distributed over the relevés; Legendre et al. 1997). However, the
variance partitioning showed that the spatial effect on species composition was
about twice the altitudinal effect. Spatial distance potentially hinders dispersal and
limits recruitment, both important in situations when regeneration through
colonization drives species composition. Because regeneration and colonization
after disturbances potentially depends on largely unpredictable processes related to
mass movements and tree mortality the lottery model of species assembly (Sale
1978; van der Maarel & Sykes 1993) seemed appropriate as well. Therefore, we
also applied a model 3 randomization (for each relevee separately the abundance
values are randomly distributed over the species; Legendre et al., 1997). The
fourthcorner results (Table 6.2) were in line with our visual interpretations of the
scatter plots (Fig. 6.3AB), evidencing that the distribution of energy balance and
fragmentation related traits was significantly correlated with altitude.
Functional diversity against elevation
All three species-based diversity indices in energy balance related traits showed a
convincing positive correlation with elevation (Fig.6.4A, Table 6.3). The
altitudinal association of the individual-based diversity of these traits was weaker
and less consistent (Fig. 6.4B, Table 6.3). The Shannon and Simpson indices
seemed negatively correlated but these patterns depended strongly on three SARF
relevés and lacked significance when tested with the lottery model of permutation.
The individual-based Fisher's alpha index of energy balance related traits was
positively related to elevation.
Regarding the diversity in fragmentation related traits, the overall tendency was
that of a negative association with elevation (Fig. 6.4A and B; Table 6.3).
However, compared to the energy balance related traits, the altitudinal correlations
were weaker and more strongly influenced by outlying diversities of SARF plots.
Positive SARF outliers (going against the trend of neutral or negative trends of
diversity with elevation) were clearly visible in the scatters of species-based
Simpson and Fisher's alpha indices (Fig.6.4A). For that reason the positive
altitudinal correlation of the species-based Fisher's alpha was not convincing, even
though it was significant in both permutation models. Negative outliers of SARF
against elevation appeared in the scatters of the individual-based Shannon and
Simpson against elevation (Fig. 6.4B).
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Flora, vegetation and ecology in the Venezuelan Andes
Figure 6.3. The association between elevation and the principal variation in trait
composition of the vascular plant species in the forests of Ramal de
Guaramacal in the Venezuelan Andes. Relevés scores for the trait
composition were the weighted average of the species scores of the DCA axes
derived from the species-to-trait state matrix, with for each relevé the number
of individuals as weight. A: energy balance related traits. B: fragmentation
related traits. Symbols of SARF relevés have been filled
6.4 DISCUSSION
Energy balance related traits
The forests in the Ramal de Guaramacal area varied altitudinally in the selected
energy balance related traits. Also they became more diverse in these traits at
higher elevations, pointing at more prominent levels of overdispersion higher up
the slopes. Community assemblage rule theory (Weiher & Keddy 1995; Díaz et al.
1998) predicts that increasing levels of overdispersion might occur when better
adapted species outcompete functionally related species from the local community.
Leaf size contributed substantially to the altitudinal variation in energy balance
related traits (Fig. 6.2A). The lower leaf size at higher altitudes in wet tropical
forests has been recorded repeatedly (Grubb et al. 1963; Vareschi 1966; Sugden
1985). In the absence of pronounced dryness (as in the situation along the slopes of
the Ramal de Guaramacal area) this can be explained by a lower upslope
temperature, more limited hydraulic conductance of stems and associated lower
mineral supplies, lower nutrient availability, and increased frost frequencies
(Cavelier 1996). Overall, at higher altitudes in montane wet forests, nitrogen
(Grubb 1977; Cavelier 1996) becomes more limiting. Therefore, our results
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Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
suggested that competition for resources, mostly related to the capture of radiation
(heat) and the uptake of minerals and nutrients, is an important driver of species
composition in this part of the Venezuelan Andes.
Figure 6.4A. Scatter plots of trait state diversity against elevation, calculated on the basis of
traits per species (left), or traits per individuals (right) for energy balance
related traits.
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Flora, vegetation and ecology in the Venezuelan Andes
Figure 6.4B: Scatter plots of trait state diversity against elevation, calculated on the basis of
traits per species (left), or traits per individuals (right) for fragmentation
related traits. Symbols of SARF relevés have been filled.
The association between functional diversity of energy related traits with elevation
further suggested that a temperature rise as a consequence Global Change might
affect the forest functionality of Andean forests. Projected higher temperatures in
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Functional diversity of Andean forests in Venezuela changes with altitude
_______________________________________________________
the coming decades (Solomon et al. 2007) might reduce the functional diversity of
energy balance related along the slopes of the Andes. This projection has only
general implications, however, because the detailed mechanisms, by which species
migrate as function of changing temperatures and ecological filtering influences
species interactions, are still poorly understood (Svenning & Condit 2008). For
example, vascular plants may migrate upslope at different speeds compared to
decomposer communities, which largely define the nutrient availability. Moreover,
our results suggests that even in undisturbed Andean forests functional diversity
varies significantly, which implies that some lower degree of functional diversity
at a certain elevation in the nearby future does not necessarily endangers the
ecosystem well-being.
Fragmentation related traits
Just as with the energy balance related traits, the forest species varied in our
selection of fragmentation related traits as function of altitude. The negative
altitudinal association of the main variation in fragmentation trait state
composition was mostly due to wind-dispersed and wind pollinated species from
SARF forests (e.g. species from Alsophila, Cyathea, Dicksonia, Diplazium,
Baccharis, Diplostephium, Pentacalia, Mikania). Because of the low human
influence in the Guaramacal area, the transition between SARF forests and páramo
vegetation is not sharp (for example compared to forest-páramo boundaries caused
by burning; Moscol & Cleef 2009). Instead, SARF and páramo vegetation occur in
a spatially well-mixed mosaic (Cuello & Cleef 2009a, b). Therefore, the
predominance of the trait states related to wind transportation in our highest
samples could be explained by the flow of plant propagules along forest-páramo
edges (Ries et al. 2004). The comparison of second-best variation in fragmentation
state composition with altitude (Fig. 6.3B) was due to the tendency that species
with larger fruits and flowers occurred at relatively low elevations (e.g.
Symbolanthus vasculosus (Griseb.) Gilg., Zygia bisingula L. Rico, Drymonia
crassa C.V. Morton, Tabebuia guayacan (Seem.) Hemsl., Macrocarpaea
bracteata Ewan, Inga edulis Mart.). We speculate that this pattern is related to a
more important role of birds in pollination and seed dispersal at elevations above
2100 m, versus a more pronounced role of mammals (including large bats) at
lower elevations.
Several SARF plots showed an outlying functional diversity compared to the
altitudinal trends in the lower forest relevés. This suggests that, in contrast to the
lower lying forests, the plants in the SARF relevés contained markedly more trait
states relative to the number of species, and/or more plant individuals relative to
the variety in trait states. Both phenomena may be caused by the increased wind
flow in SARF forests enhancing the number of plants with traits related to wind
transport.
Setting aside the outlying SARF patterns, and in contrast to the energy balance
related traits, the diversity of fragmentation related traits tended towards a negative
association with elevation, visible in both species-based and individual-based
indices of the montane forests. Hence, our results indicate that along a natural
altitudinal gradient in Andean rain forests, undisturbed by human influence,
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Flora, vegetation and ecology in the Venezuelan Andes
ecological differentiation regarding disturbance gets lower in upslope direction.
Possibly, the increased level of upslope underdispersion is due to a higher
fragmentation of the forest matrix because of stronger terrain dissections. In
comparison, Girao et al. (2007) reported that lowland forest fragments in Brazil
showed a lower functional diversity in reproductive traits compared to control
forests. Their findings pointed towards higher frequencies of self-incompatible
systems due to habitat loss. Altitudinal information on forest dynamics and forest
disturbance related to mass movements and slope instabilities in the Guaramacal
area is needed to further develop hypotheses about causal mechanisms explaining
the upward decrease in functional diversity of disturbance related traits.
Conclusions
According to our expectations, we found that functional diversity of undisturbed
Andean forests in the Guaramacal area changed with altitude. This implies that
temperature rise due to Global Change might affect the forest functionality of
Andean forests in the near future, but not necessarily in a harmful way. Functional
diversity related to energy balance traits increased in upslope direction, pointing at
increased levels of ecological differentiation. We explained this by assuming more
upslope competition in the Andean forests regarding capture of radiation and the
uptake of minerals and nutrients. Diversity in fragmentation related traits showed
an opposite pattern (more underdispersion upslope), which might relate to
discontinuities in the forest matrix due to the geomorphology of mountains. SARF
forests diverged from the altitudinal trends in fragmentation related traits, probably
as a consequence of edge effects in the SARF-páramo mosaic, created by wind.
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Synthesis
Nidia L. Cuello A.
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7.1 AIMS
The study of flora and vegetation of Ramal de Guaramacal, located in the Andes of
Venezuela, was conducted with the general aim to study montane rain forest and
páramo by exploring their structure, botanical composition and diversity and
relating these vegetation types to main environmental factors changing along
gradients. A specific goal was to examine the patterns of forest diversity along
altitudinal gradients with regard to plant functional traits. To elucidate the
phytogeographical patterns of the wet páramo flora, these vegetation types were
compared to other páramo areas in Ecuador, Colombia and the Talamancas of
Central America. This study contains basic knowledge for the conservation and
biodiversity management in the region.
This thesis provides information about forest and páramo vegetation along
altitudinal gradients using a floristic and phytosociological approach (chapters 2, 3
and 4); information about the analysis of the phytogeography of the Guaramacal
páramo flora, and relationships to floras of other páramos (chapter 5), and finally
information on montane forest diversity along an altitudinal gradient by means of a
plant functional approach (chapter 6). The results of the different chapters are
synthesized below.
7.2 ALTITUDINAL ZONATION
The TWINSPAN analysis of forest vegetation along an elevational gradient in
Ramal de Guaramacal showed an altitudinal zonation of forest types. Forest types
are grouped into discrete zones corresponding to the lower montane rain forest
(LMRF), upper montane rain forest (UMRF), and subalpine rain forests (SARF)
classes of Grubb (1977). Alternatively there is a correspondence to the subandean
forest, Andean forest, and high Andean forest, respectively, following Cuatrecasas
(1934, 1958). However, forest zonation was found variable between the northern
and southern slopes of Guaramacal.
LMRF of Ramal de Guaramacal can be found from 1350 m on the southern slope
and from 1650 m on the northern slope, to about 2300 m. However, in downslope
direction LMRF is limited to 1800 m, determined by the Park boundaries, in most
areas on the northern slope. Below 1800 m, disturbed areas occupy the potential
LMRF zone. UMRF is present from 2300 to ~2800 m on the northern slope of
Guaramacal. However, on southern or northwestern slopes near the tops of small
ranges, UMRF occurs as low as 2100 m. In Ramal de Guaramacal SARF is present
at the same altitudes as páramo vegetation, viz. from 2800 to 3050 m. In this
altitudinal range a zone with subpáramo vegetation, according to Cuatrecasas
(1934, 1958), is recognized. Subpáramo vegetation is classified into a lower
subpáramo or shrub páramo, and an upper subpáramo or dwarfshrub bunchgrass
páramo, following Cleef (1980, 1981). On the windward southern slopes, forest
zones of UMRF tend to reach lower elevations than on the opposite and drier
northern slopes and there the sequence of forest zones is shortest in distance.
Higher temperatures, almost permanent humidity, and frequent landslides on the
steeper and wetter southern slopes at mid-elevation may play a role.
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Flora, vegetation and ecology in the Venezuelan Andes
The surveys along the forest-páramo border in Ramal de Guaramacal detected an
upper forest line (UFL) position around 2950 m. This is remarkably low in
comparison with UFL positions elsewhere in the equatorial Andes and the Costa
Rican Talamancas. The low UFL is apparently caused by the “top effect” (Grubb
1971) with UMRF (including SARF) found at lower altitudes (Grubb 1977). A top
effect may also cause a compression of the montane forest zones. The history of
the low UFL in Guaramacal range may be explained by paleoecological studies of
lake sediments, e.g. the promising peat land at c. 2000 m near the Park Rangers
house of the Guaramacal National Park. Also an analysis of climatic data from the
Davis weather station near Antenas (data available from December 2006 onwards),
and analysis of soil surface temperatures on the top of Páramo de Guaramacal may
be helpful to explain the UFL position. Additional comparative analysis of the
vegetation ecology at the UFL would also provide more clues for understanding
the low altitude of the UFL on Ramal de Guaramacal.
7.3 FLORISTIC COMPOSITION AND DIVERSITY
Montane forests of Ramal de Guaramacal show different patterns of species
diversity, family composition and vegetation structure along altitudinal gradients
on the drier as well as on the wetter slopes. In LMRF the Rubiaceae, Lauraceae
and Melastomataceae are the most species rich families of woody plants, which is
the same trend as observed in other Andean forests (Gentry 1992, 1995, RangelCh. 1991). In UMRF the Lauraceae family is the most diverse in species, which is
exceptional, followed by the Melastomataceae and Myrtaceae. In SARF
Asteraceae and Ericaceae are the most species rich families.
In páramo vegetation Asteraceae and Poaceae rank as most species rich. This is the
case in almost all páramo vegetation and in the flora‟s of other alpine areas
(Rangel-Ch. 2000c; Vargas & Sánchez 2005; Rivera-Diaz 2007; Rangel-Ch. et al.
2008; Briceño & Morillo 2002, 2006; Lozano et al. 2009), followed by Ericaceae
and Orchidaceae. Pteridophytes are also species diverse, with Grammitidaceae,
Lycopodiaceae, and Dryopte-ridaceae as the most species rich families.
Species diversity and composition of montane forests of Ramal de Guaramacal
varies along the altitudinal gradient with some bias depending slope exposure.
Species richness generally decreases with elevation. However, an increase in
species richness based on 0.1 ha plots was locally observed between 2300 and
2400 m on the northern slope of Guaramacal. At this elevation is the transitional
zone from LMRF to UMRF, and here species richness may be related to the
increasing humidity from the dry interandean Boconó Valley to the top of the
mountain. This diversity pattern supports a proposed “third pattern” of altitudinal
species richness claiming highest biodiversity in the middle of an altitudinal zone
(Lomolino 2001). Wolf (2003) already pointed at high richness of epiphytic
bryophytes and lichens in the mid-altitudinal range of a zone.
With a limited altitudinal span (2820-3130 m) and a small surface area of ca. 10
km2, the Páramo de Guaramacal counts some 200 vascular páramo species (alpha
diversity). Compared to the number of 1544 vascular species reported for all
Venezuelan páramos [1437 angiosperms species reported by Briceño & Morillo
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(2002, 2006) plus 107 fern species reported by Luteyn (1999)] the low species
number of Páramo de Guaramacal seems proportional. At an elevation of about
2200 m there is a small direct connection to the Mérida Andes.
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.
Repeated isolation in the past probably triggered the development of endemic
species. The highest species diversity of Ruilopezia rosettes reported for the
Venezuelan Andes to date is in our study area and it may be speculated that during
the Pleistocene repeated isolation and merging of populations gave rise to
endemics.
7.4 VEGETATION PHYSIOGNOMY
The structure of the montane forests of Ramal de Guaramacal is more compressed
towards higher elevations, with an increase in stem density and a decrease in stem
diameter and canopy height. LMRF is dense and of medium height, with canopies
up to 25 m tall, while UMRF canopies reach up to 18 m, and those of SARF to 6-8
(10) m only.
Diversity and density of growth forms varies with elevation. More diversity and
density of palms, lianas and climbers is clearly observed in LMRF. Although
diversity and density of lianas decrease with altitude, still a substantial percentage
of the total species richness of SARF is represented by liana species. Tree ferns
show highest density in the LMRF, but highest species diversity is observed in
UMRF.
Zonal páramo vegetation is represented by shrub páramo, bunchgrass páramo and,
in our study area most commonly by bamboo páramo. Bamboo páramos are
mainly dominated by woody growth forms, particularly upright shrubs with
bamboo groves and clumps, which give an overall appearance of a mostly shrub
páramo vegetation. Low bunchgrass páramo vegetation, devoid of shrubs and with
a high density of small ground rosettes, cushion grasses and few bamboos, is found
in limited areas above 2900 m. In the study area the most representative life form,
in terms of both number of species and cover, are the phanerophytes, especially
those belonging to the microphanerophytic type, followed by hemicryptophytes
with a caespitose life form.
The dominance of shrubby growth forms in páramo of Ramal de Guaramacal may
be partly explained by the high relative humidity, the low altitudinal range, the
close proximity of the dwarf forests near the upper forest line, past disturbance
events and fire dynamics.
Azonal páramo vegetation is represented by patches of azonal bunchgrass,
Sphagnum bogs, aquatic communities, and boggy bamboo páramo.
7.5 PHYTOSOCIOLOGICAL CLASSIFICATION
The phytosociological classification of the vegetation of Ramal de Guaramacal
was based on floristic composition and species abundance or cover. Results
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Flora, vegetation and ecology in the Venezuelan Andes
revealed a total of eight new alliances and eighteen vegetation communities at
association level, which are distinguished and described according to the ZürichMontpellier method (Braun-Blanquet 1979; Westhoff & van der Maarel 1973).
We recognized and documented three new subandean forest (LMRF) communities,
and four new Andean forest (UMRF) and high Andean forest (SARF); three new
associations of lower subpáramo or shrubby páramo, and two new associations of
upper subpáramo or bunchgrass páramo dominated by rosettes and tussock plants;
one new azonal bunchgrass páramo association, two new Sphagnum bog
associations, one new bamboo páramo („chuscal‟) association, and two submerged
aquatic communities. A synoptic syntaxonomical scheme for classification of the
vegetation of Ramal de Guaramacal runs as follows:
I. Montane forest group of Meliosma tachirensis – Alchornea grandiflora
A. The alliance Geonomo undatae – Posoquerion coriaceae Cuello & Cleef 2009,
which contains the following subandean forests associations:
1. Simiro erythroxylonis – Quararibeetum magnificae Cuello & Cleef 2009
2. Conchocarpo larensis – Coussareetum moritzianae Cuello & Cleef 2009
B. The alliance Farameo killipii – Prunion moritzianae Cuello & Cleef 2009. This
alliance contains one subandean forest community and one Andean forest
community:
3. Croizatio brevipetiolatae – Wettinietum praemorsae Cuello & Cleef 2009
4. Schefflero ferrugineae – Cybianthetum laurifolii Cuello & Cleef 2009
C. The alliance Ruilopezio paltonioides – Cybianthion marginatii Cuello & Cleef
2009. This includes one Andean and two high Andean forest communities:
5. Geissantho andini – Miconietum jahnii Cuello & Cleef 2009
6. Gaultherio anastomosantis – Hesperomeletum obtusifoliae Cuello & Cleef
2009
7. Libanothamnetum griffinii Cuello & Cleef 2009
II. Zonal humid lower páramo of Ruilopezio lopez-palacii – Chusqueetalia
angustifoliae Cuello & Cleef 2009 (prov. Ord.)
D. The alliance Hyperico paramitanum – Hesperomeletion obtusifoliae Cuello &
Cleef 2009, groups the shrubby páramo associations:
8. Ruilopezio paltonioides – Neurolepidetum glomeratae Cuello & Cleef 2009
9. Disterigmo acuminatum – Arcytophylletum nitidum Cuello & Cleef 2009
E. The alliance Hyperico cardonae – Xyridion acutifoliae Cuello & Cleef 2009,
groups one shrubby páramo and two open grass páramo associations:
10. Cortaderio hapalotrichae – Hypericetum juniperinum Cuello & Cleef 2009
11. Puyo aristeguietae – Ruilopezietum lopez-palacii Cuello & Cleef 2009
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12. Rhynchosporo gollmeri – Ruilopezietum jabonensis Cuello & Cleef 2009
III. The order of azonal páramo peat bog vegetation of Geranio stoloniferum –
Caricetalia bonplandii Cuello & Cleef 2009
F. The alliance Sphagno recurvi – Paepalanthion pilosi Cuello & Cleef 2009,
groups a bunchgrass páramo association and the both new Sphagnum bog
associations:
13. Paepalantho pilosi – Agrostietum basalis Cuello & Cleef 2009
14. Sphagno recurvi – Caricetum bonplandii Cuello & Cleef 2009
15. Sphagno sparsi – Caricetum bonplandii. Cuello & Cleef 2009
G. The alliance Carici bonplandii – Chusqueion angustifolia Cuello & Cleef 2009,
contains a bamboo páramo („chuscal‟) association:
16. Carici bonplandii – Chusqueetum angustifoliae Cuello & Cleef 2009
H. The alliance Districho submersi – Isoëtion Cleef 1981 is represented by:
17. The submerged aquatic community of Sphagnum cuspidatum
18. Isoëtetum karstenii Cleef 1981
For both montane forest and páramo vegetation classes could not yet be defined on
the basis of the present number of relevés and other available information, neither
due to the lack of similar data to characterise the region and from montane forests
and Chusquea angustifolia bamboo páramos elsewhere in Venezuela and adjacent
Colombia. Comparison of zonal páramo communities of Chusquea angustifolia is
at present impossible.
More relevés, in particular in the zones of LMRF and UMRF may provide helpful
information to better classify the alliance Farameo killipii - Prunion moritzianae,
which includes forest associations of both LMRF and UMRF. More studies with
comparable aims in the surrounding mountains enable a forest classification at
order and class level.
This study represents the first attempt to classify the vegetation phytosociologically based on a quantitative data set from an entire mountain range in the Venezuelan Andes. Despite of the relatively low number of relevés and methodological
constraints, we arrived at a clear forest classification for the montane forests of
Ramal de Guaramacal.
Zonal páramo vegetation of the Guaramacal range was described on the basis of a
relatively low number of relevés from the most accessible páramo areas of Ramal
de Guaramacal (sector of Las Antenas of Páramo de Guaramacal) . The most
different physiognomic formations are found in sector Las Antenas in relatively
close proximity. This sector shows a larger altitudinal range (2820~3130 m).
However it has a past history of disturbances and fires, which may have affected
the spatial distribution of vegetation communities and consequently the current
situation of the upper limit of the forest. The resulting zonal páramo classification
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Flora, vegetation and ecology in the Venezuelan Andes
at association level may be representative of most páramo areas of Ramal de
Guaramacal.
The azonal páramo vegetation has been described on the basis of a limited number
of relevés and from two peat bogs and a small pond only located in two páramo
areas of Ramal de Guaramacal. The latter includes the Páramo de Guaramacal and
the Páramo El Pumar. The limited accessibility of the study area throughout most
of the year, together with high precipitation levels and the frequency of mist, made
the exploration of peat bog areas of Guaramacal difficult. As other unexplored peat
bogs are known to exist in the area other azonal vegetation communities may be
present in the páramos of Ramal de Guaramacal.
An integrated study of regional importance of the Sphagnum bogs of the northern
Andes is still lacking as most studies report on local peat bog types only. Despite
their presence in the páramos of the Sierra Nevada de Mérida Sphagnum bog
communities have not yet been formally reported. Sphagnum bogs in the páramos
of Ramal de Guaramacal have shown two new associations belonging to the new
alliance Sphagno recurvi - Paepalanthion pilosi. Peatbog communities in Colombia
share Carex bonplandii, Sphagnum magellanicum and S. sancto-josephense with
the Sphagnum bog communities of Guaramacal. We have not found other shared
species as a basis to establish relationships. Isolation, low altitude, and an
inadequate number of phytosociological studies account for the observed
assemblage of species in Ramal de Guaramacal.
The phytosociological classification of montane forest and páramo vegetation in
the Venezuelan Andes has just started with the present study. For a proper
management and conservation this mission needs to be continued in order to
develop a strong tool for vegetation mapping.
7.5 PHYTOGEOGRAPHY
The phytogeographical composition at genus level of páramo flora of Ramal de
Guaramacal shows a relatively high proportion of neotropical-montane elements
(ca. 40%) compared to other páramo floras. This characteristic is considered a
consequence of the humid climate and the low altitude of the Guaramacal range.
The latter could also be the cause of the low proportion of endemic páramo taxa. It
is plausible that the wet climatic conditions on the low range of Guaramacal since
Holocene time (Van der Hammen 1974; Salgado-Laboriau 1979, 1980) has served
as a filter preventing the arrival and survival of dry páramo species originating
from the Mérida páramos. The low representation of Andean-alpine (0.9%) and
Holarctic (3.7%) genera is in support of this suggestion.
The presence of Isoëtes karstenii in a small glacial lake, a submerged species
known from grass páramo up to the highest lakes in the superpáramo in Colombia
(Cleef 1981; Salamanca et al. 2003) and Venezuela (Fuchs-Eckert 1982; Small &
Hickey 2001) suggests that páramo vegetation with some form of superpáramo,
nowadays completely absent, could have occurred during glacial times at lower
altitudes along the steep slopes of the uppermost parts of Guaramacal range.
Roches moutonnées are commonly present along the ridges and are a testimony the
past glaciations on the top of Guaramacal ridge.
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The vascular flora of the Páramo de Guaramacal is largely composed of (1) a
group of neotropical widespread species (31%), (2) a group of Andean distributed
species (49%), a part of this group is confined to the northern Andes and another
part is widespread in the Andes from Colombia to Bolivia, and (3) a group of
Venezuelan endemics (20%).
The vascular páramo flora of Ramal de Guaramacal shows neither close
relationship to the flora of the dry páramos of the extended and high elevated
central part of the Sierra Nevada de Mérida nor to some other nearby páramos of
Colombian Cordillera Oriental. At the genus level some similarities may be found
between the páramo flora of the Podocarpus National Park in southern Ecuador
and the flora of the páramo/SARF mosaic of Guaramacal. Both locations do have
some genera in common and show a high proportion of neotropical-montane
elements. Both páramo areas have a permanent high humidity level, are relatively
low in altitude, and have a smooth topography. In southern Ecuador also a gradual
transition from SARF into shrub páramo has been observed.
7.6 FOREST FUNCTIONAL DIVERSITY AND ALTITUDE
Increasing deforestation and global warming are potential threats for Andean
forests. Losses in forest cover and biotic attrition might be exacerbated by
degradation in functional diversity, i.e. the variety of life-history traits presented
by an assemblage of organisms (Mayfield et al. 2005; Girao et al. 2007).
Considering the role of temperature changes in ecosystem functioning along
mountain slopes (Chapin & Körner 1995; Colwell et al 2008; Svenning & Condit
2008) and the importance of analysing changes of functional traits along altitudinal
gradient as of potential value for predicting the effects of environmental changes
on ecosystem functioning (Díaz & Cabido 1997; Díaz et al. 1999; Duckworth et al.
2000; Lavorel & Garnier 2002; McGill et al. 2006), functional diversity was
studied in relation to altitude in undisturbed Andean forests of Ramal de
Guaramacal (Guaramacal National Park). The aim was to contribute with reference
information for studies of degraded Andean systems.
Information of the vascular plant species composition of forest relevés sampled
along the altitudinal gradient (Chapter 2) was linked to different species functional
traits related to the energy balance (growth form, leaf shape and leaf size) or
fragmentation (dispersal and pollination modes, fruit type, and flower and fruit
size). This information was obtained from the literature and herbarium studies.
Information of species traits and altitude from plots surveys were summarized by
means of ordination analysis to detect the principal variation. Randomizations of
the species assemblages in relevés (Legendre et al. 1997; Dray and Legendre
2008) were used to test if the composition and diversity of energy-related traits and
fragmentation-related traits changed with elevation.
Results show that functional diversity of fragmentation-related traits decrease with
elevation (more underdispersion at higher elevations), and the energy-related traits
increase (more overdispersion at higher elevations). Overdispersion occurs when
better adapted species outcompete functionally to related species from the local
community (Weiher & Keddy 1995; Mayfield et al. 2005). Leaf size contributed
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Flora, vegetation and ecology in the Venezuelan Andes
substantially to the altitudinal variation in energy balance related traits. The
smaller leaf size at higher altitudes in wet tropical forests has been repeatedly
reported (Grubb et al. 1963; Vareschi 1966; Sugden 1985). In the absence of
pronounced dryness, as is the case in the situation along the slopes of the Ramal de
Guaramacal, smaller leave size can be explained by a lower temperatures upslope,
less hydraulic conductance of stems and associated lower mineral supplies, lower
nutrient availability, and an increased frost frequency (Cavelier 1996). Overall,
nitrogen becomes more limiting at higher altitudes in montane wet forests (Grubb
1977; Cavelier 1996). Therefore, our results suggest that competition for
resources, which is mostly related to the capture of radiation (heat) and the uptake
of minerals and nutrients, is an important driver of species composition.
The association between traits showing a functional diversity of energy balance
and traits related to elevation further suggests that a temperature increase due to
Global Change will affect forest functionality of Andean forests. Higher
temperatures in the coming decades might reduce functional diversity of energy
balance along the slopes of the Andes. This projection should be treated with
caution, however, because effects of changing temperatures on species migration
and the way ecological filtering has impact on species interactions, is unknown.
Results also suggests that even in undisturbed Andean forest functional diversity
varies significantly, which implies that in the nearby future a lower degree of
functional diversity at a certain elevation does not necessarily endangers
ecosystem well-being.
The negative altitudinal association of the main variation in fragmentation trait
state composition is considered mostly due to the presence of wind-dispersed,
wind pollinated, and fern species from SARF (e.g. species from Alsophila,
Cyathea, Dicksonia, Diplazium, Baccharis, Diplostephium, Pentacalia, Mikania).
Since SARF and páramo vegetation occur in the study area in a spatially well
mixed mosaic (Chapters 2 and 3), the predominance of the trait states in the
samples at highest elevation related to wind dispersal can be explained by the
functional role of wind as an edge effect. There is a tendency that species with
larger fruits and flowers occurr at relatively low elevations (e.g. Symbolanthus
vasculosus, Zygia bisingula, Drymonia crassa, Tabebuia guayacan,
Macrocarpaea bracteata, Inga edulis), which may be related to a more
pronounced role of mammals (including large bats) at lower elevations versus a
more dominant role of birds in pollination and seed dispersal at elevations above
2100 m.
Several SARF plots showed an outlying functional diversity compared to the
trends in the forest relevés at lower elevations. This suggests that the plants in the
SARF relevés contained markedly more trait states relative to the number of
species, and/or more plant individuals relative to the variety in trait states as is the
case at lower elevations. Both phenomena may be caused by the increased wind
vigor in SARF enhancing the number of plants with traits related to pollination and
dispersal by wind.
The diversity of fragmentation related traits tend towards a negative relationship
with elevation, visible in both species-based and individual-based indices of the
montane forests. Hence, our results indicate that along an undisturbed altitudinal
gradient in Andean rain forest ecological differentiation driven by disturbance is
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decreasing in upslope direction. Possibly, the increased level of underdispersion in
upslope direction is due to a higher fragmentation of the forest matrix related to
more frequent terrain dissections. More information on forest dynamics and forest
disturbance in relation to mass movements and slope instability in the Guaramacal
area is needed to better understand mechanisms that explain the upward decrease
of functional diversity of disturbance related traits.
According to our expectations we concluded that in the Guaramacal area
functional diversity of undisturbed Andean forest is changing with altitude. This
implies that in the nearby future a temperature rise might affect the functionality of
Andean forest, but not necessarily in a harmful way. Functional diversity related to
energy balance traits increases in upslope direction, pointing to increased levels of
ecological differentiation. We explained this by assuming more upslope
competition in the Andean forests regarding capture of radiation and the uptake of
minerals and nutrients. Diversity in fragmentation related traits showed an
opposite pattern (more underdispersion upslope), which might relate to
discontinuities in the forest matrix due to the geomorphology in the study area.
SARF diverged from the observed altitudinal trends in fragmentation related traits,
probably as a consequence of edge effects created by wind in the SARF-páramo
mosaic.
7.7 CONSERVATION IMPLICATIONS
Tropical montane forests and páramos are fragile ecosystems and hold a high and
exceptional biodiversity. In the Venezuelan Andes there is a net of national parks
and reserves that have kept UMRF and páramo ecosystems relatively well
protected. However, LMRF is most affected by human intervention and has been
largely converted into areas with an agricultural land use (Ataroff 2000). Outside
protected areas montane ecosystems have been severely affected and fragmented,
leaving natural montane forest as remnants only.
Fortunately, Ramal de Guaramacal and its montane ecosystems is among the best
conserved national parks in Venezuela. Results of this thesis are important
contributions to the knowledge of floristic diversity and botanical composition of
our forests and páramos. The information from this mountain range is basic for
conservation planning and restoration of these ecosystems. However, more studies
on ecosystem dynamics from other mountain systems are needed to develop a
proper management planning and conservation on a wider regional scale.
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205
APPENDIX
Appendix
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Appendix 1. Checklist and vouchers of vascular species recorded from plots surveys in the
montane forest of Ramal de Guaramacal. Species are alphabetically ordered by
class, family and genus. Asterisc (*) indicates annotated/collected for forest
description, but not documented in the plot surveys. Collectors NC: N. Cuello
et al., AL: A. Licata et al.
Lycopodiopsida: LYCOPODIACEAE: Huperzia sp. * NC 2822; H. sp. * NC 2268; H.
mollicoma (Spring) Holub. * NC 1471; Lycopodium contiguum Kl. * NC 2696; L. jussiaei Desv.
ex Peir * NC 1075; SELAGINELLACEAE: Selaginella difussa (C. Presl) Spring * NC 1452; S.
producta Baker. * NC 1472; S. substipitata Spring * NC 1298. Filicopsida: ASPLENIACEAE:
Asplenium sp. * NC 2814; A. alatum Alvaro Cogollo * NC 2150; A. auriculatum Sw. * AL 220,
235, 260; A. cirrhatum Rich. ex Willd * NC 2385; A. cristatum Lam. * NC 1860, 1862; A.
cuspidatum Lam. * AL 221; A. flabellulatum Kunze * AL 222; A. harpeodes Kunze * AL 228,
251; A. raddianum Gaudich. * NC 1247, 1523; A. radicans L. * NC 1631, 2014; A. uniseriale
Raddi * NC 1719. BLECHNACEAE: Blechnum ensiforme (Liebm.) C. Chr. * NC 1305; B.
schomburgkii (Klotzsch) C. Chr. CYATHEACEAE: Alsophila angelii Tryon NC 2432; A.
erinacea (Karst.) Conant. NC 1765, 2209, 2224, 2302; Cyathea aff. straminea H. Karst NC 2394;
C. caracasana (Klotzsch) Domin NC 1101, 1226, 1276, 1857, 1875, 1938, 2391, 2473, 2482,
2483; C. fulva (Mart. & Gal.) Fée NC 1032, 1701, 1887, 2011, 2041, 2096, 2365, 2396, 2461,
2582,2286,2295, 2301; C. kalbreyeri (Baker) Domin NC 1512, 1763, 2206, 2603, 2624; C.
pauciflora (Kuhn) Lellinger NC 1157, 1971, 2353, 2392, 2633; C. pungens (Willd.) Domin NC
1324; Sphaeropteris sp. NC 1051. DENNSTAEDTIACEAE: Dennstaedtia sp. * NC 1473.
DICKSONIACEAE:
Dicksonia
sellowiana
Hook.
NC
1873,
1921,
2429.
DRYOPTERIDACEAE: Arachniodes denticulata (Sw) Ching * NC 1246, 1477, 1628;
Didymochlaena truncatula J. Smith * NC 2151; Diplazium celtidifolium Kunze NC 2154; D.
hians Kunze ex Klotzsch. NC 1841, 2136; Elaphoglossum cuspidatum (Will.) Moore * AL 266;
E. eximium (Mett.) H. Christ * AL 245; Polystichum muricatum (L.) Fée * NC 2181; P.
platyphyllum (Will.) C. Presl * NC 1863. GRAMMITIDACEAE: Melpomene flabelliformis
(Poir.) A.R. Sm. & R.C. Moran * NC 2877; M. xiphopteroides (Liebm.) A.R. Sm. & R.C. Moran
* AL 270; Micropolypodium truncicola (Klotzsch) A.R. Sm. * AL 275; Terpsichore asplenifolia
(L.) A. R. Sm. * NC 1499, 2427; T. subtilis (Kunze ex Klozsch) A. R. Smith, vel aff * AL 271; T.
taxifolia (L.) A. R. Sm. * NC 1302; T. xanthotrichia (Klotzsch) A. R. Smith * AL 244.
HYMENOPHYLLACEAE: Hymenophyllum fucoides (Sw.) Sw. * NC 1317; H. microcarpum
Desv. * NC 1492; H. myriocapum Hook. * NC 1465; H. polyanthos (Sw.) Sw. * AL 248; H.
trichomanoides Bosch * NC 1489, 1491; Trichomanes capillaceum L. * NC 1490; T. radicans
Sw. * NC 2152. MARATTIACEAE: Danaea moritziana C. Presl. * NC 1533.
POLYPODIACEAE :Campyloneurum ophiocaulon (Klotzsch) Fée * NC 2153; C. serpentinum
(H. Christ) Ching. * NC 1846; Microgramma percussa (Cav.) de la Sota * NC 1630; Pecluma
divaricata (E. Fourn.) Mickel & Beitel * NC 2089; Polypodium sp.* NC 2809; P. buchtienii H.
Christ. & Rosenst. * AL 287; P. fraxinifolium Jacq. * NC 1248. PTERIDACEAE: Eriosorus
flexuosus (Kunth) Copel. * NC 2824. THELYPTERIDACEAE: Thelypteris concinna (Will.)
Ching * AL 337; T. dentata (Forssk.) E.P. St. John * AL 338. VITTARIACEAE Polytaenium
lineatum (Sw.) J. Sm. * NC 1203; Vittaria graminifolia Kaulf. * AL 274.
Pinopsida: PODOCARPACEAE: Podocarpus oleifolius D. Don ex Lambert var. macrostachyus
(Parl.) J. Bunchholz & N. E. Gray NC 1126, 2410, 2647.
Magnoliopsida: ACANTHACEAE: Aphelandra macrophylla Leonard NC 1736, 2535, 2557;
Mendoncia tovarensis (Klotzsch & Karsten ex Nees) Leonard NC 1800; Ruellia tuberosa L. *
NC 2499; R. tubiflora Kunth var. tetrastichantha (Lindau) Leon NC 1038. ACTINIDIACEAE:
Saurauia tomentosa (Kunth) Spreng. NC 1190; S. yasicae Loes NC 1026, 1143.
ANACARDIACEAE: Tapirira guianensis Aubl. NC 1546, 1854, 2281. ANNONACEAE:
Rollinia mucosa (Jacq.) Baill. NC 1814, 2558; Trigynaea duckei (R.E. Fr.) R.E. Fr. NC 1526,
2512. AQUIFOLIACEAE: Ilex guaramacalensis Cuello & Aymard, sp. nov. NC 2853; I. laurina
Kunth NC 1147, 1338, 1920, 1961, 2599, 2693; I. myricoides Kunth NC 1140; I. sp.1 NC 1189;
I. sp.2 NC 1333, 2462, 2470; I. truxillensis Turcz. subsp. bullatissima Cuatrec. NC 1120, 2412,
209
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Flora, vegetation and ecology in the Venezuelan Andes
2471. ARALIACEAE: Dendropanax arboreus (L.) Dcne. & Planch. NC 1503, 1787, 2297;
Oreopanax discolor (Kunth) Decne. & Planch. NC 1257, 2448; O. sp. NC 2768, 2826; Schefflera
ferruginea (Willd. ex Roem. & Schult.) Harms NC 1083, 1328, 1966, 2393. ASTERACEAE:
Ageratina neriifolia (B.L. Rob.) R.M. King & H. Rob. NC 2079; A. theifolia (Benth.) R. M. King
& H. Rob. NC 2881; Baccharis brachylaenoides DC. NC 2869; Critoniopsis paradoxa (Sch.
Bip.) V.M. Badillo NC 1110, 1253, 1291, 1318, 1935, 2625 Diplostephium obtusum S.F. Blake
NC 2678; Fleischmannia pratensis (Klatt) R.M. King & H. Rob. * NC 1548; Libanothamnus
griffinii (Ruiz-Teran & Lop. Fig.) Cuatr. NC 2704; Mikania banisteriae DC. NC 1004, 2010,
2379; M. bogotensis Benth. * NC 1355; M. houstonians (L.) B.L. Rob. NC 2232; M.
nigropunctulata Hieron NC 2068; M. sp.1 NC 2082; M. stuebelii Hieron NC 2075, 2316, 2363,
2852; Paragynoxys cuatrecasasii Ruiz-Teran & Lopez Figueiras NC 1217; P. venezuelae (V.M.
Badillo) Cuatrec. NC 1238; Pentacalia cachacoensis (Cuatrec.) Cuatrec. NC 1242; P.
greenmanniana (Hieron.) Cuatr. NC 2879; P. theifolia (Benth.) Cuatrec. NC 2838; P. vicelliptica
(Cuatrec.) Cuatrec. NC 2085; Ruilopezia paltonioides (Standl.) Cuatrec. NC 2616.
BEGONIACEAE: Begonia sp.* NC 2182; B. trispathulata (A. DC.) Warb. in Engler & Prantl *
NC 2553; B. vareschii Irmscher * NC 1464. BIGNONIACEAE: Schlegelia spruceana K. Schum.
NC 1816; Tabebuia guayacan (Seem.) Hemsl. NC 1609, 1790. BOMBACACEAE: Matisia sp.
NC 2508; Quararibea magnifica Pittier NC 2133, 2225, 2505. BORAGINACEAE: Cordia
cylindrostachya (Ruiz & Pav.) Roem. & Schult. * AL 283. BRASSICACEAE: Cardamine
fulcrata Greene * AL 272. BRUNELLIACEAE: Brunellia acutangula Humb. & Bonpl. NC
1241; B. cf. integrifolia Szyszyll. NC 1105, 1864, 1936, 2104, 2600. BURSERACEAE: Protium
tovarense Pittier NC 1283, 1558, 1778, 2257. CAESALPINIACEAE: Senna pendula (Humb. &
Bonpl. ex Willd.) H. Irwin & Barneby * AL 333. CAMPANULACEAE Centropogon cornutus
(L.) Druce * AL 309; C. elmanus Wimm. * NC 1420; C. solanifolius Benth. * NC 1479.
CAPRIFOLIACEAE: Viburnum tinoides L.f. var. venezuelensis (Killip & A. C. Smith) Steyerm.
NC 1258, 1919. CARICACEAE: Vasconcella microcarpa (Jacq.) A. DC. NC 1201, 2526.
CARYOPHYLLACEAE: Drymaria cordata (L.) Willd. ex Roem. & Schult. * NC 1457; D.
ovata willd. ex Roem & Schult * NC 1439; Stellaria ovata Willd. * NC 2810; Stellaria sp. * NC
2763. CELASTRACEAE: Celastrus liebmanii Standl. NC 1011, 2241, 2357; Maytenus sp. A NC
1293, 1582, 1659, 2626; Perrottetia quinduensis Kunth NC 2030, 1003, 1068.
CHLORANTHACEAE: Hedyosmum cf. gentryii D'Arcy & Liesner NC 1518, 1593, 1813, 1870,
2008, 2283, 2380, 2604; H. crenatum Occhioni NC 1115, 1933, 1949, 2101, 2254; H.
cuatrecazanum Occhioni NC 989, 1025, 2047; H. goudotianum Solms-Laubach * NC 1441; H.
racemosum (Ruiz & Pav.) G. Don NC s/n; H. sp. A NC 1232, 1323, 2450, 2629; H. translucidum
Cuatrec. NC 1223, 1326, 2799. CLETHRACEAE: Clethra fagifolia Kunth var. fagifolia NC
1059, 1148, 1330, 1931, 1959, 2356. CLUSIACEAE Clusia alata Triana & Planch. NC 1215,
1219, 1356, 1578, 1883, 1942, 1967, 2001, 2051, 2445, 2617, 2250, 2453; C. sp. 1 NC 2270,
2360; C. sp. A? (C. multiflora group) NC 1676, 1748, 1970; C. trochiformis Vesque NC 1017,
1113, 1255, 1288, 1357, 1674, 1934, 1968, 2120, 2219, 2467, 2251, 2408, 2510; Hypericum
paramitanum N. Robson NC 2831; H. thesiifolium Kunth * AL 319; Vismia baccifera (L.) Triana
& Planch subsp. dealbata (Kunth) Ewan NC 1274. CUCURBITACEAE: Elateriopsis oerstedii
(Cogn.) Pittier NC 2514. CUNONIACEAE: Weinmannia aff. balbisiana Kunth NC 1648, 1877,
2106, 2451, 2638, W. auriculata D. Don NC 2849; W. fagaroides Kunth NC 2252, 2850; W.
glabra L.f. NC 1116, 1929, 1964, 2109, 2398; W. karsteniana Szyszyll. NC 2848; W. lechleriana
Engl. NC 1351; W. sorbifolia Kunth NC 1195, 2586. DICHAPETALACEAE: Dichapetalum
pedunculatum (DC.) Baill. NC 2263. ELAEOCARPACEAE: Sloanea brevispina F. Sm. NC
2265; S. guianensis Aubl. NC 919, 1541, 1775, 1995, 2170, 2231, 2271; S. laurifolia (Benth.)
Benth. NC 937; S. rufa Planch. ex Benth. NC 1070, 1513, 2003, 2140, 2276, 2287.
EREMOLEPIDACEAE: Antidaphne viscoidea Poepp. & Endl. * NC 1014. ERICACEAE:
Diogenesia tetrandra (A. C. Jm.) Sleumer NC 2118, 2589; Disterigma alaternoides (Kunth)
Nied. NC 2861; D. sp. NC 2437; Indet. Eric-1 NC 2417; Gaultheria anastomosans (L.f.) Kunth
NC 2680; G. erecta Vent. 2827; G. myrsinoides Kunth [Pernettya prostrata (Cav.) DC. NC
2706]; Macleania rupestris (Kunth) A.C. Sm. NC 1575; Psammisia hookeriana Klotzsch. NC
1125, 1637, 1896, 2063, 2334, 2338, 2498; P. penduliflora Klotzsch * NC 1717; Themistoclesia
dependens (Benth.) A. C. Smith NC 1974; Thibaudia floribunda Kunth. NC 2693; Vaccinium
corymbodendron Dunal NC 2687. ESCALLONIACEAE: cf. Escallonia hispida (Vell.) Sleumer
210
Appendix
_______________________________________________________
NC 2431. EUPHORBIACEAE: Acalypha macrostachya Jacq. NC 1177; Alchornea glandulosa
Poepp. & Endl. NC 1855, 2536; A. grandiflora Muell. Arg. NC 927, 1186, 1273, 1565, 1584,
1667, 1806, 2129, 2326, 2354, 2415; Croizatia brevipetiolata (Secco) Dorr NC 1703; Hyeronima
cf. oblonga (Tul.) Mull. Arg. NC 953, 1309, 1655, 1999, 2035, 2073, 2291, 2628; H. moritziana
(M. Arg.) Pax & Hoffmann NC 948, 1081, 1292, 1339, 1773, 1988, 2036, 2601; H. scabrida
(Tul.) Mull. Arg. NC 1231, 1352, 2447, 2618; Mabea occidentalis Benth. NC 1519; Phyllanthus
niruri L. * AL 325; Sapium stylare Mull. Arg. NC 2029; Tetrorchidium rubrivenium Poepp. NC
1196, 1812, 2022. FABACEAE: Desmodium intortum (Mill.) Urb. * AL 300; D. molliculum
(Kunth) DC. * NC 1400, 1438; Dussia coriacea (Sw.) Roem. & Schult. NC 1607, 1977, 2053,
2138; Machaerium cf. floribundum Benth. NC 1807, 2277. FLACOURTIACEAE: Casearia
tachirensis Sleumer NC 2032, 2165, 207. GENTIANIACEAE: Macrocarpea bracteata Ewan NC
2806; Symbolanthus vasculosus (Griseb.) Gilg. NC 1329, 1649, 1891, 2122, 2474.
GESNERIACEAE: Alloplectus aff. chrysantha Planch. & Linden * NC 1424; Besleria pendula
Hanst. NC 942, 1930; Columnea sanguinea (Pers.) Hanst. * AL 307; Drymonia crassa C. V.
Morton * NC 2492; D. crassa C.V. Morton NC 2048; Heppiella viscida (Lindl. & Paxt.) Fritzsch
* NC 1622, 1724; Kohleria hirsuta (Kunth) Regel * NC 1716. HIPPOCRATEACEAE: Salacia
aff. cordata (Miers.) Mennega NC 1786, 2259. HYDRANGEACEAE: Hydrangea aff. peruviana
Moricard NC 1180; H. cf. preslii Briq. NC 2238, 2525; H. sp.1 NC 2211. ICACINACEAE:
Calatola venezuelana Pittier NC 1005, 2135, 2221, 2282, 2564; Citronella costaricensis (Donn.
Sm.) R.A. Howard NC 1165, 1914, 2033. LAMIACEAE: Hyptis vilis Kunth & Bouché * AL
331. LAURACEAE: Aiouea dubia (Kunth) Mez NC 926, 1916, 1998, 2092, 2208, 2444, 2640;
Aniba cf. cinnamomiflora C. K. Allen NC 1514, 1596, 1780, 1902, 1992, 2262; Beilschmiedia
tovarensis (Meissn.) Sa. Nishida NC 1313, 1587, 1888, 1987, 2023, 2245, 2266; Endlicheria sp.
NC 2285, 2330; Nectandra aff. membranacea (Sw.) Griseb. NC 1835, 2532; N. aff. purpurea
(Ruiz & Pav.) Mez NC 1536, 2520; N. sp. NC 1838; Ocotea aff. puberula (Rich.) Nees NC 922,
1991; O. aff. tarapotana (Meissn.) Mez NC 1799; O. auriculata Lasser NC 1178; O. calophylla
Mez NC 2460; O. cernua (Nees) Mez, vel aff. NC 1828, 2162; O. cf. hexanthera Kopp. NC
2369, 2397; O. floribunda (Sw.) Mez NC 1010, 1037, 1539, 2099, 2351; O. jelski Mez NC 1137,
1962; O. karsteniana Mez NC 1123, 1197, 1307, 1707, 1912, 1996, 2121, 2414; O. leucoxylon
(Sw.) de Lanessen, s.l. NC 1664, 1893, 2110, 2366, 2438; O. macropoda (Kunth) Mez NC 1084,
1791, 1880, 1960, 2115, 2318, 2475; O. rubrinervis Mez NC 1563, 2000; O. sericea Kunth NC
2076, 2479; O. sp. s/n; O. sp. A NC 1044, 1295, 1660, 1867, 2290; O. sp. C NC 2518, 2569,
2574; O. terciopelo C. K. Allen NC 2086; O. vaginans (Meissn.) Mez NC 1045, 1290, 1989,
2037, 2641; Persea aff. mutisii Kunth NC 1928, 1954, 1975, 2449, 2464; P. ferruginea Kunth.
NC 2480; P. meridensis Kopp. NC 943, 1985, 1897, 1885; P. peruviana Nees NC 976, 1569,
1817, 2142, 2223, 2575; P. sp.1 NC 1953, 1963, 2472; P. sp.2 NC 2434; P. sp.3 NC 1770;
Pleurothyrium costanense van der Werff NC 1188, 2128. LECYTHIDACEAE: Eschweilera
perumbonata Pittier NC 1521, 1646, 2260, 2278; E. sp. nov NC 1832. LORANTHACEAE:
Aetanthus nodosus (Desr.) Engl. * NC 1066, 1345, 2401; Dendrophtora sp. * NC 2795;
Gaiadendron punctatum (R. & P.) G. Don NC 1121, 2084, 2847; Struthanthus dichotrianthus
Eichl. * AL 320. LYTHRACEAE: Cuphea denticulata Kunth * NC 1401. MAGNOLIACEAE:
Talauma sp. NC 1745. MALPIGHIACEAE: Bunchosia armeniaca (Cav.) DC. NC 1181, 2126;
Byrsonima karstenii W. R. Anderson NC 1131; B. sp. NC 2439, 2481, 2642; Mascagnia sp. A
NC 1552. MARCGRAVIACEAE: Marcgravia brownei (Tr. & Pl.) Krug & Urb. NC 2176, 2578.
MELASTOMATACEAE: Anaectocalyx bracteosa (Naud.) Triana NC 1112, 1254, 1285, 1685,
1907, 1941, 2078, 2341, 2477; Blakea schlimii (Naud.) Triana NC 944, 1277, 1571; Chaetolepis
lindeniana (Naudin) Triana NC 2691; Henriettella cf. verrucosa Triana NC 1769; H. sp. NC
2303; H. tovarensis Cogn. NC 1550; Meriania grandidens Triana NC 2002, 2046, 2648; M.
macrophylla (Benth.) Triana NC 1997, 2141; Miconia aeruginosa Naud. * NC 1549; M.
amilcariana Almeda & Dorr NC 999, 1185, 1327; M. cf. dolichopoda Naud. NC 1174, 1836,
2025, 2098, 2299, 2442, 2534; M. donaeana Naud. NC 2577; M. elvirae Wurdack NC 1362; M.
cf. minutiflora (Bonpl.) DC. NC 960, 1598, 1662, 2045, 2594; M. jahnii Pittier NC 2828; M.
lonchophylla Naud. NC 1043, 1555, 1577, 1771, 1886, 207, 2215, 2300, 2346; M. lucida Naud.
NC 932, 941, 1657, 1900, 1978, 2333, 2342; M. mesmeana Gleason subsp. longipetiolata
Wurdack NC 1948; M. sp. C (hibrido) NC 1028; M. sp.B NC 2644; M. spinulosa Naudin * NC
1460; M. suaveolens Wurdack NC 1222; M. theaezans (Bonpl.) Cogn., s.l. NC 1151, 1237, 1271,
211
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
1349, 1540, 2637; M. tinifolia Naud. NC 1106, 1325, 1336, 2062, 2436, 2631; M. tovarensis
Cogn. NC 1278, 1231, 2621; M. tuberculata (Naud.) Triana * AL 315; M. ulmarioides Naud. NC
1119, 1225, 1320, 1939, 2097, 2390; Monochaetum discolor H. Karst. NC 2710; Mouriri
barinensis (Morley) Morley NC 1504, 1830, 2205; Ossaea micrantha (Sw.) Macfad. NC 2168.
MELIACEAE: Guarea kunthiana A. Juss. NC 1171, 1525, 1810, 2210,2555, 2568; Ruagea
glabra Triana & Planch. NC 928; R. pubescens H. Karst. NC 1270, 1642, 1801, 1881, 2361,
2523; Trichilia hirta L. NC 2517; T. pallida Sw. NC 1545; T. septentrionalis C. DC. NC 1909,
2043, 2164, 2213. MIMOSACEAE: Inga aff. densiflora Benth. NC 1595, 2226, 2605; Inga
edulis Mart. NC 1179, 2524; Zygia bisingula L. Rico NC 1737. MORACEAE: Cecropia
sararensis Cuatrec. NC 1746; C. sp. NC 2311; C. telenitida Cuatrec. NC 1826; Ficus
nymphaefolia P. Miller NC 1793, 2538; F. sp. NC 1843; F. tonduzii Standl. NC 1198, 2145,
2570; F. tovarensis Pittier NC 2294; Morus insignis Bureau NC 2527; Pseudolmedia rigida
(Planch. & Karst.) Cuatrec. subsp. rigida NC 1537, 2273.MYRICACEAE: Myrica pubescens
Willd. * AL 314; MYRSINACEAE Cybianthus cuspidatus Miq. NC 1652, 1895; Cybianthus
iteoides (Benth.) Agost. NC 1141, 2403; C. laurifolius (Mez) Agost. NC 1153, 1926, 2409; C.
marginatus (Benth.) Pipoly NC 2679; C. stapfii (Mez) Agostini NC 2832; Geissanthus fragans
Mez NC 1199, 2620; Myrsine dependens (Ruiz & Pav.) Spreng NC 2792; M. coriacea (Sw.) R.
Br. ex Roem & Schult. NC 1107, 2117; Parathesis venezuelana Mez NC 1744, 2173, 2506;
Stylogyne longifolia (Mart. ex Miq.) Mez NC 2530, 2559; S. sp. A NC 2236, 2645; Geissanthus
andinus Mez NC 2863. MYRTACEAE: Calyptranthes cf. meridensis Steyerm. NC 1104, 1951,
1965, 2343, 2359, 2646; Calyptranthes sp. NC 2446; Eugenia albida Humb. & Bonpl. NC 1951,
1980; E. cf. oerstediana O. Berg. NC 1560, 1739, 1788,2264, 2264, 2298, 2584; E. cf. patens
Poir. NC 1337, 1766, 1792, 1798,1815; E. cf. tamaensis Steyerm. NC 951, 954,1021, 1142, 1133,
1127,1874, 2114, 2220, 2348, 2340, 2345, 2406,2430; E. grandiflora O. Berg. NC 1535, 1625,
1591, 1603; E. moritziana H. Karst. NC 1173, 2519; E. sp. 1 NC 1567, 1617, 2292, 2350, 2350;
E. sp. 2 NC 1889, 1905, 2222; E. sp. 3 NC 2161, 2166, 2218; E. triquetra Berg NC 2804; Myrcia
acuminata (Kunth) DC. NC 1616, 1670, 1673, 1680, 2595, 2608, 2650; M. aff. guianensis
(Aubl.) DC. NC 1130, 1144, 1280, 2413; M. cf. sanisidrensis Steyerm. NC 1272, 1649, 1709,
1876, 2202, 2368, 24692, 2399, 2404, 2443; M. sp.1 NC 1866, 1899, 1903, 2615; Myrcianthes
sp. NC 1023; Myrtaceae sp 2 p14 NC 1510, 1774, 1840, 2513, 265; Myrtaceae-indet. ‘hojita
chiquita’ NC 2854. NYCTAGINACAE: Neea sp. NC 1851; OLACACEAE: Heisteria acuminata
(Humb. & Bonpl.) Engler NC 2511. OLEACEAE: Chionanthus sp. NC 2230. ONAGRACEAE:
Fuchsia nigricans Linden * NC 1493; Ludwigia peruviana (L.) H.Hara * AL 316.
PICRAMNIACEAE: Picramnia sp. A NC 1802, 1839, 2314; Picramnia sp. C NC 2137, 228,
2515, 2579. PIPERACEAE: Peperomia * NC 1391; Peperomia acuminata Ruiz & Pav. * NC
2204; P. ouabianae C. DC. * NC 1202; P. peltoidea H. B. K. * NC 2489; P. portuguesensis
Steyerm. * NC 2187; P. rotundata Kunth * NC 1861; P. tetraphylla (G. Frost.) Hook. & Arn. *
NC 1467; Piper aduncum L. var. cordulatum (C. DC.) Yunck. NC 1183; P. aequale Vahl * NC
1450; P. dilatatum L. C. Rich. * AL 334; P. hispidum Sw. NC 2567; P. longispicum C. DC. var.
glabratum (Yunck.) Steyerm. NC 1538, 2174; P. phytolaccifolium Opiz NC 1735; P. sp. NC
2019; P. sp. Liana NC 2144, 2214; P. veraguense C. DC. NC 2242. PLANTAGINACEAE:
Plantago australis Lam * NC 1402. POLYGALECEAE: Bredemeyera sp. NC 2267; Monnina
meridensis Planch. & Lindl. ex Wedd. NC 1221; M. smithii Chodat * AL 282; M. sp. NC 2825.
POLYGONACEAE: Coccoloba cf. llewelynii R.A. Howard NC 1906, 2344, 2623; Coccoloba
sp. NC 2516. PROTEACAE: Panopsis sp. NC 2616; Panopsis suaveolens (H. Karst.) Pittier NC
1911, 2081; Roupala barnettiae Dorr NC 2637. RANUNCULACEAE Clematis guadeloupae
Pers * NC 1757. RHAMNACEAE: Rhamnus sphaerosperma Sw. var. polymorpha (Reiss.) M.C.
Johnst NC 2441, 2395. ROSACEAE: Hesperomeles obtusifolia (Pers.) Lindl. NC 2685;
Hesperomeles sp. NC 3080; Prunus moritziana Koehne NC 1245,1314, 1650, 1915, 1990, 2031,
2100, 2352, 2485, 2597, 2619, 2639. RUBIACEAE: Arachnothrix reflexa (Benth.) Planchon *
AL 301; Borreria laevis (Lam.) Griseb. * AL 332; Coffea arabica L. NC 1561; Coussarea
moritziana (Benth.) Standl. NC 935, 1574, 1767, 1986; Dioicodendron dioicum (K. Schum. &
Krause) Steyerm. NC 1135, 2095, 2418; Elaeagia karstenii Standl. NC 1590, 2362; E. myriantha
(Standl.) Hammel & C. M. Taylor NC 1192; E. ruiz-teranii Steyerm. NC 921, 991, 1644, 1918,
2009, 2237; Faramea guaramacalensis Taylor NC 1297, 1623, 1699, 2364, 2607, 2635; F.
killipii Standl. NC 938, 1030; Guettarda crispiflora Vahl subsp. discolor (Rusby) Steyerm. NC
212
Appendix
_______________________________________________________
1007, 2055, 2216; Hillia parasitica Jacq. * NC 1389; Hippotis albiflora H. Karst. NC 1733,
2554; Hoffmannia pauciflora Standl. NC 2167; Ladenbergia cf. buntingii Steyerm. NC 1842;
Manettia moritziana (Schum.) Wernham * NC 2491; Notopleura patria (Standl. & Steyerm.)
C.M. Taylor * NC 1462; N. steyermarkiana C.M. Taylor * NC 1094; Palicourea angustifolia
Kunth NC 1239, 2240, 2602; P. apicata Kunth NC 1894, 2339, 2609; P. demissa Standl. NC
994, 1109, 1184, 2049; P. jahnii Standl. NC 2830; P. petiolaris Kunth * NC 2549; P.
puberulenta Steyerm. NC 917; Posoqueria coriacea M. Mart. & Galeotti subsp. formosa NC
1187, 1559, 1740, 2132; Psychotria amita Stand. NC 2217; P. aubletiana Steyerm. * NC 1095;
P. dunstervilleorum Steyerm. * NC 2807; P. fortuita Standl. NC 1172, 2160, 2235; P. lindenii
Standley * NC 1448; NC 1554; P. longirostris (Rusby) Standl. NC 1200, 1794; P. macrophylla
Ruiz & Pav. * NC 1483; P. molliramis (Schum. & Kr.) Steyerm. * NC 1403; P. trichotoma Mart.
& Gal. NC 1849, 2560; Randia cf. dioica H. Karst. NC 1782; Rudgea nebulicola Steyerm. NC
1794, 2177, 2556; R. tayloriae Aymard, Dorr & Cuello NC 1331, 2332, 2347, 2358, 2630; Simira
erythroxylon (Willd.) Brem. var. meridensis Steyerm. NC 1176, 1833, 2507; S. lezamae Steyerm.
NC 1979, 2312; Tammsia anomala Karst. NC 2533; Tocoyena costanensis Steyerm. subsp.
andina Steyerm. NC 1777,1827. RUTACEAE: Conchocarpus larensis (Tamayo & Croizat)
Kallunki & Pirani NC 1451; Zanthoxylum acuminatum (Sw.) Sw subsp. juniperinum (Poepp.)
Reynel NC 1230; Z. melanostictum Schltdl. & Cham. NC 1687, 2411. SABIACEAE: Meliosma
meridensis Lasser NC 1086; Meliosma pittierana Steyerm. NC 1562, 1804, 2052; M. tachirensis
Steyerm. & Gentry NC 1080, 1250, 2243, 2591; M. venezuelensis Steyerm. NC 1354, 2112.
SAPINDACEAE: Allophylus cf. glabratus (Kunth) Radlk NC 2018; Billia columbiana Planch. &
Lindl. ex Triana & Planch. NC 1583, 2172; Cupania cf. scrobiculata Rich. NC 1829; Matayba
camptoneura Radlk. NC 1544, 2566, 2593; Paullinia capreolata (Aubl.) Radlk. NC 1856; P. cf.
latifolia Benth. ex Radlk NC 1823, 2563. SAPOTACEAE: cf. Elaeoluma nuda (Baehni) Aubr.
NC 1087, 1671; Chrysophyllum cf. cainito L. NC 939, 950, 1542, 2258, 2337; C. sp. NC 2239;
Pouteria baehniana Monachino NC 1501, 2146. SCROPHULARIACEAE: Sibthorpia repens
(Mutis ex L.) Kuntze * NC 2801. SMILACACEAE: Smilax kunthii Killip & C. V. Morton NC
2071. SOLANACEAE: Browallia americana L. * NC 1434; Cestrum bigibbosum Pittier NC
2163, 2233; C. buxifolium Kunth NC 2457; C. darcyanum Benitez & N.W. Sawyer NC 1053,
2056; Cuatresia riparia (Kunth.) Hunz NC 2149, 2158, 2529, 2562; Deprea paneroi Benitez &
Martinez * NC 1218; Lycianthes radiata (Sendtn.) Bitter * NC 1212; Markea sp. NC 2057;
Solanum acerifolium Dunal * AL 330; S. aturense Humb. & Bonpl. ex Dunal NC 2034; S.
confine Dunal NC 995, 1024,1411; S. nudum Dunal NC 2130, 2249, 2234; S. pentaphyllum Bitter
* NC 2496; S. torvun Sw. * AL 322. STAPHYLEACEAE: Huertea glandulosa Ruiz & Pav. NC
1194, 1837; Turpinia occidentalis (Sw.) G. Don. NC 2024, 2246. SYMPLOCACEAE:
Symplocos bogotensis Brand. NC 1129, 2059; Symplocos tamana Steyerm. NC 1957.
THEACEAE: Freziera serrata A. L. Weitzman, ined. NC 2459; Gordonia fruticosa (Schrader)
H. Keng NC 1191, 1576; Ternstroemia acrodantha Kobuski & Steyerm. NC 1154; T. sp. A NC
1917; T. sp.B NC 1973. THYMELAEACEAE: Daphnopsis sp.* NC 1714; Schoenobiblus
suffruticosa Barringer & Nevling, vel aff. * NC 1484. URTICACEAE: Pilea A * NC 1470; Pilea
B * NC 1399; P. C * NC 1408; P. rhombea (L.f.) Liebm. * NC 1436; Urera caracasana (Jacq.)
Griseb. NC 1175, 2531. VERBENACEAE: Aegiphila floribunda Moritz & Moldenke NC 1170,
1853, 2576; Aegiphila moldenkeana Lopez-Pal. NC 1243; A. ternifolia (Kunth) Moldenke NC
1214, 2028; Citharexylum venezuelense Mol. NC 2539; Petrea pubescens Turcz. NC 1502;
Verbenaceae indet. NC 1532. VITACEAE: Cissus trianae Planch NC 1872. WINTERACEAE:
Drimys granadensis L.f. NC 1118, 1947, 2484.
Liliopsida: ARACEAE: Anthurium bernardii Croat. * AL 688; Anthurium eminens Schott NC
2580; A. gehrigeri Croat. * NC 1469, 1606; A. ginesii Croat NC 2465; A. humboldtianum Kunth
NC 2435; A. nymphaeifolium K. Koch & Bouche NC 1890; A. scandens (Aubl.) Engl. * NC
1469, 2307; A. smargdianum Bunting NC 2159; Philodendron fraternum Schott * AL 238.
ARECACEAE: Aiphanes stergiosii Niño, Dorr & Stauffer NC 1868, 2592; Chamaedorea
pinnatifrons (Jacq.) Oersted * AL 288; Geonoma jussieuana Mart. NC 1651, 1901, 2105; G.
orbigniana Mart. NC 1950, 2244; G. undata Klotzsch NC 1613, 1731, 2193, 2509; Prestoea
acuminata (Willd.) H.E. Moore var. acuminata NC 1282, 2428; Wettinia praemorsa (Willd.)
Wess. Boer NC 1275. BROMELIACEAE: Greigia albo-rosea (Griseb) Mez NC 2080; Pitcairnia
brevicalycina Mez. * NC 1359; Racinaea sp.* NC 2425. COMMELINACEAE: Tradescantia
213
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
zanonia (L.) Sw. * AL 292. CYCLANTHACEAE: Asplundia vagans Harling NC 1803;
Sphaeradenia laucheana (Mast.) Harling NC 1509, 1898, 2192, 2416, 2561. CYPERACEAE:
Carex jamesonii Boot, s.l. * NC 2819; Rhynchospora immensa Kük. * NC 1159; R. tuerckheimii
* NC 1160;Uncinia hamata * NC 2845. DIOSCOREACEAE: Dioscorea lisae Dorr & Stergios *
NC 2433. HELICONIACEAE: E Heliconia hirsuta L. F. * NC 1747, 2548; H. meridensis
Klotzsch * NC 1481. ORCHIDACEAE: Anathallis sclerophylla (Lindl.) Pridgeen & M. W.
Chase * NC 2306; Brachionidium tuberculatum Lindl. * NC 2823; Cirtochilum megalophium
(Lind.) Känzl. * NC 2794; Corymborkis flava (Sw.) Kuntze * NC 2155; Dichaea camaridioides
Schlechter * NC 1466; Elleanthus flavescens (Lindl.) Rchb.f. * NC 1430; Epidendrum
cereiflorum Garay & Dunst. * AL 689; E. unguiculatum (C. Schweinf.) Garay & Dunst. * NC
2185; Gomphichis costaricensis (Schltr). Ames, F. T. Hubbard & C. Schweinf. * NC 2754;
Jacquiniella teretifolia (Sw.) Britton & P. Wilson * NC 1732; Malaxis licatae Carnevali &
Ramirez * AL 218; M. nidiae Carnevali & Ramirez * NC 2372; Maxillaria nigrescens Lindl. *
NC 2184; Odontoglossum crocidipterum (Rchb. f.) * AL 230; O. schillerianum Rchb.f. * NC
2817; Ornithidium ruberrimum Reichb. F. * NC 2322; Pachyphyllum sp. * NC 2760; Platystele
pisifera (Lindl.) Luer. * NC 2374; Pleurothallis biserrula Rchb.f. * NC 2279; P. bivalvis Lindl. *
NC 1413; P. calamifolia Luer y R. Escobar R. * NC 2327; P. semiscabra Lindl. * NC 2308,
2328; Scaphyglottis summersii L.O. Williams * NC 2269; Scelochilus ottonis Kl. * AL 231;
Sobralia sp.* NC 2310; Stelis chamaestelis (Rchb.f.) Garay & Dunst. * NC 1604; S. oblonga
Willd. * AL 329; S. vulcanii Rchb.f. * AL 327; Trichocentrum pulchrum Poepp. & Endl. * NC
1528. POACEAE: Arthrostylidium venezuelae (Steud.) McClure * NC 2426; 2426; Chusquea
angustifolia (Soderstr. & C.E. Calderon) L. G. Clark NC 2757, 2884; Ch. purdieana Munro NC
1240; Ichnanthus nemorosus (Sw.) Doll * NC 2503; Muhlenbergia diversiglumis Trin. * NC
2551; Rhipidocladum geminatum (McClure) McClure 2466. INDETS. bejuco NC 2178; Indet.
Liana NC s/n.
214
Appendix
_______________________________________________________
Appendix 2. Location of montane forest plots in Ramal de Guaramacal, Andes, Venezuela.
(*) Indicates plots selected for vegetation profiles. (1) Indicates areas outside
Guaramacal National Park borders.
No.
Altitude
Plot
size
Slope
Park Sector
Plot
m
m2
1*
1960
1000
N
Guaramacal
2*
2100
1000
N
Guaramacal
3
2300
1000
N
Guaramacal
4*
2400
1000
N
5*
1850
1000
N
6*
2470
1000
7
1950
1000
8
2300
9
10*
Date of survey Geograp. Pos. UTM Zone
19
East
North
May. 1995
365900
1022968
Jun 1995
366824
1022883
Jun 1995
367166
1022341
Guaramacal
Jun 1995
367452
1022406
Guaramacal
Jun. 1995
365980
1022056
S
Guaramacal
Dec 1995
371908
1019855
S
Guaramacal
Jan 1996
372280
1018020
1000
S
Guaramacal
Dec 1995
371741
1019544
2100
1000
S
Guaramacal
Dec 1995
372345
1018661
1600
1000
S
Agua Fría (El Alto)
Mar 1999
381199
1031364
11
1800
1000
S
Agua Fría (El Alto)
Mar 1999
381241
1031827
12
1950
1000
S
Agua Fría (El Alto)
May 1999
381390
1032534
13
1550
1000
S
Agua Fría (La
Divisoria)
Dec. 1999
380077
1027005
14
1830
1000
N
Agua Fría (El Mogote1)
Dec. 1999
382173
1033526
15
1880
1000
S
Agua Fría (La
Divisoria)
Jan 2000
379322
1027005
16
2580
1000
N
Guaramacal
Mar 2000
368580
1022299
17
2480
1000
N
Guaramacal
Feb 2000
368011
1022672
18
2170
1000
N
Guaramacal
Mar 2000
367020
1022769
19
2070
1000
N
Guaramacal
Feb. 2000
366542
1022989
20
2350
1000
N
Guaramacal
Mar 2000
367166
1022341
21*
1880
1000
NO
Guaramacal (El
Santuario)
Nov. 2001
359185
1012298
22
2100
1000
NO
Guaramacal (El
Santuario)
Mar 2002
359318
1013285
23
2250
1000
NO
Guaramacal (El
Santuario)
Aug 2002
359120
1013013
24
2580
1000
S
Guaramacal
Jan 2002
371718
1022127
25
1900
1000
N
Agua Fría (Laguna
Negra)
Apr 2002
371016
1028548
26
2100
1000
N
Agua Fría (Laguna
Negra)
Apr. 2002
371722
1028117
215
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
No.
Altitude
Plot
size
Slope
Park Sector
Date of survey Geograp. Pos. UTM Zone
19
Plot
m
m2
27
2260
1000
N
Agua Fría (Laguna
Negra)
28
1330
1000
S
Agua Fría (Río Frío1)
1
East
North
Jun. 2002
371703
1027261
Nov 2002
378834
1025276
29
1450
1000
S
Agua Fría (Río Frío )
Dec. 2002
378380
1024991
30
1875
1000
S
Agua Fría (La Peña)
Jan 2003
376567
1023983
31
1770
1000
S
Agua Fría (La Peña)
Jan 2003
376797
1023835
32
2125
1000
S
Agua Fría (La Peña)
Feb. 2003
375862
1024743
33
2474
300
N
Agua Fría (Laguna
Negra)
Apr. 2003
371661
1026420
34*
3050
200
N
Guaramacal
May. 2004
369585
1020286
35
2890
1000
N
Pumar
Feb. 2005
368063
1018997
36
2870
400
N
Pumar
Mar. 2005
368111
1018943
37*
2870
1000
N
Pumar
Mar. 2005
368024
1018958
38*
2810
200
N
Guaramacal
Apr. 2005
369881
1021648
39
2750
1000
N
Guaramacal
Apr. 2005
369545
1021382
40
2950
200
S
Guaramacal
Apr. 2005
370212
1020519
41
2950
400
S
Guaramacal
May. 2005
370209
1020530
PL3
2830
100
N
Guaramacal
Dec. 2004
369784
1021281
43
3060
50
S
Guaramacal
Dec, 2006
369951
1020437
44
3050
100
N
Guaramacal
Jan. 2007
369474
1020570
216
Appendix
_______________________________________________________
Appendix 3. Morphological and chemical characteristics of some soil profiles representative
from Ramal de Guaramacal (from Marvez & Schargel 1999). ls: loamy sand,
sl: sandy loam, l: loam, scl: sandy clay loam, cl: clay loam; c: clay
4-H2
5-H1
2-H2
3-H1
37-48
7.5YR5/1
48-56
7.5YR4/1
15 2.450
5
7
56-70
> 70
0-15
1.950 15-30
> 30
0-10
10-20
20-50
50-70
2.100
70-100
100-115
5YR2.5/2
7.5YR4/2
10YR4/6
5YR3/3
5YR4/6
7.5YR4/6
7.5YR5/1
7.5YR5/2
10YR4/6
10YR3/3
7.5YR4/4
10YR4/2
7.5YR5/3
7.5YR6/4
7.5YR5/6
10YR6/4
7.5YR6/2
MARNR48
115-150 7.5YR6/4
60 1.820
0-10
-25
-45
-73
-102
-125
-150
10YR2/2
10YR3/2
7.5YR5/8
7.5YR5/8
7.5YR5/8
7.5YR5/8
7.5YR5/8
6,2 3,8 1,2(*)
Rock
32,5 5,6 65,8
Limestone boulders 0,5-1m
sl
10
5,5 5,3 16,9
l
3,5 6,7 8,2
Fractured rock
sl
20
7,5 4,0 1,6
sl
30
6,2 4,4 0,9
sl
50
3,0 4,4 1,1
Fractured rock
ls
6,5 3,8 1,1
sl
1,4 4,1 1,1
7.5YR
l
2,0 4,2 1,3
4/4-5/8
7.5YR
l
3,2 4,3 0,8
4/4-5/8
10YR4/3 scl
4,2 4,5 1,3
Fractured rock
sl
5
5,5 3,5 1,2
scl
15
3,5 3,7 0,8
Fractured rock
l
6,2 4,7 3,6
l
4,4 4,5 2,0
2.5YR6/4 cl
3,0 4,4 1,3
7.5YR6/2 cl
1,0 4,4 1,0
7.5YR6/2 scl
0,4 4,3 0,9
7.5YR6/8 sl
0,4 4,5 1,1
7.5YR7/1
sl
0,4 4,5 1,2
2.5YR4/6
sl
2
15,6 4,2
10YR5/4 cl
2
9,6 4,1
c
2
4,0 4,1
c
5
1,6 3,8
c
10
1,2 4,1
c
15
0,5 4,2
c
25
0,3 4,0
-
Exchangeable
Aluminum
me/100gr
ls
Sum Bases
me/100gr
-
pH
7.5YR3/1
Orgánic matter
(%)
0-14
> 14
0-10
30 1.850
> 10
0-18
40 1.850 18-60
> 60
0-20
20-40
20 2.300
40-100
> 100
0-25
25-37
<5 1.950
% of Coarse
fragments
Depth of layer
(cm)
Elevation (m)
Texture
1-H2
Color * *
(Mottling)
1-H4
Color **
(moist)
2-H1
Slope (%)
Profile
North Slope
0,3
1,4
0,8
0,4
1,0
0,6
1,0
2,0
0,8
2,3
3,2
0,3
1,0
1,6
2,1
1,6
1,7
1,0
-
217
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Appendix 3. Cont.
MARNR 54
*
8-H1
40-50
> 50
0-20
20-35
40 2.450
35-90
> 90
0-15
15-27
15 2.300 27-50
50-80
80-110
0-9
9-26
26-45
45-62
62-81
45 1.670 81-94
94-97
97-99
99-111
111-191
191-195
4,6
2,0
0,3
4,2
4,1
4,1
2,5
1,3
0,9
0,7
0,5
0,4
4,8
4,8
4,8
1,1
1,2
1,2
0,4
0,6
0,6
3,8
4,1
4,2
4,6
4,5
4,0
3,8
3,8
4,2
4,6
4,5
4,3
4,7
4,6
-
1,0
1,4
0,9
1,0
1,0
0,4
0,2
0,2
0,1
t
t
t
t
t
-
0,9
1,0
1,1
1,8
0,6
0,4
0,8
0,6
0,3
0,2
1,0
1,1
6,8
2,9
1,8
4,9
4,8
4,8
1,3
0,9
1,2
0,2
1,3
1,0
5,6
1,7
1,4
4,6
4,5
4,7
0,5
0,2
0,2
1,9
2,5
1,8
Orgánic
matter (%)
Texture
% of Coarse
fragments
Color * *
(Mottling)
ls
Exchangeable
Aluminum
me/100gr
7-H2
35 2.100
-
Sum Bases
me/100gr
9-H1
7.5YR4/1
pH
10-H1 45 1.950
0-10
> 10
0-10
10-20
20-40
> 40
0-18
18-40
Color **
(moist)
10-H2 45 1.950
Depth of layer
(cm)
Elevation (m)
Slope (%)
Profile
South Slope
2,2
Rock
7.5YR4/1
ls
10
4,2
5YR4/1
ls
40
3,9
5YR5/1
ls
50
2,8
Fractured rock
10YR4/1
l
9,6
10YR5/1 7.5 YR5/6 l
3,4
5YR6/27.5 YR5/6 l
2,2
7/1
Fractured rock
5YR4/2
sl
50
4,7
7.5YR6/6
sl
60
4,4
7.5YR6/8
sl
60
3,4
Fractured rock
5YR4/2
l
20
6,9
7.5YR6/8 5YR5/2
l
25
4,8
7.5YR6/8 7.5YR6/1 l
30
1,8
7.5YR6/8
scl
50
1,0
7.5YR6/6
scl
30
0,9
10YR2/2
scl
2
4,6
10YR4/2
sl
20
2,9
10YR4/1
ls
20
2,4
10YR7/1
ls
40
0,5
10YR7/1
ls
20
0,1
10YR7/1
ls
20
0,1
10YR5/2
sl
20
0,4
Plácico
10YR6/4 10YR7/1 sl
20
0,1
10YR7/1
sl
20
0
Placic
-
4,5
4,0
4,2
0,8
1,2
-
Páramo
MARNR
57 *
6-H2 >60 3.100
30 2.830
0-23
23-38
38-60
7.5YR4/1
7.5YR4/4 7.5YR5/1
5YR5/6 5YR5/2
sl
l
l
> 60
0-15
15-28
28-56
10YR4/1
5YR7/1 5YR5/8
7.5YR6/6 10YR7/3
sl
l
scl
>56
* Soils with placic horizons
** Munsell color chart
218
Rock
-
Rock. Placic horizon at 40 cm
Appendix
_______________________________________________________
Appendix 4. Checklist and vouchers of all plant species diversity recorded from the studies
of páramo vegetation of Ramal de Guaramacal (including both zonal and
azonal vegetation). All vouchers numbers correspond to N. Cuello et al.
Lichenized Ascomycetes. BAEOMYCETACEAE: Phyllobaeis imbricata (Hook. in Kunth) Kalb
& Gierl 3173. CLADONIACEAE: Cladia aggregata (Sw.) Nyl. 2966, 2977, 3067, 3148;
Cladina arcuata (Ahti) Ahti & Follmann 2973; cf. Cladina rangiferina (L.)Nyl. 2973; Cladonia
andesita Vain. 2971; Cladonia cf. pyxidata (L.) Hoffm. 2972; Cladonia co-rymbites Nyl. 3195;
Cladonia crispata (Ach.) Flot. 3146; Cladonia didyma (Fee) Vain 3146; Cladonia furcata
(Huds.) Schrad. 3036, 3049; Cladonia isabellina Vain. 2975; Cla-donia squamosa (Scop.)
Hoffm. 2964, Cladonia sp. 3068. ICMADOPHILACEAE: Siphula pteruloides Nyl.3145.
PARMELIACEAE: Rimelia reticulata (Tayl.) Hale & Fletcher 3015, 3172; Usnea sp.
3025.PELTIGERACEAE: Peltigera neopolydactyla (Gyeln.) Gyeln. 3162. PERTUSARIACEAE: Pertusaria sp. 3165. STEREOCAULACEAE: Stereocaulon didymi-cum Lamb 3195.
cf. Stereocaulon microcarpum Müll.Arg. 3024.
Hepaticae. ANEURACEAE: Riccardia spp. 2955, 2965. HERBERTACEAE: Herbertus pensilis
(Tayl.) Spruce 2986; Herbertus grossispinus (Steph.) Fulf. 3129, 3236, 2950; Her-bertus
juniperoideus (Sw.) Grolle 3078, 3149, 3270, 2980; Herbertus acanthelius Spruce 3242, 3119;
Herbertus subdentatus (Steph.) Fulf. 3246. LEPIDOZIACEAE: Bazzania latidens (Gottsche ex
Stephani) Fulford 3257, 3258; Bazzania spp. 3066, 3136. METZGE-RIACEAE: Metzgeria sp
3250. PLAGIOCHILACEAE: Plagiochila cf aerea Taylor 3126; Plagiochila tabinensis Steph.
3071; Plagiochila spp. 2948, 2952, 2957, 2958, 2961, 2962, 2969, 2979, 2982, 2985, 3042, 3132,
3163, 3239. SCAPANIACEAE: Scapania porto-ricensis Hampe & Gottsche 3130, 2967, 3073,
3124, 3159. JUNGERMANIACEAE: Jamesoniella rubricaulis (Nees) Grolle 2949, 2959, 2963,
3021. JUBULACEAE: Frullania spp. 2970, 2976, 3022, 3038, 3039. Hepaticae indet. 2979,
3132.
Musci. BARTRAMIACEAE: Breutelia rhythidoides Herz. 2978, 3035, 3075, 3099; Breu-telia
squarrosa Jaeg. 2954, 2960, 3030, 3110. BRYACEAE: Bryum grandifolium (Tayl.) C. Muell.,
2946, 3120.DICRANACEAE: Campylopus albidovirens Herz. 3100, 3101, 3107, 3109;
Campylopus cuspidatus (Hsch.) Mitt. var. dicnemioides (C. Muell.) J.-P. Frahm 3105, 3108;
Campylopus flexuosus (Hedw.) Brid. 3065; Campylopus fragilis (Brid.) B.S.G. 3048;
Campylopus insignis Herz., 2974 Campylopus nivalis (Brid.) Brid. 3072, 3128; Campylopus
pilifer Brid. 3033, 3122; Campylopus richardii Brid. 3016, 3079;Campylopus subjugorum Herz.
3031, 3046, 3076; Campylopus trichophorus Hampe ex Herz., 2983, 3050; Dicranum frigidum C.
Muell., 3118. LEUCOBRYACEAE: Leucobryum antillarum Schimp. ex Besch., 3055.
POLYTRICHACEAE: Polytrichadelphus longisetus (Hook.) Mitt., 3020; Polytrichum commune
Hedw., 3098; Polytrichum juniperinum Hedw., 3103, 3117. POTTIACEAE: Leptodontium
viticulosoides (P. Beauv.) Wijk & Marg. var. panamense (Lor.) Zander, 3054. RHACOCARPACEAE: Rhacocarpus purpurascens (Brid.) Par., 3017. SEMATHOPHYLLACEAE:
Sematophyllum swartzii (Schwaegr.) Welch & Crum, 3018; Semathophyllum sp. 3077.
SPLACHNOBRYACEAE: Tetraplodon mnioides (Hedw.) B.S.G., 3121. SPHAGNACEAE:
Sphagnum cuspidatum Ehrhart ex Hoffm. 3051, 3089, 3113; S. magellanicum Brid. 3041, 3104,
3115; Sphagnum meridense (Hpe.) C. Muell. 3074; Sphagnum recurvum P. Beauv. 3106;
Sphagnum sancto-josephense Crum & Crosby; Sphagnum sparsum Hpe., 3114.
Ferns. BLECHNACEAE: Blechnum schomburgkii (Kl.) C. Chr. 2937, 2940. DRYOPTERIDACEAE: Elaphoglossum appressum Mickel 2936; Elaphoglossum cf. lingua (C. Presl)
Brack. 2893. GRAMMITIDACEAE: Lellingeria myosuroides (Sw.) A.R. Sm. & R.C. Moran
2933; Melpomene flabeliformis (Lag. Ex Sw.) A.R. Sm & R.C. Moran 2732, 2877; M.
moniliformis (Lag. Ex Sw.) A.R. Sm & R.C. Moran 2720; M. xiphopteroides (Liebm.) A.R. Sm.
2896; Melpomene sp. 3157. HYMENOPHYLLACEAE: Hymenophyllum myrio-carpum Hook.;
H. trichomanoides Bosch. 2728, 2895; Hymenophyllum sp. ISOETACEAE: Isoetes karstenii A.
Braun 3367, 3386; LYCOPODIACEAE: Huperzia amentacea (B.Øllg.) Holub.; H. riobanbensis
(Herter) B.Øllg. 2723; Lycopodium clavatum subsp. contiguum Kl. 2696. OPHIOGLOSSA-
219
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
CEAE: Ophioglossum crotalophorioides Walter 2931. POLYPODIACEAE: Polypodium funckii
Mett. 2809; Polypodium sp. 3070. PTERIDACEAE: Eriosorus flexuosus (Kunth) Copel var.
flexuosus 2824, 2897, 2934; Jamesonia imbricata (Sw.) Hook. & Grev. 2722.
Magnoliopsida. APIACEAE: Daucus montanus H. & B. ex Spreng. 2997. AQUIFOLIACEAE:
Ilex guaramacalensis Cuello & Aymard 2701, 2942. ASTERACEAE: Ageratina theifolia
(Benth.) R.M. King & H. Rob. 2746; Diplostephium obtusum S.F. Blake 2748; D. venezuelense
Cuatrec. 2688, 3096; Hieracium avilae Kunth 2890; Libanothamnus griffinii (Ruiz-Teran &
Lopez-Fig.) Cuatrec. 2704; Pentacalia cachacoensis (Cuatrec.) Cuatrec. 2694; P. greenmaniana
(Hieron.) Cuatrec. 2747; Ruilopezia jabonensis (Cuatrec.) Cuatrec. 2695, 2715; R. lopez-palacii
(Ruiz-Teran & Lopez-Figueiras) Cuatrec. 2899; R. paltonioides (Standl.) Cuatrec. 2716; R.
viridis (Aristeguieta) Cuatr. 2717; Asteraceae indet. 3299. CARYOPHYLLACEAE: Arenaria
venezuelana Briq. 3082. CLUSIACEAE: Hype-ricum cardonae Cuatrec. 2700; H. juniperinum
Kunth; H. paramitanum N. Robson; H. sp. ERICACEAE: Bejaria aestuans L. Disterigma
acuminatum (Kunth) Nied. D. alaternoides (kunth) Nied. 2730; Gaultheria anastomosans (L.f.)
Kunth 2724, 2752; G. erecta Vent. 3058; G. hapalotricha A.C. Sm. 2714; Pernettya prostrata
(Cav.) DC. 2706, 2713, 2727; Sphyrospermum buxifolium Poepp. & Endl. 3321; Themistoclesia
dependens (Benth.) A.C. Sm. 2733; Vaccinium corymbodendron Dunal 2708, 2729.
GENTIANIACEAE: Gentianella nevadensis (Gilg.) Weaver & Rudenberg. 2915. GERANIACEAE: Geranium stoloniferum Standl. 2913. LENTIBULARIACEAE: Utricularia alpina Jacq.
2901. MELASTOMATACEAE: Chaetolepis lindeniana (Naud.) Triana 2705; Miconia tinifolia
Naud. 2702, 3064; Monochateum discolor H. Karst. 2710, 2750. MYRSINACEAE: Cybianthus
laurifolius (Mez) G. Agostini; C. marginatus (Benth.) Pipoly; C. stapfii (Mez) Agostini 2703;
Myrsine dependens (Ruiz & Pav.) Spreng. 2792; 2796. MYRTACEAE: Ugni myricoides
(Kunth.) O. Berg. 2721. POLYGALACEAE: Monnina sp. 3061. POLYGONACEAE: Muehlenbeckia thamnifolia (Kunth) Meisn. 2731. ROSACEAE: Hesperomeles obtusifolia (Pers.) Lind.
var. obtusifolia 2707; Hesperomeles sp. 3080; Lachemilla verticillata (Fielding & Gardner)
Rothn. 3116.; Rubus acanthophyllos Focke 3060. RUBIACEAE: Arcytophyllum nitidum (Kunth)
Schltdl.2718; Galium hypocarpium (L.) Endl. Ex Griseb.; Nertera granadensis (Mutis ex L.f.)
Druce 2734; Palicourea jahnii Standl. 2943; SCHROPHULARIACEAE: Castilleja fissifolia L.f.
2902; VALERIANACEAE: Valeriana quirorana Xena 2935, 3322.
Liliopsida. BROMELIACEAE: Greigia sp.; Puya aristeguietae L.B. Sm.; Puya sp.;
CYPERACEAE: Carex bonplandii Kunth 2741; Eleocharis acicularis (L.) Roem. & Schult.
3088, 3368; Eleocharis sp. 3174; Oreobolus venezuelensis Steyerm. 2891; Rhynchospora
gollmeri Boeck. 2739; R. guaramacalensis M. Strong 2889; Rhynchospora macrochaeta Steud.
Ex Boeck. 2697, 2740, 2743.; Rhynchospora spp.2697, 2735. ERIOCAULACEAE: Paepalanthus pilosus (Kunth) Kunth 2996, 3097; IRIDACEAE: Orthrosanthus acorifolius (Kunth)
Ravenna 2904; Sisyrrhinchium tinctorium Kunth 2911. Sisyrhynchium sp. 3391. ORCHIDACEAE: Cyrtochilum ramosissimum (Rchb.f.) Kränl. 3001; Epidendrum frutex Rchb.f. 2719;
Pterichis multiflora (Lindl.) Schltr. 2916. POACEAE: Agrostis basalis Luces. 2803, 3085, 3389;
A. perennans (Walter) Tucker 2917, 3189, 3084, 3092; Agrostis sp. B 3155, 3093; Calamagrostis
bogotensis (Pilg.) Pilg.; C. planifolia (Kunth) Trin. Ex Steud. 2905, 2914, 2925, 29991;
Calamagrostis sp. A, 2926; Calamagrostis sp. 3388; Chusquea angustifolia (Sodestr. & C.E.
Calderon) L.G. Clark 2941, 2995; Ch. tessellata Munro 3153; Cortaderia hapalotricha (Pilg.)
Conert. 2737; Festuca guaramacalana Stancik 2900; Neurolepis glomerata Swallen 2726;
Ortachne erectifolia (Swallen) Clayton 3390; Polypogon elongatus Kunth 2990; Indets. 3090,
3142. TOFIELDIACEAE: Isidrogalvia robustior (Steyerm.) Cruden. XYRIDACEAE: Xyris
subulata Ruiz & Pav.var. acutifolia Heimerl. 2699, 2738.
220
Appendix
_______________________________________________________
Appendix 5. Species list from páramo areas [zonal (Pzo) and azonal (Paz) páramo
vegetation, including, subpáramo (SP) and páramo-connected dwarf
forest (SARF) vegetation islands] present in the summit of Ramal de
Guaramacal in the Venzuelan Andes. Species distribution group (1-10.4)
as presented in Table 5.4, Group 0 for unknown distribution. Introduced
species indicated with asterisk (*).
FAMILY/SPECIES
ASPLENIACEAE
Asplenium serra Langsd. &
Fisch.
BLECHNACEAE
Blechnum aff. atropurpureum
A.R. Sm.
B. auratum (Fee) R.M. Tryon &
Stolze
B. binervatum (Poir) C.V.
Morton subsp. fragile (Desv).
R.M. Tryon & Stolze
B. schomburgkii (Klotzsch) C.
Chr.
CYATHEACEAE
Cyathea fulva (Mart. & Gal.)
Fee
DENNSTAEDTIACEAE
Histiopteris incisa (Thunb.) J.
Sm.
Paesia acclivis (Kunze) Kuhn
DICKSONIACEAE
Culcita coniifolia (Hook.)
Maxon
DRYOPTERIDACEAE
Diplazium hians Kunze ex
Klotzsch
Elaphoglossum andicola (Fee)
T. Moore
E. appressum Mickel
E. cf. lingua (C. Presl) Brack.
E. cuspidatum (Willd) Moore
E. minutum (Pohl ex Fee) T.
Moore
E. muscosum (Sw.) T. Moore
E. nigrocostatum Mickel
E. paleaceum (Hook. & Grev.)
Sledge
E. papillosum (Baker) H. Christ.
E. rhynchophyllum H. Christ.
EQUISETACEAE
Equisetum bogotense Kunth
GLEICHENIACEAE
Sticherus revolutus (Kunth)
Ching
GRAMMITIDACEAE
Ceradenia intonsa L.E. Bishop,
ined
Cochlidium pumilum L.E.
Bishop
Grammitis leptopoda (C.H.
Wright) Copel.
G. xanthotrichia (Kl.) A.R. Sm.
Lellingeria major (Copel.) A.R.
Sm. & R.C. Moran
Distr.
Group
VEG.
TYPE
2
SARF
10.2
SP
6
SARF/Pzo
2
SARF
5
Pzo/SARF
6
SARF
1
7
Pzo
Pzo/SARF
2
SARF
2
SP/SARF
7
7
4
6
SARF
Pzo
Pzo
SARF
4
4
7
Pzo
Pzo
SARF
4
6
10.2
Pzo
SARF
SARF/Pzo
2
Pzo
4
Pzo
8
SP/Pzo
7
SARF
5
10.2
Pzo
SARF
7
SARF/Pzo
Distr.
FAMILY/SPECIES
Group
L. myosuroides A.R. Sm. & R.C.
Moran
1
Melpomene flabelliformis (Lag.
ex Sw.) A.R. Sm & R.C.
Moran
1
M. moniliformis (Lag. ex Sw.)
A.R. Sm & R.C. Moran
4
M. xiphopteroides (Liebm.) A.R.
Sm.
4
M. sp.
0
Terpsichore cultrata (Bory ex
Willd.) A.R. Sm.
2
T. longisetosa (Hook.) A.R. Sm.
6
T. semihirsuta (Kl.) A.R. Sm.
6
HYMENOPHYLLACEAE
Hymenophyllum aff. apiculatum
Mett. ex Kuhn
5
H. fucoides (Sw.) Sw.
2
H. karstenianum J.W. Sturm
7
H. myriocarpum Hook.
6
H. sp.
0
H. tegularis (Desv.) Proctor &
Lourteig
6
H. trichomanoides Bosch.
2
ISOETACEAE
Isoëtes karstenii A. Braun
9
LYCOPODIACEAE
Huperzia amentacea (B. Øllg.)
Holub
6
H. cf. capellae (Herter) Holub.
7
H. eversa (Poir.) B. Øllg.
6
H. molongensis (Herter) Holub.
7
H. ocanana (Herter) Holub
9
H. riobambensis (Herter) B.
Øllg.
9
H. rufescens Hook. Trevis
7
H. sp.
0
Lycopodiella cernua (L.) Pic.
Serm.
0
L. pendulina (Hook.) B. Øllg.
0
L. riofrioi (Sodiro) B. Øllg.
6
Lycopodium clavatum L. subsp.
contiguum Kl.
6
L. jussiaei Desv. ex Poir.
4
L. thyoides H. & B. ex Willd.
2
OPHIOGLOSSACEAE
Ophioglossum crotalophorioides
Walter
1
PLAGIOGYRIACEAE
Plagiogyria pectinata (Liebm.)
Lellinger
4
POLYPODIACEAE
Campyloneurum amphostenon
(Kunze ex Klotzsch) Fée
4
VEG.
TYPE
SARF/Pzo
Pzo
Pzo
Pzo
Pzo
SARF
SARF
SARF/SP
SARF
SARF
SARF
SARF/Pzo
Pzo
SARF
Pzo
Paz
Pzo
Pzo
Pzo
SARF/Pzo
Pzo
Pzo
Pzo
SARF
SP
SP
B-P
Paz/Pzo
B-P
Pzo
Paz/Pzo
SARF/Pzo
Pzo
221
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Distr.
VEG.
FAMILY/SPECIES
Group
TYPE
Polypodium funckii Mett.
7
Pzo
P. sp.
0
Pzo
PTERIDACEAE
Eriosorus flexuosus (Kunth)
Copel. var. flexuosus
4
SARF/Pzo
Jamesonia imbricata (Sw.)
Hook. & Grev.
7
Pzo
THELYPTERIDACEAE
Thelypteris cheilanthoides
(Kunze) Proctor
4
Pzo
T. frigida (H. Christ) A.R. Sm.
6
Pzo/SP
T. prolatipedis Lellinger
6
SARF
APIACEAE
Daucus montanus H. & B. ex
Spreng.
1
Pzo
Hydrocotyle venezuelensis Rose
ex Mathias
10.1 Pzo/SARF
AQUIFOLIACEAE
Ilex guaramacalensis Cuello &
Aymard
10.4 Pzo/SARF
ARALIACEAE
Oreopanax discolor (Kunth)
Decne. & Planch.
9
SARF
O. sp.1
0
SARF
O. sp.2
0
SARF
ASTERACEAE
Achyrocline moritzianum Klatt
8
Pzo
A. vargasiana DC.
3
SP
Ageratina theifolia (Benth.) R.
M. King & H. Rob.
9
Pzo/SARF
Baccharis prunifolia Kunth
7
SP/SARF
Cotula mexicana L.
2
Paz
Diplostephium obtusum S.F.
Paz/Pzo/SAR
Blake
10.3
F
D. venezuelense Cuatrec.
10.3
Pzo
Gamochaeta americana (Mill.)
Wedd.
1
Pzo
Hieracium avilae Kunth
8
Paz/Pzo
H. erianthum Kunth
7
Pzo
Libanothamnus griffinii (RuízTerán & Lóp. Fig.) Cuatrec.
10.4 Pzo/SARF
Mikania nigropunctulata Hieron 5 LMRF/SARF
M. stuebelii Hieron
7
SARF
Munnozia senecionidis Benth.
6
Pzo
Paragynoxys cuatrecasasii RuízTerán & López-Fig.
0
SARF
Pentacalia cachacoensis
(Cuatrec.) Cuatrec.
9
Pzo/SARF
P. greenmaniana (Hieron.)
Cuatrec.
10.3 Pzo/SARF
P. theaefolia (Benth.) Cuatrec.
7
SARF
P. vicelliptica (Cuatrec.)
Cuatrec.
10.3
SARF
Ruilopezia jabonensis (Cuatrec.)
Cuatrec.
10.3
Paz/Pzo
R. lopez-palacii (Ruíz-Terán &
López-Figueiras) Cuatrec.
10.4
Paz/Pzo
R. paltonioides (Standl.)
Cuatrec.
10.3 Pzo/SARF
R. viridis (Aristeguieta) Cuatrec. 10.4
Pzo
Sonchus oleraceus L.*
BALANOPHORACEAE
Corynaea crassa Hook.f
6
SARF
222
Distr.
VEG.
FAMILY/SPECIES
Group
TYPE
BEGONIACEAE
Begonia formosissima Sandwith 10.3 SARF-Pzo
B. lipolepis L.B. Sm. var.
luteynorum (L.B. Sm. &
Wassh.) Dorr
10.3 SP/SARF
BORAGINACEAE
Cynoglossum amabile Stapf &
J.R. Drumm.
1
Pzo
CAMPANULACEAE
Centropogon aff. elmanus E.
Wimm.
10.3
SARF
C. lanceolatus E. Wimm.
10.3
SP/Pzo
Siphocampylus odontosepalus
Vatke
7
Pzo/SP
CARYOPHYLLACEAE
Arenaria venezuelana Briq.
9
Paz/Pzo
Stellaria cuspidata Willd. ex
Schltdl.
2
SARF
CHLORANTHACEAE
Hedyosmum translucidum
Cuatrec.
7
SARF
CLETHRACEAE
Clethra fagifolia Kunth var.
fagifolia
6 SARF/UMRF
CLUSIACEAE
Hypericum cardonae Cuatrec.
6
Paz/Pzo
H. juniperinum Kunth
9
Paz/Pzo
H. juniperinum x cardonae
10.4
PAz
H. paramitanum N. Robson
10.3 Pzo/SARF
CUNONIACEAE
Weinmannia auriculata D. Don
7
SARF
W. fagaroides Kunth
2
SARF
W. karsteniana Szyszyll.
9
SARF
W. lechleriana Engl.
7
SARF
ERICACEAE
Bejaria aestuans L.
6
Pzo
Cavendishia bracteata (Ruíz &
Pavon ex St.-Hil.) Hoerold
6
Pzo/SARF
Disterigma acuminatum (Kunth)
Nied.
3
Pzo
D. alaternoides (Kunth) Nied.
6
Pzo/SARF
Gaultheria anastomosans (L.f.)
Kunth
6
Pzo/SARF
G. buxifolia Willd.
7
Pzo
G. erecta Vent.
2
Pzo/SARF
G. glomerata (Cav.) Sleum.
7
Pzo
G. hapalotricha A.C. Sm.
7
Pzo
Pernettya prostrata (Cav.) DC.
2
Pzo/Paz
Psammisia hookeriana Klotzsch. 9
SARF
Sphyrospermum buxifolium
Poepp. & Endl.
5
Pzo
Themistoclesia dependens
(Benth.) A. C. Smith
8
Pzo/SARF
Thibaudia floribunda Kunth.
7
SARF
Vaccinium corymbodendron
Dunal
7
Pzo/SARF
GENTIANIACEAE
Gentianella nevadensis (Gilg.)
Weaver & Rudenberg.
8
Paz/Pzo
Halenia sp.
0
Pzo
Macrocarpaea bracteata Ewan 10.3 UMRF/SARF
GERANIACEAE
Geranium stoloniferum Standl. 10.3
Paz/Pzo
Appendix
_______________________________________________________
Distr.
VEG.
FAMILY/SPECIES
Group
TYPE
GESNERIACEAE
Glossoloma chrysanthus (Pl. &
Tr.) J. Clark
10.3
SARF
LENTIBULARIACEAE
Utricularia alpina Jacq.
2
Pzo
LORANTHACEAE
Dendrophtora sp. A.
0
Pzo/SARF
Gaiadendron punctatum (R. &
P.) G. Don
5 UMRF/SARF
Phoradendron sp.
0
Pzo/SARF
MELASTOMATACEAE
Chaetolepis lindeniana (Naudin)
Triana
9
Pzo/SARF
Miconia arbutifolia Naud.
10.1 SARF/SP
M. elvirae Wurdack
10.4
SARF
M. jahnii Pittier
8
SARF
M. tinifolia Naud.
8
Pzo/SARF
Monochaetum discolor H. Karst. 10.3 Pzo/SARF
MYRSINACEAE
Cybianthus laurifolius (Mez)
Agost.
9
SARF
C. marginatus (Benth.) Pipoly
7
Pzo/SARF
C. stapfii (Mez) Agostini
9
SARF
Geissanthus andinus Mez
8
SARF
Myrsine dependens (R. & P.)
Spreng
6
Pzo/SARF
MYRTACEAE
Myrcianthes myrsinoides
(Kunth) Grifo
7
SARF
Ugni myricoides (Kunth.) O.
Berg.
4
Pzo
ONAGRACEAE
Epilobium denticulatum Ruíz &
Pavon
6
Pzo
Fuchsia membranacea Hemsl.
10.3
SARF
OXALIDACEAE
Oxalis sp.
0
Pzo
PHYTOLACCACEAE
Phytolacca rugosa A. Braun &
C.D. Bouche
6
Paz
PIPERACEAE
Peperomia acuminata Ruíz &
Pavon
4
SARF
P. sp. 1
0
SARF
P. sp. 2
0
SARF
PLANTAGINACEAE
Plantago australis L.
2
Pzo
POLYGALACEAE
Monnina meridensis Planch. &
Lind. ex Wedd
10.3
Pzo
M. sp.1
0
SARF
M. sp.2
0
SARF
POLYGONACEAE
Muehlenbeckia tamnifolia
(Kunth) Meisn.
2
Pzo
Rumex acetosella L.*
ROSACEAE
Hesperomeles obtusifolia (Pers.)
Paz/Pzo/SAR
Lindl.
6
F
H. sp.
0
Pzo/SARF
Lachemilla verticillata (Fielding
& Gardner) Rothm.
6
Paz
Rubus acanthophyllos Focke
7
Pzo
Distr.
VEG.
FAMILY/SPECIES
Group
TYPE
RUBIACEAE
Arcytophyllum nitidum (Kunth)
Schltdl.
9
Pzo
Galium hypocarpium (L.) Endl.
ex Griseb.
6
Pzo
Manettia lindenii Sprague
10.3
Pzo
M. moritziana (K. Schum.)
Werham.
10.1
Pzo/SP
Nertera granadensis (Mutis ex
L.f.) Druce
1
Paz/Pzo
Palicourea jahnii Standl.
10.3 Pzo/SARF
Psychotria dunstervilleorum
Steyerm.
10.3
SARF
SCROPHULARIACEAE
Calceolaria tripartita R. & P.
2
Pzo
Castilleja fissifolia L.f.
2
Pzo
Sibthorpia repens (L.) Kuntze
6
SARF/Pzo
SOLANACEAE
Cestrum buxifolium Kunth
9
SARF/SP
Deprea paneroi Benitez et
Martinez
10.4 Pzo/SARF
Solanum macrotonum Bitter
5
Pzo
SYMPLOCACEAE
Symplocos tamana Steyerm.
10.3
SARF
THEACEAE
Freziera serrata A. L.
Weitzman, ined.
10.3 UMRF/SARF
TROPAEOLACEAE
Tropaeolum deckerianum Moritz
& H. Kar
8
Pzo
URTICACEAE
Pilea sp.
0
SARF
VALERIANACEAE
Valeriana quirorana Xena
10.3
Pzo
VIOLACEAE
Viola stipularis Sw.
4
Pzo
WINTERACEAE
Drimys granadensis L.f.
6
SARF
ALSTROEMERIACEAE
Bomarea amilcariana Stergios
& Dorr
10.4
Pzo
B. edulis (Tussac) Herb.
2
SARF
BROMELIACEAE
Greigia sp.
0
Pzo
Guzmania squarrosa (Mez &
Sodiro) L.B. Sm. & Pittdn.
7
Pzo/SARF
Puya aristeguietae L.B. Sm.
8
Pzo
Puya sp. nov.
10.4
Pzo
Tillandsia complanata Benth
4
SP
CYPERACEAE
Carex bonplandii Kunth
1
Paz/Pzo
C. jamesonii Boott
6
Pzo
Eleocharis acicularis (L.) Roem.
& Schult.
1
PAz
Oreobolus venezuelensis
Steyerm.
6
Paz/Pzo
Rhynchospora gollmeri Boeck
10.2
Paz/Pzo
R. guaramacalensis M. Strong
10.4
Pzo
R. cf. lechleri Steud.
5
Pzo
R. macrochaeta Steud. ex
Boeck.
6
Pzo
R. ruiziana Boeck
6
Pzo
R. sp.
0
Pzo
223
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Distr.
FAMILY/SPECIES
Group
ERIOCAULACEAE
Paepalanthus pilosus (Kunth)
Kunth
6
HYPOXIDACEAE
Hypoxis decumbens L.
0
IRIDACEAE
Orthrosanthus acorifolius
(Kunth) Ravenna
9
Sisyrinchium tinctorium Kunth
6
S. sp.
0
JUNCACEAE
Juncus bufonius L.
1
J. stipulatus Nees & Meyen
3
Luzula gigantea Desv.
6
LILIACEAE
Excremis coarctata (Ruíz &
Pav.) Baker
7
ORCHIDACEAE
Brachionidium tuberculatum
Lindl.
7
Cranichis antioquensis Schltr.
6
Elleanthus aurantiacus (Lindl.)
Rchb.f
2
E. flavescens (Lindl.) Rchb.f.
7
E. maculatus (Lindl.) Rchb. f.
6
Epidendrum frutex Rchb.f.
7
E. guaramacalensis Hagsater
10.4
Gomphichis costaricensis
(Schltr.) Ames, F.T. Hubb. &
Schweinf.
6
Odontoglossum megalophium
Lindl.
7
O. ramosissimun Rchb.f.
7
O. schillerianum Rchb.f.
9
Pachyphyllum crystallinum
Lindl.
6
Pleurothallis glossopogon Rchb.
f.
8
Pterichis multiflora (Lindl.)
8
224
VEG.
TYPE
Paz/Pzo
SP
Paz/Pzo
Pzo
Paz
Pzo
Pzo
Pzo
Pzo
SARF
SARF
Pzo
Pzo
Pzo
Pzo
Pzo
SP/SARF
SARF
Pzo
SARF
SARF
SARF
Pzo
Distr.
VEG.
FAMILY/SPECIES
Group
TYPE
Schltr.
POACEAE
Agrostis basalis Luces
10.1
PAz
A. perennans (Walter) Tucker
1
PAz
A. sp. B
0
PAz
Aulonemia trianae (Munro)
McClure
9
SP
Calamagrostis bogotensis (Pilg.)
Pilg.
6
Paz/Pzo
C. planifolia (Kunth) Trin. ex
Steud.
7
Pzo
C. sp. A
0
Paz/Pzo
Chusquea aff. fendleri Munro
9
Pzo
C. angustifolia (Soderstr. & C.E.
Pzo/Paz/SAR
Calderon) L. G. Clark
9
F
C.mollis (Swallen) L.G. Clark
9
Pzo/SP
C. spectabilis L.G. Clark
8
Pzo/SP
C. spencei Ernst.
9
Pzo/SARF
C.steyermarkii L.G. Clark
10.2
Pzo
C. tessellata Munro
7
Pzo
Cortaderia hapalotricha (Pilg.)
Conert.
6
Paz/Pzo
Danthonia secundiflora J. Presl.
subsp. secundiflora
2
Pzo
Festuca guaramacalana Stancik 10.4
Pzo
Festuca sp.
0
Pzo
Ortachne erectifolia (Swallen)
Clayton
6
PAz
Poa annua L.*
1
Pzo
Polypogon elongatus Kunth*
TOFIELDIACEAE
Isidrogalvia robustior
(Steyerm.) Cruden.
10.3
Pzo
XYRIDACEAE
Xyris subulata Ruíz & Pav. var.
acutifolia Heimerl.
6
Paz/Pzo
Appendix
_______________________________________________________
Appendix 6. List of species and their trait states (trait state codes are in Table 6.1). The
DCA axis columns give the sample scores (following CANOCO 4.5 terminology) of the
DCA analyses of the species-to-trait state matrices (compare Fig. 6.2).
Family
Species name
Acanthaceae
Aphelandra macrophylla Leonard
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
1, 4, 5, 16, 20, 30, 38, 52, 56, 64
2.51
1.68
3.21
2.38
1, 4, 5, 16, 20, 31, 35, 52, 56, 64
2.51
1.68
3.50
2.25
Actinidiaceae
Saurauia tomentosa (Kunth) Spreng.
4, 5, 16, 19, 30, 36, 52, 56, 64
2.51
1.68
3.75
1.46
Anacardiaceae
Tapirira guianensis Aubl.
4, 5, 13, 21, 30, 34, 52, 58, 64
1.67
2.58
2.88
0.70
Annonaceae
Rollinia mucosa (Jacq.) Baill.
4, 5, 12, 27, 33, 35, 52, 56, 64
2.51
1.68
3.88
2.39
4, 5, 16, 19, 31, 35, 52, 56, 64
2.51
1.68
3.72
1.54
4, 5, 13, 21, 30, 34, 52, 56, 61
3.10
2.49
2.88
0.70
4, 5, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.88
0.70
Ilex myricoides Kunth
4, 5, 13, 21, 30, 34, 52, 56, 61
3.10
2.49
2.88
0.70
Ilex sp. 1
4, 5, 13, 21, 29, 34, 52, 56, 62
3.48
2.29
2.60
0.72
Ilex sp. 2
4, 5, 13, 21, 29, 34, 52, 56, 62
3.48
2.29
2.60
0.72
Ilex truxillensis Turcz. subsp.
bullatissima Cuatrec.
4, 5, 13, 21, 30, 35, 52, 56, 61
3.10
2.49
3.26
0.86
Araceae
Anthurium eminens Schott
4, 5, 16, 19, 30, 34, 41, 58, 65
0.67
2.43
3.22
1.13
Anthurium ginesii Croat
4, 5, 16, 19, 30, 34, 41, 56, 63
2.96
1.04
3.22
1.13
Anthurium humboldtianum Kunth
4, 5, 16, 19, 30, 34, 41, 56, 65
1.51
1.54
3.22
1.13
Anthurium nymphaeifolium K. Koch
& Bouche
Anthurium smaragdinum Bunting
4, 5, 16, 19, 30, 34, 45, 56, 64
1.83
0.73
3.22
1.13
4, 5, 16, 19, 30, 34, 41, 56, 64
2.10
0.74
3.22
1.13
4, 5, 14, 22, 30, 34, 52, 56, 63
3.36
1.99
3.24
1.80
4, 5, 7, 16, 21, 30, 35, 52, 56, 64
2.51
1.68
3.73
0.92
4, 11, 13, 19, 30, 35, 52, 56, 64
2.51
1.68
3.04
1.03
4, 5, 16, 21, 30, 34, 52, 58, 64
1.67
2.58
2.92
1.05
4, 5, 16, 21, 30, 35, 48, 58, 65
0.00
4.17
3.31
1.20
4, 5, 13, 19, 30, 34, 48, 58, 65
0.00
4.17
3.17
0.79
Ruellia tubiflora Kunth var.
tetrastichantha (Lindau) Leon
Trigynaea duckei (R.E. Fr.) R.E. Fr.
Aquifoliaceae
Ilex guaramacalensis Cuello &
Aymard
Ilex laurina Kunth
Araliaceae
Dendropanax arboreus (L.) Dcne. &
Planch.
Oreopanax discolor (Kunth) Decne.
& Planch.
Oreopanax sp.
Schefflera ferruginea (Willd. ex
Roem. & Schult.) Harms
Arecaceae
Aiphanes stergiosii Nino, Dorr &
Stauffer
Geonoma jussieuana Mart.
225
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Geonoma orbignyana Mart.
4, 5, 13, 19, 30, 34, 48, 58, 65
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
0.00 4.17 3.17 0.79
Geonoma undata Klotzsch
4, 5, 14, 19, 30, 34, 48, 58, 65
0.00
4.17
3.07
1.21
Prestoea acuminata (Willd.) H.E.
Moore var. acuminata
Wettinia praemorsa (Willd.) Wess.
Boer
1, 4, 5, 14, 21, 31, 35, 48, 58, 65
0.00
4.17
3.33
1.82
1, 4, 5, 16, 21, 31, 35, 48, 58, 65
0.00
4.17
3.46
1.75
2, 5, 16, 18, 28, 35, 52, 56, 63
3.36
1.99
2.03
1.74
2, 5, 16, 18, 29, 35, 55, 56, 61
4.15
2.51
2.38
1.71
Family
Species name
Asteraceae
Ageratina neriifolia (B.L. Rob.) R.M.
King & H. Rob.
Ageratina theifolia (Benth.) R. M.
King & H. Rob.
Baccharis brachylaenoides DC.
trait state code
2, 10, 13, 18, 28, 34, 52, 56, 61
3.10
2.49
1.49
0.78
Critoniopsis paradoxa (Sch. Bip.)
V.M. Badillo
Diplostephium obtusum Blake.
2, 5, 16, 18, 30, 36, 52, 56, 63
3.36
1.99
2.80
1.87
2, 5, 14, 18, 29, 34, 52, 56, 61
3.10
2.49
1.84
1.64
Libanothamnus griffinii (Ruiz-Teran
& Lopez-Fig.) Cuatrec.
Mikania banisteriae DC.
2, 5, 14, 18, 29, 35, 52, 56, 63
3.36
1.99
2.23
1.79
2, 5, 16, 18, 29, 35, 47, 56, 63
3.72
2.07
2.38
1.71
Mikania houstoniana (L.) B.L. Rob.
2, 5, 16, 18, 29, 34, 47, 56, 63
3.72
2.07
1.99
1.56
Mikania nigropunctulata Hieron
2, 5, 16, 18, 29, 35, 47, 56, 63
3.72
2.07
2.38
1.71
Mikania sp. 1
2, 5, 16, 18, 29, 35, 47, 56, 62
3.84
2.37
2.38
1.71
Mikania stuebelii Hieron
2, 5, 16, 18, 30, 35, 47, 56, 61
3.46
2.57
2.65
1.69
Paragynoxis cuatrecasasii Ruiz-Teran
& Lopez Figueiras
Paragynoxis venezuelae (V.M.
Badillo) Cuatrec.
Pentacalia cachacoensis (Cuatrec.)
Cuatrec.
Pentacalia greenmaniana (Hieron.)
Cuatrec.
Pentacalia theaefolia (Benth.)
Cuatrec.
Pentacalia vicelliptica (Cuatrec.)
Cuatrec.
Ruilopezia paltonioides (Standl.)
Cuatrec.
2, 5, 16, 18, 29, 35, 52, 56, 64
2.51
1.68
2.38
1.71
2, 5, 16, 18, 29, 35, 52, 56, 64
2.51
1.68
2.38
1.71
2, 5, 14, 18, 29, 35, 55, 56, 61
4.15
2.51
2.23
1.79
2, 5, 14, 18, 28, 35, 55, 56, 63
4.42
2.01
1.87
1.82
2, 5, 16, 18, 29, 35, 47, 56, 61
3.46
2.57
2.38
1.71
2, 5, 14, 18, 29, 35, 47, 56, 61
3.46
2.57
2.23
1.79
1, 5, 14, 18, 29, 34, 50, 56, 63
4.38
1.22
2.34
1.76
4, 5, 16, 19, 30, 37, 47, 56, 63
3.72
2.07
3.92
1.46
Tabebuia guayacan (Seem.) Hemsl.
2, 5, 16, 20, 33, 38, 52, 58, 64
1.67
2.58
2.89
3.08
Blechnaceae
Blechnum schomburgkii (Kl.) C. Chr.
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Bombacaceae
Matisia cf. ochrocalyx K. Schum
4, 11, 16, 21, 32, 38, 52, 56, 63
3.36
1.99
2.43
2.20
4, 11, 16, 21, 32, 38, 52, 56, 64
2.51
1.68
2.43
2.20
Bignoniaceae
Schlegelia spruceana K. Schum.
Quararibea magnifica Pittier
226
Appendix
_______________________________________________________
Family
Species name
Bromeliaceae
Greigia albo-rosea (Griseb) Mez
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
4, 11, 16, 19, 30, 38, 45, 56, 65
1.24
1.53
2.86
1.78
4, 5, 13, 23, 31, 34, 52, 58, 63
2.52
2.88
2.85
1.55
4, 5, 13, 23, 29, 34, 52, 58, 64
1.67
2.58
2.46
1.31
1, 4, 5, 13, 20, 31, 35, 52, 58, 63
2.52
2.88
3.45
1.96
4, 5, 7, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.99
0.99
4, 5, 7, 13, 14, 15, 19, 32, 37, 52,
56, 64
2.51
1.68
3.69
1.45
4, 5, 13, 14, 20, 30, 34, 47, 56,
63
3.72
2.07
2.81
1.48
1, 4, 5, 13, 14, 20, 30, 34, 52, 56,
62
4, 7, 13, 19, 29, 34, 52, 56, 63
3.48
2.29
2.92
1.74
3.36
1.99
3.44
0.34
4, 10, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.77
0.24
4, 10, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.77
0.24
Hedyosmum cuatrecazanum Occhioni 4, 10, 13, 21, 30, 35, 52, 56, 63
3.36
1.99
3.16
0.40
Hedyosmum racemosum (Ruiz &
Pav.) G. Don
Hedyosmum sp. A
4, 10, 13, 21, 30, 35, 52, 56, 63
3.36
1.99
3.16
0.40
4, 10, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.77
0.24
Hedyosmum translucidum Cuatrec.
4, 10, 13, 21, 31, 34, 52, 56, 63
3.36
1.99
2.89
0.50
Clethraceae
Clethra fagifolia Kunth var. fagifolia
2, 5, 16, 20, 30, 34, 52, 56, 63
3.36
1.99
2.49
1.89
Clusiaceae
Clusia alata Triana & Planch.
Brunelliaceae
Brunellia acutangula Humb. &
Bonpl.
Brunellia cf. integrifolia Szyszyll.
Burseraceae
Protium tovarense Pittier
Caprifoliaceae
Viburnum tinoides L.f. var.
venezuelensis (Killip & A. C. Smith)
Steyerm.
Caricaceae
Vasconcella microcarpa (Jacq.) A.
DC.
Celastraceae
Celastrus liebmannii Standl.
Maytenus macrocarpa (Ruiz & Pav.)
Briq.
Perrottetia quinduensis Kunth
Chloranthaceae
Hedyosmum cf. gentryii D'Arcy &
Liesner
Hedyosmum crenatum Occhioni
4, 5, 13, 22, 32, 38, 46, 56, 64
3.24
0.88
3.38
2.44
Clusia sp. 1
4, 5, 13, 22, 32, 38, 47, 56, 63
3.72
2.07
3.38
2.44
Clusia sp. A? (C. multiflora group)
4, 5, 13, 22, 31, 38, 46, 56, 63
4.09
1.18
3.63
2.20
Clusia trochiformis Vesque
4, 5, 13, 22, 30, 35, 46, 56, 63
4.09
1.18
3.73
1.54
3.72
2.00
2.92
2.18
3.36
1.99
3.61
1.28
Hypericum paramitanum N. Robson
1, 2, 9, 10, 16, 20, 30, 36, 52, 56,
60
Vismia baccifera (L.) Triana & Planch 4, 5, 16, 19, 30, 35, 52, 56, 63
subsp. dealbata (Kunth) Ewan
Cucurbitaceae
227
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
4, 11, 14, 19, 32, 36, 47, 56, 63
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
3.72 2.07 2.94 2.12
Cunoniaceae
Weinmannia aff. balbisiana Kunth
2, 5, 16, 23, 30, 34, 52, 56, 62
3.48
2.29
2.31
1.90
Weinmannia auriculata D. Don
2, 5, 16, 23, 30, 34, 52, 58, 61
2.26
3.38
2.31
1.90
Weinmannia fagaroides Kunth
2, 5, 16, 23, 29, 34, 52, 58, 61
2.26
3.38
2.04
1.91
Weinmannia glabra L.f.
2, 5, 16, 23, 29, 34, 52, 58, 61
2.26
3.38
2.04
1.91
Weinmannia karsteniana Szyszyll.
2, 5, 16, 23, 30, 34, 52, 56, 61
3.10
2.49
2.31
1.90
Weinmannia lechleriana Engl.
2, 5, 16, 23, 30, 35, 52, 58, 62
2.63
3.18
2.69
2.05
Weinmannia sorbifolia Kunth
2, 5, 16, 23, 30, 34, 52, 58, 63
2.52
2.88
2.31
1.90
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Alsophila erinacea (Karst.) Conant.
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Cyathea aff. straminea H. Karst
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Cyathea caracasana (Klotzsch)
Domin
Cyathea fulva (Mart. & Gal.) Fee
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Cyathea kalbreyeri (Baker) Domin
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Cyathea pauciflora (Kuhn) Lellinger
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Cyathea pungens (Willd.) Domin
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Sphaeropteris sp.
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
4, 5, 14, 27, 32, 35, 41, 57, 64
0.99
0.00
3.14
2.35
4, 5, 14, 27, 32, 36, 45, 57, 65
0.12
0.79
3.29
2.52
4, 5, 16, 21, 30, 34, 47, 56, 63
3.72
2.07
2.92
1.05
Dicksoniaceae
Dicksonia sellowiana Hook.
2, 11, 17, 26, 28, 34, 53, 58, 65
0.00
4.17
0.00
1.83
Dryopteridaceae
Diplazium celtidifolium Kunze
2, 11, 17, 26, 28, 34, 45, 58, 65
0.40
2.43
0.00
1.83
2, 11, 17, 26, 28, 34, 45, 58, 65
0.40
2.43
0.00
1.83
Elaeocarpaceae
Sloanea brevispina F. Sm.
1, 4, 5, 16, 20, 31, 35, 52, 56, 63
3.36
1.99
3.50
2.25
Sloanea guianensis Aubl.
1, 4, 5, 16, 20, 29, 35, 52, 56, 62
3.48
2.29
3.17
2.04
Sloanea laurifolia (Benth.) Benth.
1, 4, 5, 16, 20, 30, 34, 52, 56, 61
3.10
2.49
3.07
1.90
Sloanea rufa Planch. Ex Benth.
1, 4, 5, 16, 20, 32, 35, 52, 56, 64
2.51
1.68
3.29
2.46
Family
Species name
Elateriopsis oerstedii (Cogn.) Pittier
Cyatheaceae
Alsophila engelii Tryon
Cyclanthaceae
Asplundia vagans Harling
Sphaeradenia laucheana (Mast.)
Harling
Dichapetalaceae
Dichapetalum pedunculatum (DC.)
Baill.
Diplazium hians Kunze ex Klotzsch.
228
trait state code
Appendix
_______________________________________________________
Family
Species name
Ericaceae
Diogenesia tetrandra (A. C. Jm.)
Sleumer
Disterigma alaternoides (Kunth)
Nied.
Disterigma sp.
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
4, 5, 16, 19, 30, 35, 47, 56, 62
3.84
2.37
3.61
1.28
4, 7, 16, 19, 30, 35, 55, 56, 61
4.15
2.51
4.15
0.82
4, 7, 16, 19, 30, 35, 47, 56, 60
4.07
2.08
4.15
0.82
Gaultheria anastomosans (L.f.) Kunth 2, 8, 16, 20, 30, 35, 55, 56, 60
4.77
2.03
2.87
2.48
Gaultheria erecta Vent.
4, 5, 16, 20, 30, 35, 55, 56, 62
4.53
2.31
3.35
1.79
Macleania rupestris (Kunth) A.C.
Sm.
Psammisia hookeriana Klotzsch.
4, 7, 16, 19, 31, 36, 47, 56, 63
3.72
2.07
4.41
1.25
4, 7, 16, 19, 30, 35, 47, 56, 63
3.72
2.07
4.15
0.82
Themistoclesia dependens (Benth.)
A.C. Sm.
Thibaudia floribunda Kunth.
4, 7, 16, 19, 30, 36, 55, 56, 61
4.15
2.51
4.29
1.00
4, 7, 16, 19, 31, 36, 47, 56, 63
3.72
2.07
4.41
1.25
Vaccinium corymbodendron Dunal
4, 7, 16, 19, 30, 35, 55, 56, 60
4.77
2.03
4.15
0.82
3.84
2.37
2.87
2.03
1, 10, 13, 14, 20, 29, 35, 52, 56,
63
Alchornea glandulosa Poepp. & Endl. 4, 7, 10, 13, 20, 30, 34, 52, 56,
64
Alchornea grandiflora Muell. Arg.
4, 7, 10, 13, 20, 30, 34, 52, 56,
63
Croizatia brevipetiolata Govaerts
1, 5, 13, 20, 31, 36, 52, 56, 63
3.36
1.99
2.84
1.56
2.51
1.68
3.31
0.59
3.36
1.99
3.31
0.59
3.36
1.99
3.58
2.25
Hieronyma cf. oblonga (Tul.) Mull.
Arg.
Hieronyma fendleri Briq.
4, 5, 13, 21, 29, 34, 52, 56, 63
3.36
1.99
2.60
0.72
4, 5, 13, 21, 29, 34, 52, 56, 62
3.48
2.29
2.60
0.72
Hieronyma scabrida (Tul.) Mull. Arg. 4, 5, 13, 21, 30, 35, 52, 56, 63
3.36
1.99
3.26
0.86
Mabea occidentalis Benth.
1, 6, 14, 20, 30, 34, 52, 56, 62
3.48
2.29
3.12
2.42
Sapium stylare Mull. Arg.
1, 5, 14, 20, 30, 34, 52, 56, 62
3.48
2.29
2.83
2.09
Tetrorchidium rubrivenium Poepp.
1, 4, 5, 14, 20, 30, 34, 52, 56, 63
3.36
1.99
2.94
1.97
1, 4, 5, 16, 24, 32, 36, 52, 58, 65
1.08
3.38
4.00
2.79
2, 5, 16, 18, 31, 35, 47, 58, 64
2.03
2.66
2.77
1.95
Flacourtiaceae
Casearia tachirensis Sleumer
4, 5, 16, 20, 30, 35, 52, 56, 63
3.36
1.99
3.35
1.79
Gentianiaceae
Macrocarpaea bracteata Ewan
1, 6, 16, 20, 31, 36, 52, 56, 63
3.36
1.99
3.92
2.92
1, 6, 16, 20, 32, 38, 52, 56, 63
3.36
1.99
3.31
3.40
Escalloniaceae
cf. Escallonia hispida (Vell.) Sleumer 2, 3, 4, 5, 16, 20, 30, 35, 47, 56,
62
Euphorbiaceae
Acalypha macrostachya Jacq.
Fabaceae
Dussia coriacea (Sw.) Roem. &
Schult.
Machaerium cf. floribundum Benth.
Symbolanthus vasculosus (Griseb.)
Gilg.
229
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Family
Species name
Gesneriaceae
Besleria pendula Hanst.
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
4, 7, 16, 19, 30, 36, 52, 56, 62
3.48
2.29
4.29
1.00
4, 6, 8, 16, 22, 31, 38, 47, 56, 62
3.84
2.37
3.82
3.11
Hippocrateaceae
Salacia aff. cordata (Miers.) Mennega 4, 11, 16, 21, 29, 35, 47, 56, 63
3.72
2.07
2.51
1.31
Hydrangeaceae
Hydrangea aff. peruviana Moricard
2, 5, 16, 20, 29, 34, 52, 56, 63
3.36
1.99
2.22
1.90
Hydrangea cf. preslii Briq.
2, 5, 16, 20, 29, 34, 47, 56, 63
3.72
2.07
2.22
1.90
Hydrangea sp. 1
2, 5, 16, 20, 29, 34, 47, 56, 63
3.72
2.07
2.22
1.90
4, 11, 14, 15, 21, 32, 34, 52, 56,
63
4, 11, 14, 15, 21, 31, 35, 52, 56,
63
3.36
1.99
2.14
1.69
3.36
1.99
2.68
1.61
4, 5, 12, 19, 31, 34, 52, 56, 63
3.36
1.99
3.55
1.38
Aniba cf. cinnamomiflora C. K. Allen
4, 5, 12, 19, 31, 34, 52, 56, 63
3.36
1.99
3.55
1.38
Beilschmiedia tovarensis (Meissn.)
Sa. Nishida
Endlicheria sp.
4, 5, 16, 19, 31, 34, 52, 56, 63
3.36
1.99
3.34
1.39
4, 5, 16, 21, 31, 34, 52, 56, 63
3.36
1.99
3.04
1.30
Nectandra aff. membranacea (Sw.)
Griseb.
Nectandra aff. purpurea (Ruiz &
Pav.) Mez
Nectandra sp.
4, 5, 12, 19, 31, 34, 52, 56, 64
2.51
1.68
3.55
1.38
4, 5, 12, 19, 31, 34, 52, 56, 64
0.00
0.00
0.00
0.00
4, 5, 16, 21, 31, 34, 52, 56, 63
3.36
1.99
3.04
1.30
Ocotea aff. puberula (Rich.) Nees
Drymonia crassa C.V. Morton
Icacinaceae
Calatola venezuelana Pittier
Citronella costaricensis (Donn. Sm.)
R.A. Howard
Lauraceae
Aiouea dubia (Kunth) Mez
4, 5, 13, 19, 30, 35, 52, 56, 63
3.36
1.99
3.56
0.94
Ocotea aff. tarapotana (Meissn.) Mez 4, 5, 16, 19, 31, 35, 52, 56, 62
3.48
2.29
3.72
1.54
Ocotea auriculata Lasser
4, 5, 16, 19, 30, 34, 52, 56, 64
2.51
1.68
3.22
1.13
Ocotea calophylla Mez
4, 5, 13, 19, 30, 35, 52, 56, 63
3.36
1.99
3.56
0.94
Ocotea cernua (Nees) Mez, vel aff.
4, 5, 13, 19, 30, 34, 52, 56, 63
3.36
1.99
3.17
0.79
Ocotea cf. hexanthera Kopp.
4, 5, 16, 19, 31, 35, 52, 56, 64
2.51
1.68
3.72
1.54
Ocotea floribunda (Sw.) Mez
4, 5, 13, 19, 30, 35, 52, 56, 62
3.48
2.29
3.56
0.94
Ocotea jelskii Mez
4, 5, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.61
1.28
Ocotea karsteniana Mez
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Ocotea leucoxylon (Sw.) de Lanessen, 4, 5, 13, 19, 30, 35, 52, 56, 64
s.l.
Ocotea macropoda (Kunth) Mez
4, 5, 13, 19, 30, 35, 52, 56, 63
2.51
1.68
3.56
0.94
3.36
1.99
3.56
0.94
Ocotea rubrinervis Mez
4, 5, 13, 19, 30, 35, 52, 56, 63
3.36
1.99
3.56
0.94
Ocotea sericea Kunth
4, 5, 13, 19, 31, 35, 52, 56, 63
3.36
1.99
3.67
1.20
230
Appendix
_______________________________________________________
Ocotea sp.
4, 5, 16, 19, 31, 35, 52, 56, 63
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
3.36 1.99 3.72 1.54
Ocotea sp. A
4, 5, 16, 19, 31, 35, 52, 56, 62
3.48
2.29
3.72
1.54
Ocotea sp. B
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Ocotea sp. C
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Ocotea terciopelo C. K. Allen
4, 5, 16, 19, 31, 34, 52, 56, 63
3.36
1.99
3.34
1.39
Ocotea vaginans (Meissn.) Mez
4, 5, 16, 19, 30, 34, 52, 56, 63
3.36
1.99
3.22
1.13
Persea aff. mutisii Kunth
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Persea ferruginea Kunth.
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Persea meridensis Kopp.
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Persea peruviana Nees
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Persea sp. 1
4, 5, 16, 19, 31, 35, 52, 56, 63
3.36
1.99
3.72
1.54
Persea sp. 2
4, 5, 16, 19, 31, 35, 52, 56, 62
3.48
2.29
3.72
1.54
Persea sp. 3
4, 5, 16, 19, 31, 35, 52, 56, 64
2.51
1.68
3.72
1.54
Pleurothyrium costanense van der
Werff
4, 11, 16, 19, 31, 35, 52, 56, 64
2.51
1.68
3.20
1.63
3.36
1.99
3.29
2.46
3.36
1.99
2.96
2.32
4, 5, 16, 19, 30, 35, 52, 56, 60
3.72
2.00
3.61
1.28
Magnoliaceae
Talauma sp.
4, 5, 16, 23, 32, 38, 52, 56, 64
2.51
1.68
2.82
2.70
Malpighiaceae
Bunchosia armeniaca (Cav.) DC.
4, 5, 16, 21, 31, 35, 52, 56, 63
3.36
1.99
3.42
1.46
Byrsonima karstenii W. R. Anderson
4, 5, 16, 21, 30, 35, 52, 56, 63
3.36
1.99
3.31
1.20
Byrsonima sp.
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Hiraea sp.
2, 5, 16, 18, 30, 35, 47, 56, 62
3.84
2.37
2.65
1.69
4, 5, 7, 16, 20, 30, 37, 47, 56, 63
3.72
2.07
4.03
1.57
4, 5, 16, 19, 30, 36, 52, 56, 63
3.36
1.99
3.75
1.46
4, 5, 16, 19, 30, 36, 47, 56, 63
3.72
2.07
3.75
1.46
2, 5, 16, 20, 29, 35, 55, 56, 59
5.08
2.10
2.60
2.06
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Family
Species name
trait state code
Lecythidaceae
Eschweilera aff. antioquensis Dugand 1, 4, 5, 16, 20, 32, 35, 52, 56, 63
& Daniel
Eschweilera perumbonata Pittier
1, 4, 5, 16, 20, 32, 34, 52, 56, 63
Loranthaceae
Gaiadendron punctatum (R. & P.) G.
Don
Marcgraviaceae
Marcgravia brownei (Tr. & Pl.) Krug
& Urb.
Melastomataceae
Anaectocalyx bracteosa (Naud.)
Triana
Blakea schlimii (Naud.) Triana
Chaetolepis lindeniana (Naud.)
Triana
Henriettella cf. verrucosa Triana
231
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Henriettella sp.
4, 5, 16, 19, 30, 35, 52, 56, 63
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
3.36 1.99 3.61 1.28
Henriettella tovarensis Cogn.
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
Meriania grandidens Triana
Family
Species name
trait state code
1, 2, 5, 16, 20, 30, 37, 52, 56, 63
3.36
1.99
3.26
2.40
Meriania macrophylla (Benth.) Triana 1, 2, 5, 16, 20, 30, 34, 52, 56, 64
2.51
1.68
2.66
2.12
Miconia amilcariana Almeda & Dorr
4, 5, 16, 19, 28, 34, 52, 56, 63
3.36
1.99
2.60
1.18
Miconia cf. dolichopoda Naud.
4, 5, 16, 19, 29, 34, 52, 56, 63
3.36
1.99
2.95
1.14
Miconia cf. minutiflora (Bonpl.) DC.
4, 5, 16, 19, 28, 34, 52, 56, 63
3.36
1.99
2.60
1.18
Miconia donaeana Naud.
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
Miconia elvirae Wurdack
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
Miconia jahnii Pittier
4, 5, 16, 19, 29, 34, 52, 56, 63
3.36
1.99
2.95
1.14
Miconia lonchophylla Naud.
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
Miconia lucida Naud.
4, 5, 16, 19, 29, 35, 52, 56, 64
2.51
1.68
3.34
1.30
Miconia mesmeana Gleason subsp.
longipetiolata Wurdack
Miconia sp. C (hibrido)
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
4, 5, 16, 19, 29, 34, 52, 56, 62
3.48
2.29
2.95
1.14
Miconia sp. B
4, 5, 16, 19, 32, 35, 52, 56, 63
3.36
1.99
3.48
1.78
Miconia suaveolens Wurdack
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Miconia theaezans (Bonpl.) Cogn., s.l. 4, 5, 16, 19, 28, 34, 52, 56, 63
3.36
1.99
2.60
1.18
Miconia tinifolia Naud.
4, 5, 16, 19, 29, 34, 52, 56, 62
3.48
2.29
2.95
1.14
Miconia tovarensis Cogn.
4, 5, 16, 19, 29, 34, 52, 56, 63
3.36
1.99
2.95
1.14
Miconia ulmarioides Naud.
4, 5, 16, 19, 29, 35, 52, 56, 63
3.36
1.99
3.34
1.30
Monochaetum discolor H. Karst.
4, 5, 16, 20, 30, 37, 55, 56, 61
4.15
2.51
3.66
1.96
Ossaea micrantha (Sw.) Macfad.
4, 5, 16, 20, 28, 35, 52, 56, 63
3.36
1.99
2.72
1.83
4, 5, 13, 20, 32, 36, 52, 58, 65,
1.08
3.38
3.31
2.12
Ruagea glabra Triana & Planch.
4, 5, 7, 13, 20, 31, 35, 52, 58, 64
1.67
2.58
3.82
1.34
Ruagea pubescens H. Karst.
4, 5, 7, 13, 20, 31, 35, 52, 58, 64
1.67
2.58
3.82
1.34
Trichilia hirta L.
4, 5, 13, 20, 30, 35, 52, 58, 64
1.67
2.58
3.30
1.44
Trichilia pallida Sw.
4, 5, 13, 14, 20, 30, 35, 52, 58,
64
4, 5, 13, 14, 20, 31, 35, 52, 58,
64
1.67
2.58
3.14
1.62
1.67
2.58
3.24
1.84
4, 7, 16, 21, 31, 36, 47, 56, 63
3.72
2.07
4.11
1.17
4, 5, 16, 24, 32, 35, 52, 58, 65
1.08
3.38
3.90
2.50
Meliaceae
Guarea kunthiana A. Juss.
Trichilia septentrionalis C. DC.
Mendonciaceae
Mendoncia tovarensis (Klotzsch &
Karsten ex Nees) Leonard
Mimosaceae
Inga aff. densiflora Benth.
232
Appendix
_______________________________________________________
Inga edulis Mart.
4, 5, 16, 24, 33, 36, 52, 58, 64
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
1.67 2.58 4.41 2.80
Zygia bisingula L. Rico
1, 5, 16, 24, 33, 36, 52, 58, 64
1.67
2.58
4.43
3.18
4, 5, 13, 18, 28, 34, 52, 56, 64
2.51
1.68
2.06
0.99
Cecropia sp.
4, 5, 13, 18, 28, 34, 52, 56, 64
2.51
1.68
2.06
0.99
Cecropia telenitida Cuatrec.
4, 5, 13, 18, 28, 34, 52, 56, 65
1.92
2.48
2.06
0.99
Ficus nymphaeifolia P. Miller
4, 5, 14, 27, 31, 34, 52, 56, 63
3.36
1.99
3.00
1.95
Ficus sp.
4, 5, 14, 27, 31, 34, 52, 56, 64
2.51
1.68
3.00
1.95
Ficus tonduzii Standl.
4, 5, 14, 27, 31, 34, 52, 56, 64
2.51
1.68
3.00
1.95
Ficus tovarensis Pittier
4, 5, 14, 27, 30, 34, 52, 56, 63
3.36
1.99
2.89
1.69
Morus insignis Bureau
4, 5, 13, 14, 27, 29, 34, 52, 56,
63
4, 5, 13, 21, 30, 35, 52, 56, 63
3.36
1.99
2.64
1.48
3.36
1.99
3.26
0.86
4, 5, 13, 21, 29, 34, 52, 56, 63
3.36
1.99
2.60
0.72
Cybianthus iteoides (Benth.) Agost.
4, 5, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.88
0.70
Cybianthus laurifolius (Mez) Agost.
4, 5, 13, 21, 30, 34, 52, 56, 63
3.36
1.99
2.88
0.70
Cybianthus marginatus (Benth.)
Pipoly
Cybianthus stapfii (Mez) Agostini
4, 5, 14, 21, 29, 34, 52, 56, 63
3.36
1.99
2.50
1.14
4, 5, 13, 21, 29, 34, 52, 56, 61
3.10
2.49
2.60
0.72
Geissanthus andinus Mez
4, 5, 15, 21, 30, 34, 52, 56, 61
3.10
2.49
2.79
0.94
Geissanthus fragrans Mez
4, 5, 15, 21, 30, 34, 52, 56, 64
2.51
1.68
2.79
0.94
Myrsine coriacea (Sw.) R. Br. ex
Roem & Schult.
Myrsine dependens (Ruiz & Pav.)
Spreng.
Parathesis venezuelana Mez
4, 5, 13, 21, 29, 34, 52, 56, 61
3.10
2.49
2.60
0.72
4, 5, 13, 21, 29, 34, 52, 56, 60
3.72
2.00
2.60
0.72
4, 5, 16, 21, 29, 34, 52, 56, 64
2.51
1.68
2.65
1.06
Stylogyne longifolia (Mart. Ex Miq.)
Mez
Stylogyne sp. A
4, 5, 16, 21, 30, 34, 52, 56, 64
2.51
1.68
2.92
1.05
4, 5, 16, 21, 29, 34, 52, 56, 63
3.36
1.99
2.65
1.06
Myrtaceae
Calyptranthes cf. meridensis Steyerm. 4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Family
Species name
Moraceae
Cecropia sararensis Cuatrec.
Pseudolmedia rigida (Planch. &
Karst.) Cuatrec. subsp. rigida
Myrsinaceae
Cybianthus cuspidatus Miq.
trait state code
Calyptranthes sp.
4, 5, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.61
1.28
Eugenia albida Humb. & Bonpl.
4, 5, 16, 19, 30, 36, 52, 56, 63
3.36
1.99
3.75
1.46
Eugenia cf. oerstediana O. Berg.
4, 5, 16, 19, 29, 34, 52, 56, 62
3.48
2.29
2.95
1.14
Eugenia cf. patens Poir.
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Eugenia cf. tamaensis Steyerm.
4, 5, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.61
1.28
Eugenia grandiflora O. Berg.
4, 5, 16, 19, 30, 36, 52, 56, 64
2.51
1.68
3.75
1.46
233
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Eugenia moritziana H. Karst.
4, 5, 16, 19, 31, 35, 52, 56, 63
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
3.36 1.99 3.72 1.54
Eugenia sp.
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Eugenia sp. 1
4, 5, 16, 19, 30, 35, 52, 56, 61
3.10
2.49
3.61
1.28
Eugenia sp. 2
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
Eugenia sp. 3
4, 5, 16, 19, 32, 35, 52, 56, 63
3.36
1.99
3.48
1.78
Eugenia sp.?
4, 5, 16, 19, 30, 35, 52, 56, 61
3.10
2.49
3.61
1.28
Eugenia triquetra Berg
4, 5, 16, 19, 30, 35, 52, 56, 61
3.10
2.49
3.61
1.28
Myrcia acuminata (Kunth) DC.
4, 5, 16, 19, 30, 34, 52, 56, 63
3.36
1.99
3.22
1.13
Myrcia aff. guianensis (Aubl.) DC.
4, 5, 16, 19, 30, 34, 52, 56, 61
3.10
2.49
3.22
1.13
Myrcia cf. sanisidrensis Steyerm.
4, 5, 16, 19, 30, 34, 52, 56, 63
3.36
1.99
3.22
1.13
Myrcia sp. 1
4, 5, 16, 19, 30, 34, 52, 56, 61
3.10
2.49
3.22
1.13
Myrcianthes sp.
4, 5, 16, 19, 30, 34, 52, 56, 61
3.10
2.49
3.22
1.13
4, 5, 16, 18, 29, 35, 52, 56, 63
3.36
1.99
2.85
1.45
4, 5, 16, 21, 29, 34, 52, 56, 63
3.36
1.99
2.65
1.06
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
4, 5, 7, 16, 19, 31, 38, 47, 56, 61
3.46
2.57
3.90
1.56
3.36
1.99
2.07
1.33
Family
Species name
Nyctaginaceae
Neea sp.
Olacaceae
Heisteria acuminata (Humb. &
Bonpl.) Engler
Chionanthus sp.
trait state code
Onagraceae
Fuchsia membranacea Hemsl.
Piperaceae
Piper aduncum L. var. cordulatum (C. 2, 4, 5, 16, 21, 28, 34, 52, 56, 63
DC.) Yunck.
Piper arboreum Aubl.
4, 5, 16, 21, 28, 34, 52, 56, 64
2.51
1.68
2.30
1.10
Piper hispidum Sw.
4, 5, 16, 21, 28, 34, 52, 56, 64
2.51
1.68
2.30
1.10
Piper longispicum C. DC. var.
glabratum (Yunck.) Steyerm.
Piper phytolaccifolium Opiz
4, 5, 16, 21, 28, 34, 52, 56, 64
2.51
1.68
2.30
1.10
4, 5, 16, 21, 29, 34, 52, 56, 63
3.36
1.99
2.65
1.06
Piper sp.
4, 5, 16, 21, 29, 34, 52, 56, 62
3.48
2.29
2.65
1.06
Piper sp. 1- Liana
4, 5, 16, 21, 29, 34, 47, 56, 62
3.84
2.37
2.65
1.06
Piper veraguense C. DC.
4, 5, 16, 21, 28, 34, 52, 56, 63
3.36
1.99
2.30
1.10
4, 10, 14, 15, 18, 30, 35, 39, 56,
63
4, 10, 16, 18, 30, 35, 39, 56, 61
4.32
2.32
2.79
1.12
4.06
2.82
3.02
0.98
4, 10, 16, 18, 30, 34, 39, 56, 61
4.06
2.82
2.64
0.82
Rhipidocladum geminatum (McClure) 4, 10, 16, 18, 30, 35, 39, 56, 61
McClure
4.06
2.82
3.02
0.98
Poaceae
Arthrostylidium venezuelae (Steud.)
McClure
Chusquea angustifolia (Soderstr. &
C.E. Calderon) L.G. Clark
Chusquea purdieana Munro
234
Appendix
_______________________________________________________
Family
Species name
Podocarpaceae
Podocarpus oleifolius D. Don ex
Lambert var. macrostachyus (Parl.) J.
Bunchholz & N. E. Gray
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
4, 10, 13, 25, 30, 35, 52, 56, 61
3.10
2.49
3.70
0.00
2, 10, 16, 18, 30, 36, 47, 56, 61
3.46
2.57
2.69
1.41
Monnina meridensis Planch. & Lindl.
ex Wedd.
Monnina sp. 1
4, 5, 16, 21, 30, 35, 52, 56, 61
3.10
2.49
3.31
1.20
4, 5, 16, 21, 30, 35, 52, 56, 62
3.48
2.29
3.31
1.20
Monnina sp. 2?
4, 5, 16, 21, 30, 35, 52, 56, 62
3.48
2.29
3.31
1.20
Polygonaceae
Coccoloba cf. llewelynii R.A. Howard 4, 5, 16, 18, 30, 34, 52, 56, 63
3.36
1.99
2.74
1.28
4, 5, 14, 18, 30, 34, 52, 56, 63
3.36
1.99
2.59
1.36
1, 5, 16, 23, 32, 35, 52, 56, 64
2.51
1.68
3.06
2.67
1, 5, 16, 23, 32, 35, 52, 56, 63
3.36
1.99
3.06
2.67
2, 5, 16, 23, 32, 36, 52, 56, 63
3.36
1.99
2.71
2.72
1, 4, 5, 16, 21, 29, 34, 52, 56, 63
3.36
1.99
2.80
1.41
3.10
2.49
3.78
1.88
3.72
2.00
3.78
1.88
Polygalaceae
Bredemeyera sp.
Coccoloba sp.
Proteaceae
Panopsis sp.
Panopsis suaveolens (H. Karst.)
Pittier
Roupala barnettiae Dorr
Rhamnaceae
Rhamnus sphaerosperma Sw. var.
polymorpha (Reiss.) M.C. Johnst
Rosaceae
Hesperomeles obtusifolia (Pers.) Lind. 4, 5, 16, 22, 30, 35, 52, 56, 61
var. obtusifolia
Hesperomeles sp.
4, 5, 16, 22, 30, 35, 52, 56, 60
Prunus cf. skutchii Johnston
4, 5, 16, 21, 30, 35, 52, 56, 63
3.36
1.99
3.31
1.20
Prunus moritziana Koehne
4, 5, 16, 21, 31, 35, 52, 56, 63
3.36
1.99
3.42
1.46
Rubiaceae
Coussarea moritziana (Benth.) Standl. 4, 5, 16, 21, 31, 36, 52, 56, 64
2.51
1.68
3.57
1.63
Dioicodendron dioicum (K. Schum. & 1, 5, 13, 20, 30, 34, 52, 56, 63
Krause) Styerm.
Elaeagia karstenii Standl.
1, 5, 16, 20, 30, 35, 52, 56, 64
3.36
1.99
2.94
1.67
2.51
1.68
3.37
2.17
Elaeagia myriantha (Standl.) Hammel 1, 5, 16, 20, 29, 35, 52, 56, 64
& C. M. Taylor
Elaeagia ruizteranii Steyerm.
1, 5, 16, 20, 30, 35, 52, 56, 64
2.51
1.68
3.10
2.18
2.51
1.68
3.37
2.17
Faramea guaramacalensis Taylor
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Faramea killipii Standl.
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Guettarda crispiflora Vahl subsp.
discolor (Rusby) Steyerm.
Hippotis albiflora H. Karst.
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
4, 5, 16, 19, 31, 38, 52, 56, 64
2.51
1.68
3.50
1.95
Hoffmannia pauciflora Standl.
4, 5, 7, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.99
0.99
235
_______________________________________________________
Flora, vegetation and ecology in the Venezuelan Andes
Ladenbergia cf. buntingii Steyerm.
2, 5, 16, 20, 30, 38, 52, 56, 65
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
1.92 2.48 2.65 2.45
Palicourea angustifolia Kunth
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Palicourea apicata Kunth
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Palicourea demissa Standl.
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Palicourea jahnii Standl.
4, 5, 16, 21, 30, 35, 52, 56, 61
3.10
2.49
3.31
1.20
Palicourea puberulenta Steyerm.
4, 5, 16, 21, 30, 36, 52, 56, 63
3.36
1.99
3.45
1.37
Posoqueria coriacea M. Mart. &
Galeotti subsp. formosa
Psychotria amita Stand.
4, 5, 16, 19, 32, 36, 52, 56, 64
2.51
1.68
3.62
1.96
4, 5, 16, 21, 30, 35, 52, 56, 61
3.10
2.49
3.31
1.20
Psychotria cf. lindenii Standl.
4, 5, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.61
1.28
Psychotria fortuita Standl.
4, 5, 16, 21, 30, 35, 52, 56, 62
3.48
2.29
3.31
1.20
Psychotria longirostris (Rusby)
Standl.
Psychotria trichotoma Mart. & Gal.
4, 5, 16, 19, 30, 35, 52, 56, 62
3.48
2.29
3.61
1.28
4, 5, 16, 21, 30, 35, 52, 56, 64
2.51
1.68
3.31
1.20
Randia cf. dioica H. Karst.
4, 5, 13, 19, 31, 36, 52, 56, 63
3.36
1.99
3.82
1.37
Rudgea nebulicola Steyerm.
4, 5, 16, 19, 31, 36, 52, 56, 64
2.51
1.68
3.87
1.71
Rudgea tayloriae Aymard, Dorr &
Cuello
Simira erythroxylon (Willd.) Brem.
var. meridensis Steyerm.
Simira lezamae Steyerm.
4, 5, 16, 19, 30, 35, 52, 56, 63
3.36
1.99
3.61
1.28
1, 2, 5, 16, 20, 32, 35, 52, 56, 64
2.51
1.68
2.88
2.68
1, 2, 5, 16, 20, 32, 35, 52, 56, 63
3.36
1.99
2.88
2.68
Tammsia anomala Karst.
4, 5, 16, 19, 30, 36, 52, 56, 63
3.36
1.99
3.75
1.46
Tocoyena costanensis Steyerm. subsp. 4, 5, 16, 19, 32, 36, 52, 56, 64
andina Steyerm.
2.51
1.68
3.62
1.96
1, 2, 5, 16, 20, 31, 36, 52, 58, 64
1.67
2.58
3.21
2.62
4, 5, 16, 20, 31, 35, 52, 58, 64
1.67
2.58
3.46
2.04
4, 5, 16, 20, 30, 35, 52, 58, 64
1.67
2.58
3.35
1.79
4, 5, 16, 21, 30, 34, 52, 56, 63
3.36
1.99
2.92
1.05
Meliosma pittieriana Steyerm.
4, 5, 16, 21, 32, 34, 52, 56, 63
3.36
1.99
2.79
1.55
Meliosma tachirensis Steyerm. &
Gentry
Meliosma venezuelensis Steyerm.
4, 5, 16, 21, 31, 34, 52, 56, 63
3.36
1.99
3.04
1.30
4, 5, 16, 21, 31, 34, 52, 56, 63
3.36
1.99
3.04
1.30
4, 5, 15, 20, 30, 34, 52, 58, 63
2.52
2.88
2.83
1.52
1, 4, 5, 16, 20, 32, 37, 52, 58, 63
2.52
2.88
3.55
2.61
Family
Species name
Rutaceae
Conchocarpus larensis (Tamayo &
Croizat) Kallunki & Pirani
Zanthoxylum acuminatum (Sw.) Sw
subsp. juniperinum (Poepp.) Reynel
Zanthoxylum melanostictum Schltdl.
& Cham.
Sabiaceae
Meliosma meridensis Lasser
Sapindaceae
Allophylus cf. glabratus (Kunth)
Radlk
Billia rosea (Planch. & Linden) C.
Ulloa & P. Jørg.
236
trait state code
Appendix
_______________________________________________________
Family
Species name
Cupania cf. scrobiculata Rich.
trait state code
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
1.67 2.58 3.16 1.45
Matayba camptoneura Radlk.
4, 5, 13, 15, 20, 30, 35, 52, 58,
64
4, 5, 15, 20, 30, 34, 52, 58, 64
1.67
2.58
2.83
1.52
Paullinia capreolata (Aubl.) Radlk.
4, 5, 15, 20, 31, 34, 47, 58, 64
2.03
2.66
2.95
1.78
Paullinia cf. latifolia Benth. ex Radlk
4, 5, 15, 20, 31, 34, 47, 58, 64
2.03
2.66
2.95
1.78
3.36
1.99
3.33
1.61
Chrysophyllum cf. cainito L.
4, 5, 6, 13, 14, 19, 31, 34, 52, 56,
63
4, 6, 16, 19, 30, 34, 52, 56, 63
3.36
1.99
3.51
1.45
Chrysophyllum sp.
4, 6, 16, 19, 30, 34, 52, 56, 64
2.51
1.68
3.51
1.45
Pouteria baehniana Monachino
4, 6, 16, 19, 32, 36, 52, 56, 64
2.51
1.68
3.91
2.28
Simaroubaceae
Picramnia sp. A
4, 5, 13, 19, 30, 34, 52, 58, 64
1.67
2.58
3.17
0.79
Picramnia sp. C
4, 5, 13, 19, 31, 34, 52, 58, 64
1.67
2.58
3.29
1.04
Smilacaceae
Smilax kunthii Killip & C. V. Morton
4, 5, 16, 19, 30, 35, 47, 56, 63
3.72
2.07
3.61
1.28
Solanaceae
Cestrum bigibbosum Pittier
4, 5, 16, 19, 30, 37, 52, 56, 62
3.48
2.29
3.92
1.46
Cestrum buxifolium Kunth
4, 5, 16, 19, 31, 36, 55, 56, 62
4.53
2.31
3.87
1.71
Cestrum darcyanum Benitez & N.W.
Sawyer
Cuatresia riparia (Kunth.) Humz
4, 5, 16, 19, 30, 36, 52, 56, 62
3.48
2.29
3.75
1.46
4, 5, 16, 19, 30, 36, 52, 56, 63
3.36
1.99
3.75
1.46
Markea sp.
4, 11, 16, 19, 30, 36, 52, 56, 64
2.51
1.68
3.23
1.54
Solanum aturense Humb. & Bonpl. ex 4, 5, 16, 19, 31, 36, 47, 56, 62
Dunal
Solanum confine Dunal
4, 5, 16, 19, 30, 36, 52, 56, 63
3.84
2.37
3.87
1.71
3.36
1.99
3.75
1.46
Solanum nudum Dunal
4, 5, 16, 19, 30, 36, 52, 56, 63
3.36
1.99
3.75
1.46
4, 5, 16, 20, 30, 34, 52, 58, 64
1.67
2.58
2.96
1.63
4, 5, 16, 20, 30, 35, 52, 58, 64
1.67
2.58
3.35
1.79
Symplocaceae
Symplocos bogotensis Brand.
4, 5, 16, 21, 30, 35, 52, 56, 61
3.10
2.49
3.31
1.20
Symplocos tamana Steyerm.
4, 5, 16, 21, 30, 35, 52, 56, 61
3.10
2.49
3.31
1.20
2, 11, 13, 20, 30, 36, 52, 56, 62
3.48
2.29
2.45
1.96
2, 5, 16, 20, 31, 38, 52, 56, 63
3.36
1.99
2.77
2.71
4, 5, 16, 20, 30, 36, 52, 56, 62
3.48
2.29
3.49
1.96
Sapotaceae
cf. Elaeoluma nuda (Baehni) Aubr.
Staphyleaceae
Huertea glandulosa Ruiz & Pav.
Turpinia occidentalis (Sw.) G. Don.
Theaceae
Freziera serrata A. L. Weitzman,
ined.
Gordonia fruticosa (Schrader) H.
Keng
Ternstroemia acrodantha Kobuski &
Steyerm.
237
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Flora, vegetation and ecology in the Venezuelan Andes
Ternstroemia sp. A
4, 5, 16, 20, 30, 36, 52, 56, 63
Energy
Fragmentati
balance traits
on traits
DCA
DCA
axis axis axis axis
1
2
1
2
3.36 1.99 3.49 1.96
Ternstroemia sp. B
4, 5, 16, 20, 30, 36, 52, 56, 63
3.36
1.99
3.49
1.96
4, 10, 13, 14, 18, 30, 35, 52, 56,
64
2.51
1.68
2.86
0.92
4, 5, 16, 21, 31, 35, 52, 56, 64
2.51
1.68
3.42
1.46
4, 5, 16, 21, 30, 34, 52, 56, 63
3.36
1.99
2.92
1.05
Aegiphila ternifolia (Kunth)
Moldenke
Citharexylum venezuelense Mol.
4, 5, 16, 21, 30, 35, 52, 56, 63
3.36
1.99
3.31
1.20
4, 5, 16, 21, 31, 36, 52, 56, 63
3.36
1.99
3.57
1.63
Petrea pubescens Turcz.
2, 5, 16, 21, 31, 35, 52, 56, 63
3.36
1.99
2.95
1.72
Vitaceae
Cissus trianae Planch
4, 5, 16, 19, 30, 34, 47, 58, 61
2.61
3.46
3.22
1.13
Winteraceae
Drimys granadensis L.f.
4, 5, 16, 19, 30, 37, 52, 56, 62
3.48
2.29
3.92
1.46
Family
Species name
Urticaceae
Urera baccifera (L.) Gaudich ex
Wedd.
Verbenaceae
Aegiphila floribunda Moritz &
Moldenke
Aegiphila moldenkeana Lopez-Pal.
238
trait state code
Summary
_______________________________________________________
SUMMARY
Ramal de Guaramacal is an outlier and lower elevation mountain range up to 3,130
m located at the northeastern end of the Venezuelan Andes.
In Chapter 2, montane forest community composition of Ramal de Guaramacal
was studied along the altitudinal gradient on both sides of the range with different
slope expositions. Thirty five 0.1 ha plots were surveyed, with variable intervals of
30 to 150 meters between 1350 m and 2890 m and nine plots of variable size (50
m2 to 400 m2) were surveyed in dwarf forests located between 2800-3050 m. A
total of 388 morphospecies with dbh ≥ 2.5 cm, corresponding to 189 genera and 78
families of vascular plants, were recorded from a total of 44 forest plots. The
TWINSPAN phytosociological clustering, based on both floristic composition and
species relative abundance, revealed seven forest communities at association level,
grouped in three alliances and one montane forest order group. Three subandean
forest (LMRF) communities and four Andean - high Andean forest (UMRFSARF) communities are distinguished and described according to the ZürichMontpellier method. The Geonomo undatae-Posoquerion coriaceae alliance
contains two subandean forest communities (Simiro erythroxylonis-Quararibeetum
magnificae and Conchocarpo larensis-Coussareetum moritzianae); the Farameo
killipii - Prunion moritzianae alliance contains one subandean forest community
(Croizatio brevipetiolatae-Wettinietum praemorsae) and one Andean forest
community (Schefflero ferrugineae-Cybianthetum laurifolii) and the Ruilopezio
paltonioides-Cybianthion marginati alliance includes one Andean (Geissantho
andini-Miconietum jahnii) and two high Andean forest communities (Gaultherio
anastomosantis-Hesperomeletum obtusifoliae and the Libanothamnetum griffinii).
Altitudinal zonation, forest floristic diversity, composition and forest structure is
discussed between slopes and along the altitudinal gradient and compared, where
possible, to other montane forests. In LMRF, Rubiaceae, Lauraceae and
Melastomataceae are the most speciose of woody families. In UMRF, the
Lauraceae family is still the most diverse, followed by Melastomataceae and
Myrtaceae, while in SARF the Asteraceae and Ericaceae are the most species rich
families. The structure of the montane forests of Ramal de Guaramacal becomes
more compressed towards higher elevations. LMRF are dense and of medium
height, with canopies up to 25 m tall, while UMRF canopies can reach up to 18 m,
and those of SARF are only 6-8 (10) m tall. Basal area was slightly increased on
the North than on the South slopes and shows different patterns against altitude
between slopes. More diversity and density of palms, lianas and climbers is clearly
observed in LMRF, but richness of liana species is also important in SARF
(forests). Forest altitudinal zonation is variable between the North and South
slopes of Guaramacal, with the forest zones of UMRF on the windward South
slope, tending toward reaching lower elevations than on the opposite and drier
North slope. There is a low altitudinal limit of the uppermost forest (Upper Forest
Line or UFL) apparently caused by the “top effect”.
In Chapter 3 zonal páramo vegetation communities present on top of Ramal de
Guaramacal, were studied with the aim to provide a syntaxonomic scheme or
classification, based on analysis of the physiognomy, floristic composition,
ecological relations and spatial distribution of the different vegetation
239
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Flora, vegetation and ecology in the Venezuelan Andes
communities. A total of 91 vascular species, 33 species of bryophytes and 11
species of lichens have been documented from fifty 10 m-line intercept transects,
each surveying 10 m of altititudinal interval on zonal páramo vegetation present
between 2800 and 3100 m altitude. The interpretation of the TWINSPAN
clustering allowed the recognition of five vegetation communities at association
level grouped into two alliances and one order. Three associations of lower
subpáramo or shrubby páramo and two of upper subpáramo or bunchgrass páramo
dominated by rosettes and tussocks have been documented. The alliance Hyperico
paramitanum-Hesperomeletion obtusifoliae groups the shrubby páramo
associations: Ruilopezio paltonioides-Neurolepidetum glomeratae and Disterigmo
acuminatum-Arcytophylletum nitidum, present on wind protected slopes, dwarf
forests edges or along streams. The alliance Hyperico cardonae-Xyridion
acutifoliae groups one widely distributed shrubby páramo association Cortaderio
hapalotrichae-Hypericetum juniperinum and two open grass páramo associations:
Puyo aristeguietae-Ruilopezietum lopez-palacii and Rhynchosporo gollmeriiRuilopezietum jabonensis, present on wind exposed slopes. Asteraceae and
Ericaceae are the most speciose of families, followed by Poaceae and Cyperaceae.
The most diverse genera are Ruilopezia (Asteraceae), Rhynchospora (Cyperaceae)
and Hypericum (Clusiaceae). Diversity of species and growth forms is greater
among the shrubby communities, decreasing in the bunch grass-rosette
communities. Canonical correspondence analysis (CCA) indicates that floristic
composition of zonal vegetation communities is mostly related to slope angle and
altitude than to other observed variables such as pH, soil depth and humus
thickness. The generic and species composition is that of a rain bamboo páramo.
In Chapter 4 the azonal páramo vegetation present at the top of Ramal de
Guaramacal was studied by means of observations, plant collections and surveys
consisting of a total of 71 small plots of between 0.25 to 6 m 2. Azonal vegetation is
represented in the study area by habitats experiencing water stress (peat bogs and
aquatic vegetation). The azonal vegetation present in two peat bogs areas of
Páramo El Pumar (Laguna El Pumar y Laguna Seca), and in a small valley where
water collects in Páramo de Guaramacal, near the „Las Antenas‟ area between
~2900 and ~3000 m were analyzed. A total of 53 morphospecies, belonging to 30
species of vascular plants, 20 species of cryptogams and 3 undetermined species of
algae have been documented for the azonal vegetation. The interpretation of a
TWINSPAN clustering, based on affinities of floristic composition and species
cover, allowed the recognition of six azonal vegetation communities grouped into
three alliances and one order. The new alliance Sphagno recurvi-Paepalanthion
pilosi groups the new bunchgrass association Paepalantho pilosi-Agrostietum
basalis and the both new Sphagnum bog associations: Sphagno recurvi-Caricetum
bonplandii and Sphagno sparsi-Caricetum bonplandii. The new alliance Carici
bonplandii-Chusqueion angustifoliae contains a bamboo páramo („chuscal‟)
association Carici bonplandii-Chusqueetum angustifoliae growing close to the lake
shores, in periodically flooded areas, and characterized almost exclusively by
Chusquea angustifolia. The alliance Ditricho submersi-Isoëtion Cleef 1981 is
represented by the submerged aquatic community of Sphagnum cuspidatum and
the Isoëtetum karstenii Cleef 1981.
240
Summary
_______________________________________________________
Chapter 5 presents the study of the phytogeographical patterns and affinities of the
low altitude and wet páramo vascular flora of Ramal de Guaramacal with emphasis
in to the analysis of the floristic connections of the Guaramacal páramo flora with
the neighboring dry páramos of the Sierra Nevada de Mérida and other páramo
floras of the northern Andes and Central America. A total of 251 vascular plant
taxa belonging to 150 genera and 69 families were recorded from the vegetation
formations existing in the study area. The most species rich families are Asteraceae, Poaceae, Ericaceae and Orchidaceae, followed by the ferns families Grammitidaceae and Lycopodiacae. The most diverse genera are the ferns and fern ally
Elaphoglossum, Huperzia and Hymenophyllum. The analysis of phytogeographical
composition of páramo flora at genus level showed that 52.8% of the genera are
Tropical. The Temperate component is represented by 33.3% of the genera and the
Cosmopolitan component by 13.9%. The Neotropical montane element (38.9%) is
high in Guaramacal páramo, the Páramo endemic element (1.9%) and the Andean
alpine element (0.9% and represented by only one genus (Lachemilla)) are low
compared to other páramo areas. The vascular flora of Páramo de Guaramacal is
largely composed of (1) a group of Neotropical widespread species (31%), (2) a
group of Andean distributed species (49%), part of them confined to the northern
Andes and part widespread in the Andes from Colombia to Bolivia, and (3) a
group of Venezuelan endemics (20%). From an eight páramo flora comparative
dataset, the closest relationships among páramos is observed between the generic
páramo floras of the Colombian Cordillera Oriental of Sumapáz and Sierra Nevada
del Cocuy, which are both closely related to that of the Sierra Nevada de Mérida in
Venezuela. The generic páramo flora of Ramal de Guaramacal shows the closest
relationship to southern Ecuador páramo flora of Podocarpus National Park.
According to Detrended Correspondance Analysis and Principal Component
Analysis ordination results, most of the variations in páramo floras may represent a
response to differences in ambient humidity.
Chapter 6 presents the analysis of functional diversity of mountain forests of
Ramal de Guaramacal as a function of altitude. Decreasing functional diversity is
generally seen as indication of ecosystem degradation. This study aimed to
examine if functional diversity changed with altitude in undisturbed Andean
forests as reference information for studies of degraded Andean systems. We
studied the vascular plant composition of 44 small plots located between 1330 m
and 3060 m in a well-protected forest reserve. We linked each species to their
functional traits related to energy balance and fragmentation, by means of
literature and herbarium studies. Detrended correspondence analysis was used to
detect the principal variation in the trait information. Using fourthcorner analysis,
we randomized the species assemblages in our relevées using two permutation
models, to test if trait composition changed with elevation. Functional trait
diversity was calculated on the basis of species and individuals in the relevées,
using Shannon, Simpson (1-D) and Fisher's alpha indices. Applying the same
permutations models as in the fourthcorner analysis, we tested the relationship of
functional diversity with elevation. Results show that forests in the Ramal de
Guaramacal area became more diverse in the energy balance related traits at higher
elevations, pointing at more prominent levels of overdispersion higher up the
slopes. Leaf size contributed substantially to the altitudinal variation in these traits.
241
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Flora, vegetation and ecology in the Venezuelan Andes
The diversity in fragmentation related traits showed an opposite altitudinal pattern.
Subalpine rain forests (SARF) diverged from the altitudinal trends in
fragmentation related traits, probably as a consequence of edge effects in the
SARF-páramo mosaic, created by wind. We conclude that functional diversity of
undisturbed Andean forests in the Guaramacal area changed with altitude. Global
temperature rises might thus affect the functionality of Andean forests, but not
necessarily in a harmful way.
242
Samenvatting
_______________________________________________________
SAMENVATTING
Ramal de Guaramacal is een ‘outlier’ en lage tot ca. 3130 m hoge bergrug aan het
noordoostelijke einde van de Venezolaanse Andes.
In Hoofdstuk 2 werd de samenstelling van het montane bos van Ramal de
Guaramacal aan beide zijden van de bergrug bestudeerd langs de hoogtegradiënt
en met verschillende hellingexposities. 35 0.1 ha plots met variabele hoogte
intervals van 30 tot 150 m tussen 1350 en 2890 m en 9 plots variabel van afmeting
(50 m2 tot 400 m2) werden bemonsterd in hoogandiene dwergbossen tussen 2850
en 3050 m. In totaal werden 388 morfospecies met dbh≥ 2.5 cm gevonden, die
betrekking hadden op 189 genera en 78 vaatplanten families uit een totaal van 44
bosopname oppervlakken.
De TWINSPAN phytosociologische classificatie gebaseerd op floristische
samenstelling en relatieve abundantie van soorten leidde tot zeven
bosgezelschappen op associatieniveau gegroupeerd in drie verbonden en een
ordegroep van montane bossen. Drie subandiene (LMRF) en vier Andiene –
hoogandiene (UMRF-SARF) bosgezelschappen werden onderscheiden en
beschreven volgens de Zürich-Montpellier methode. (De acronymen SARF,
UMRF en LMRF staan voor respectievelijk Subalpine Rain Forest, Upper
Montane Rain Forest en Lower Montane Rain Forest). Het verbond Geonomo
undatae-Posoquierion coriaceae bevat twee subandiene bosgezelschappen (Simiro
erythroxylonis-Quararibietum magnifoliae en Conchocarpo larensis-Coussareetum
moritzianae), het verbond Farameo killipii-Prunion moritzianae) bevat een
subandiene bosgemeenschap (Croizatio brevipetiolatae-Wettinion praemorsae) en
een andien bosgezelschap (Schefflero ferrugineae-Cybianthetum laurifolii) en het
verbond Ruilopezio paltonoides-Cybianthion marginati) omvat een andien
bosgezelschap (Geissantho andini-Miconietum jahnii) en twee hoogandiene
bosgezelschappen (Gaultherio anastomosantis-Hesperomeletum obtusifoliae en
Libanothamnetum griffinii).
Hoogtezonering, floristische diversiteit van de bossen, samenstelling en
bosstructuur wordt besproken tussen de tegenoverstaande hellingen en langs de
hoogtegradiënt, en waar mogelijk, vegeleken met ander montane bossen.
In LMRF zijn de Rubiaceae, Lauraceae en Melastomataceae de meest soortenrijke
houtige families. In het UMRF is de Lauraceae familie nog steeds de meest
diverse, gevolgd door Melastomataceae en Myrtaceae, terwijk in het SARF de
Asteraceae en Ericaceae de meest soortenrijke families zijn.
De structuur van de bossen van Ramal de Guaramacal wordt naar grotere hoogten
steeds meer gecomprimeerd. LMRF zijn dichte bossen en van en met kronendak
tot 25 m, terwijl UMRF kronendaken tot 18 m reiken en die van SARF slechts 6-8
(10) m hoog.De ‘basal area’ was licht verhoogd op de Noordhelling en vertoont
verschillende patronen in relatie met de hoogte tussen beide hellingen.
IN LMRF is de hogere diversiteit en dichtheid van palmen, lianen en klimmers
duidelijk zichtbaar, maar de lianensoortenrijkdom is ook belangrijk in het SARF.
243
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Flora, vegetation and ecology in the Venezuelan Andes
De altitudinale zonering van de bossen is variabel tussen Noord- en Zuidhellingen
van de Guaramacal bergrug met de UMRF zone op de winderige zuidkant wat
lager voorkomend dan op de andere drogere noordhelling. Er is een altitudinale
lage bovenste bosgrens (Upper Forest Line of UFL), die waarschijnlijk
veroorzaakt wordt door het ‘top effect’.
In Hoofdstuk 3 werden de zonale páramo vegetatie gezelschappen op de toppen
van Ramal de Guaramacal bestudeerd met als doel een syntaxonomisch overzicht
of classificatie gebaseerd op de fysiognomie, floristische samenstelling,
ecologische relaties en ruimtelijke verdeling van de verschillende
vegetatiegezelschappen. In totaal werden 91 vaatplantsoorten, 33 soorten
bryofieten en 11 lichenensoorten gedocumenteerd van 50 10 mlijnintercepttransecten van de zonale páramovegetatie elk met een 10 m hoogte
interval tussen 2800 en 3100 m. De interpretatie van de TWINSPAN clustering
leidde tot het vaststellen van vijf vegetatiegezelschappen op associatieniveau
behorend tot twee verbonden en een orde. Drie associaties van de lage subpáramo
of struikpáramo en twee van de hoge subpáramo of horstgraspáramo gedomineerd
door rozetten en horstgrassen werden onderscheiden. Het verbond Hyperico
paramitanum – Hesperomelion obtusifoliae omvat de struikpáramo associaties:
Ruilopezio paltonioides-Neurolepidetum conglometae en Disterigmo acuminatumArcytophylletum nitidi voorkomend op wind-beschutte hellingen, randen van
dwergbossen en langs beekjes. Het verbond Hyperico cardonae-Xyridion
acutifoliae omvat de wijd verbreide struikpáramo associatie Cortaderio
hapalotriche- Hypericetum juniperinum, alsmede twee open graspearamo
associaties: Puyo arisitguietae- Ruilopezietum lopez-palacii en Rhynchosperoa
gollmeri-Ruilopezietum jabonensis van de wind geëxponeerde hellingen.
Asteraceae en Ericaceae zijn de meest soortenrijke families gevold door de
Poaceae en Cyperaceae. De meest diverse genera zijn Ruilopezia (Asteraceae),
Rhynchospora (Cyperaceae) en Hypericum (Clusiaceae). De diversiteit van soorten
en groevormen is groter in de struikgezelschappen en neemt af naar de open
horstgras-rozet gemeenschappn. Canonical Correspondence Analysis (CCA) geeft
aan dat de floristische samenstelling van de zonale páramo gezelschappen het
meest gerelateerd is aan de hellinghoek en hoogte (boven zeeniveau) dan naar
andere variabelen zoals pH, bodemdiepte en humusdikte.
In Hoofdstuk 4 werd de azonale páramo van de bergrug van Ramal de Guaramacal
bestudeerd met behulp van observaties, plantencollecties en vegetatieopnames van
in totaal 71 vlakken van 0.25 tot 6 m2. De azonale vegetatie in het studiegebied is
vooral gerepresenteerd door habitats met waterstress (venen en watervegetaties).
De azonale vegetatie is vooral bestudeerd in twee veengebieden van Páramo El
Pumar (Laguna El Pumar en Laguna Seca), als ookin een kleine vlakke vallei bij
‘Las Antenas’ in Páramo de Guaramacal tussen 2900 en 3000 m. In totaal werden
53 morfospecies waarvan 30 soorten vaatplanten, 20 soorten cryptogamen en 3
niet gedetermineerde algen aangetroffen in de azonale vegetatie. De interpretatie
van een TWINSPAN clustering, gebaseerd op floristische samenstelling en
bedecking door soorten, resulteerde in zes azonale gezelschappen, behorend tot
drie verbonden en een orde. Het nieuwe verbond Sphagni recurvi-Paepalanthion
pilosi bevat de nieuwe horstgrasassociatie Paepalantho pilosi-Agrostietum basalis
244
Samenvatting
_______________________________________________________
en de beide nieuwe Sphagnumveen associaties: Sphagno recurvi-Caricetum
bonplandii en Sphagno sparsi-Caricetum bonplandii. Het nieuwe Carici
bonplandii-Chusqueion angustifoliae verbond bevat een venige bamboepáramo
associatie (‘chuscal’), Carici bonplandii-Chusqueetum angustifoliae, dicht bij de
meeroevers met periodieke inundatie en vrijwel exclusief gekarakteriseerd door
Chusquea angustifolia.
Het verbond Ditricho submersi-Isoétion Cleef 1981 is gerepresenteerd door het
ondergedoken gezelschap van Sphagnum cuspidatum en het Isoëtetum karstenii
Cleef 1981.
Hoofdstuk 5 heeft betrekking op de studie van de fytogeografische patronen en
verwantschappen van de vaatplantenflora van de lage en natte bamboepáramo van
Ramal de Guaramacal. Hierbij ligt de nadruk op de analyse van de floraconnecties
met de naburige droge páramos van de Sierra Nevada de Mérida en andere páramo
floras in de noordelijke Andes en Centraal Amerika.
In totaal zijn 251 vaatplant taxa behorend tot 150 genera en 69 families
gedocumenteerd van de páramovegetatie van het Guaramacal studiegebied. De
meest soortenrijke families zijn: Asteraceae, Poaceae, Ericaceae en Orchidaceae
gevolgd door de varenfamilies Grammitidaceae en Lycopodiaceae. De meest
diverse genera zijn Elaphoglossum en Hymenophyllum van de varens en Huperzia
van de wolfsklauwfamilie. De analyse van de fytogeografische samenstelling op
genusniveau van de páramoflora toonde aan, dat 52.8% van de genera tot de
Tropische component behoren. De Gematigde componentis vertegenwoordigd
door 33.3% van de genera en de Kosmopolitische component door 13.9%. Het
Neotropisch montane element (38.9%) is hoog in de Guaramacal páramo; het
endemische Páramo element (1.9%) en het Andien-alpiene element (0.9%,
vertegenwoordigd door slechts een genus, Lachemilla) hebben lage warden
vergeleken met andere páramogebieden (behalve Podocarpus Nationaal Park in
Zuid Ecuador). De vaatplantflora van Páramo de Guaramacal bestaat grotendeels
uit (1) een groep van Neotropisch wijd verbreide soorten (31%), (2) een groep van
soorten uit de Andes (49%), waarvan een deel uit de noordelijke Andes en een
ander deel van Colombia tot Bolivia en (3) een groep Venezolaanse endemische
soorten (20%).
Van een comparatieve dataset van acht verschillende páramofloras is de meeste
verwantschap gevonden tussen de generische páramofloras van de Colombiaanse
Cordillera Oriental met Sumapaz en Sierra Nevada del Cocuy, die beide nauw
verwant zijn aan de páramoflora van de Sierra Nevada de Mérida, Venezuela. De
genera van de páramoflora van Ramal de Guaramacal vertoont een nauwe
verwantschap met die van het Nationale Park Podocarpus in Zuid Ecuador. In
overeenstemming met de ordinaties met Detrended Correspondence Analysis
(DCA) en Principal Component Analysis (PCA) lijken de varieties in
fytogeografische properties in de páramoflora een respons te zijn op verschillen in
milieuvochtigheid.
Hoofdstuk 6 tenslotte betreft de analyse van de functionele diversiteit van het
bergbos van Ramal de Guaramacal in relatie tot en als functie van de hoogte.
Afnemende functionele diversiteit wordt in het algemeen gezien als een aanwijzing
245
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Flora, vegetation and ecology in the Venezuelan Andes
van ecosystem degradatie. Deze studie heeft als doel te onderzoeken of functionele
diversiteit verandert met de hoogte in niet verstoorde montane bossen als referentie
informatie voor studies van gedegradeerde andiene systemen. We bestudeerden de
vaatplanten samenstelling van 44 kleine plots tussen 1330 en 3060 m in een goed
beschermd natuur park. Elke soort werd gekoppeld aan hun functionele
kenmerken wat betreft energiebalans en mate van fragmentatie met behulp van
literatuur en herbarium studies. Detrended Correspondence Analysis (DCA) werd
gebruikt om de belangrijkste variatie in de kenmerken set te ontdekken. Met
behulp van ‘fourthcorner’ analyse randomiseerden wij de soortenassemblages in
onze opnamen met gebruik van twee permutatiemodellen om te testen of de
kenmerken samenstelling veranderde met de hoogte. Functionele kenmerken
diversiteit werd berekend op basis van de soorten en individuele planten in de
opnames met behulp van de indexen van Shannon, Simpson (1-D) en Fisher’s
alpha. Door dezelfde permutatiemodellen toe te passen als in de ‘fourthcorner’
analyse testten we de associatie van functionele diversiteit met de hoogte.
De resultaten laten zien dat de bossen van de Ramal de Guaramacal meer divers
worden op grotere hoogten wat betreft kenmerken verband houdend met de
energiebalans die zelfs tot opvallende niveaus van overdispersie leiden op de
bovenste hellingen. De bladgrootte droeg substantieel bij tot de altitudinale
variatie van deze kenmerken. De kenmerken verband houdend met fragmentatie
vertoonden juist een tegenover gesteld beeld. Subalpiene regenbossen (SARF)
weken af van de altitudinale trends van de kenmerken gerelateerd aan
fragmentatie, vermoedelijk als gevolg van randeffecten in het SARF-páramo
mozaiek, veroorzaakt door de wind.
We concluderen dat de functionele diversiteit van niet-verstoorde montane bossen
in het Guaramacal gebied veranderde met de hoogte. Globale
temperatuurverhoging zou dus de functionaliteit van Andes bossen kunnen
aantasten, maar niet noodzakelijkerwijs op een schadelijke wijze.
246
Resumen
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RESUMEN
El Ramal de Guaramacal es una pequeña ramificación montañosa con altura
máxima de 3130 m, ubicada al extremo nor-oriental de los Andes venezolanos.
En el Capítulo 2 se estudiaron las comunidades de bosques montanos del Ramal de
Guaramacal, Andes, Venezuela, a lo largo de un gradiente altitudinal y entre
diferentes vertientes. Se analizaron treinta y cinco parcelas de 0.1-ha ubicadas, con
intervalos variables de 30 m a 150 m, entre 1350 m y 2890 m de altitud, y nueve
parcelas de tamaño variable (50 m2 hasta 400 m2) ubicadas entre 2800-3050 m. De
las 44 parcelas estudiadas, se registró un total de 388 morfoespecies con DAP ≥2.5
cm correspondientes a 189 géneros y 78 familias de plantas vasculares. La
clasificación fitosociológica mediante TWINSPAN basado en la composición
florística y abundancia relativa de las especies, reveló siete comunidades de
bosque, agrupadas en tres alianzas y un grupo bosques montanos equivalente a
orden. Se distinguen y se describen, según la metodología Zürich-Montpellier, tres
comunidades de bosque subandino (LMRF), y cuatro comunidades de bosque
andino/alto-andino (UMRF-SARF). La allianza Geonomo undatae-Posoquerion
coriaceae incluye dos comunidades de bosque subandino (Simiro erythroxylonisQuararibeetum magnificae y Conchocarpo larensis-Coussareetum moritzianae); la
alianza Farameo killipii-Prunion moritzianae incluye una comunidad de bosque
subandino(Croizatio brevipetiolatae-Wettinietum praemorsae) y una de bosque
andino(Schefflero ferrugineae-Cybianthetum laurifolii) y la alianza Ruilopezio
paltonioides-Cybianthion marginatii incluye una comunidad de bosque andino
(Geissantho andini-Miconietum jahnii) y dos comunidades de bosque altoandino
(Gaultherio anastomosantis-Hesperomeletum obtusifoliae y el Libanothamnetum
griffinii). Se discuten la zonificación altitudinal, diversidad y composición
florística y estructura del bosque con respecto a la altitud y se compara, cuando
posible, con otros bosques de montaña. En el bosque subandino, las familias de
plantas leñosas más diversas en especies son Rubiaceae, Lauraceae y
Melastomataceae. En el bosque andino, Lauraceae es la familia más diversa,
seguido de Melatomataceae y Myrtaceae, mientras que en el bosque alto-andino
las familias con mayor riqueza de especies son Asteraceae y Ericaceae. La
estructura de los bosques montanos del Ramal de Guaramacal se comprime hacia
las partes más altas. Los bosques subandinos son densos y de altura media, con un
dosel hasta 25 m de alto, mientras que en el bosque andino el dosel puede alcanzar
hasta 18 m y en el bosque alto-andino el dosel alcanza solo 6-8 (10) m de alto. El
área basal se encontró ligeramente mayor en la vertiente Norte que en la Sur y
presenta patrones diferentes respecto a la altitud en cada vertiente. En los bosques
subandinos se observa claramente mayor diversidad y densidad de palmas, lianas y
trepadoras, pero en el bosque alto-andino la riqueza de especies de lianas es
también importante. La zonificación altitudinal del bosque varía entre las
vertientes Norte y Sur de Guaramacal, observándose que la zona de bosque andino
o montano alto tiende a alcanzar altitudes menores en la vertiente sur más húmeda
que en la vertiente Norte más seca. El límite superior del bosque en el Ramal de
Guaramacal es bajo, aparentemente causado por el efecto de cumbre.
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Flora, vegetation and ecology in the Venezuelan Andes
En el Capítulo 3 se presenta el estudio de las comunidades de vegetación de
paramo zonal existentes en las cimas del Ramal de Guaramacal, con el fin de
proporcionar un esquema de clasificación sintaxonómico basado en el análisis de
la fisonomía, composición florística, relaciones ecológicas y distribución espacial
de las diferentes comunidades vegetales. Mediante el levantamiento de cincuenta
líneas de intersección de 10 m de largo, establecidas a cada 10 m de altitud, en la
vegetación zonal entre 2800 y 3100 m, se encontraron un total de 91 especies de
plantas vasculares, 33 de briofitas y 11 de líquenes. La interpretación de la
clasificación con TWINSPAN permitió reconocer cinco comunidades de
vegetación zonal al nivel de asociación, agrupadas en dos alianzas y un orden. Tres
de las asociaciones son del subpáramo bajo arbustivo y dos del subpáramo alto con
pajonal, dominadas por rosetas y hierbas en macollas. La alianza Hyperico
paramitanum-Hesperomeletion obtusifoliae agrupa las asociaciones de subpáramo
arbustivo: Ruilopezio paltonioides-Neurolepidetum glomeratae and Disterigmo
acuminatum-Arcytophylletum nitidum, presentes en vertientes protegidas del
viento, a los bordes de bosques enanos o a lo largo de cursos de agua. La alianza
Hyperico cardonae-Xyridion acutifoliae agrupa una asociación de subpáramo
arbustivo, la asociación Cortaderio hapalotrichae-Hypericetum juniperinum,
ampliamente distribuida en el área, y dos asociaciones de subpáramo alto con
pajonal: Puyo aristeguietae-Ruilopezietum lopez-palacii, y Rhynchosporo
gollmeri–Ruilopezietum jabonensis, presentes sobre vertientes expuestas. Las
familias Asteraceae y Ericaceae son las más ricas en especies seguido de Poaceae
y Cyperaceae. Los géneros más diversos son Ruilopezia (Asteraceae),
Rhynchospora (Cyperaceae) e Hypericum (Clusiaceae). Tanto la diversidad de
especies como de formas de crecimiento es mayor en las comunidades arbustivas y
disminuye en los pajonales-acaulirrosuletales. El análisis de correspondencia
canónica (CCA) indica que la composición florística de las comunidades de
vegetación zonal del Páramo de Guaramacal se relaciona principalmente con la
pendiente y altitud, más que con otras variables observadas como profundidad y
pH de los suelos. La composición genérica y de especies es propia de un páramo
muy húmedo de bambúes.
En el Capítulo 4, se estudió la vegetación de páramo azonal presente en la cima del
Ramal de Guaramacal, mediante observaciones, colecciones botánicas y muestreos
de un total de 71 pequeñas parcelas de tamaño entre 0.25 a 6 m2. La vegetación
azonal está representada en el área de estudio por habitats donde existe un estrés
por exceso de agua (turberas y vegetación acuática). Se analizó las vegetaciones
azonales presente en dos áreas de turberas del Páramo El Pumar (Laguna El Pumar
y Laguna Seca) y en un pequeño valle con acumulación de agua cerca del área de
‘Las Antenas’ del Páramo de Guaramacal, ubicadas entre aprox. 2900 y 3000 m de
altitud. Se documentó un total de 53 morfoespecies correspondientes a 30 especies
de plantas vasculares, 20 de briofitas y líquenes y 3 especies indeterminadas de
algas presentes en la vegetación azonal. La interpretación de la clasificación de
TWINSPAN, basada en afinidades de composición florística y cobertura de
especies, permitió reconocer seis comunidades de vegetación azonal agrupadas en
tres alianzas y un orden. La alianza nueva Sphagno recurvi-Paepalanthion pilosi
agrupa la asociación nueva de pajonal de páramo Paepalantho pilosi-Agrostietum
basalis y las dos asociaciones nuevas de turberas de Sphagnum: Sphagno recurvi248
Resumen
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Caricetum bonplandii y Sphagno sparsi-Caricetum bonplandii. La alianza nueva
Carici bonplandii-Chusqueion angustifolia contiene una asociación de páramo de
bambues (‘chuscal’), Carici bonplandii-Chusqueetum angustifoliae, que crece
cerca de las orillas de las lagunas, en áreas inundadas periódicamente,
caracterizada casi exclusivamente por la presencia de Chusquea angustifolia. La
alianza Districho submersi-Isoëtion Cleef 1981 está representada en el área de
estudio por la comunidad acuática sumergida de Sphagnum cuspidatum y la
asociación Isoëtetum karstenii Cleef 1981.
El Capítulo 5 presenta el estudio de los patrones fitogeográficos y afinidades de la
flora vascular de páramo húmedo y de baja altitud del Ramal de Guaramacal, con
énfasis en el análisis de sus conexiones florísticas con páramos secos cercanos de
la Sierra Nevada de Mérida y otras floras de páramo de los Andes del Norte y
Centroamérica. Un total de 251 taxa de plantas vasculares pertenecientes a 150
géneros y 69 familias se han registrado en el área de estudio. Las familias más
ricas en especies son Asteraceae, Poaceae, Ericaceae y Orchidaceae, seguido por
las familias de helechos Grammitidaceae y Lycopodiacae. Los géneros más
diversos son Elaphoglossum, Huperzia, Hymenophyllum y Chusquea. El análisis
de composición fitogeográfica a nivel de género de la flora de páramo mostró que
52,8% de los géneros son Tropical. El componente Templado está representado
por 33,3% de los géneros y el componente Cosmopolita está representado por
13,9%. El elemento Montano Neotropical (38.9%) es alto en el páramo de
Guaramacal, los elementos Endémico de Páramo (1,9%) y Alpino Andino (0,9%),
representado por sólo un género (Lachemilla), son bajos comparado con otros
páramos. La flora vascular de Páramo de Guaramacal está integrada en gran
medida por (1) un grupo de especies de distribución amplia Neotropical (31%), (2)
un grupo de especies de distribución Andina (49%), parte de ellos se limita a los
Andes del Norte y parte generalizada en los Andes desde Colombia hasta Bolivia y
(3) un grupo de especies endémicas de Venezuela (20%). De la comparación del
conjunto de datos de flora de ocho páramos, las relaciones más cercanas entre los
páramos se observa entre las floras genéricas de los páramos de la Cordillera
Oriental colombiana, Sumapáz y Sierra Nevada del Cocuy, los cuales están
estrechamente relacionadas con Sierra Nevada de Mérida en Venezuela. La flora
genérica de páramo del Ramal de Guaramacal muestra la relación más cercana con
la flora de páramo de la Reserva de Biósfera Podocarpus al sur del Ecuador. Según
los resultados de ordenación DCA y PCA, la mayoría de las variaciones en las
floras de los páramos analizados pueden representar una respuesta a diferencias de
humedad ambiental.
El Capítulo 6 presenta el análisis de la diversidad funcional de los bosques
montanos del Ramal de Guaramacal en relación y en función de la altitud. La
disminución de la diversidad funcional es generalmente vista como una indicación
de degradación del ecosistema. Este estudio pretende examinar si la diversidad
funcional cambió con la altitud en bosques andinos sin intervención antropica,
como información de referencia para estudios de sistemas Andinos degradados. Se
estudió la composición de plantas vasculares de 44 parcelas pequeñas situadas
entre 1330 m y 3060 m en una reserva de bosques bien protegidos. Se vinculó cada
especie a sus rasgos funcionales relacionados con balance energético y
fragmentación, por medio de estudios de literatura y de material de herbario.
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Flora, vegetation and ecology in the Venezuelan Andes
Análisis de correspondencia linearizadas (DCA) fue usado para detectar la
variación principal en la información de los rasgos. Se utilizó análisis de
fourthcorner, para aleatorizar los ensambles de especies en los relevés usando dos
modelos de permutación, para probar si la composición de rasgos cambió con la
elevación. La diversidad de rasgos funcionales se calculó sobre la base de las
especies y los individuos en los relevés, mediante los índices de Shannon, Simpson
(1-D) y alfa de Fisher. Al aplicar los mismos modelos de permutaciones, como en
el análisis de fourthcorner, se probó la asociación de la diversidad funcional con
elevación. Los resultados muestran que los bosques en la zona del Ramal de
Guaramacal se vuelven más diversos hacia las elevaciones más altas, en aquellos
rasgos funcionales relacionados con balance energético, apuntando a los niveles
más prominentes de sobredispersión en las partes más altas de las laderas. El
tamaño de las hojas contribuye sustancialmente a la variación altitudinal en estos
rasgos. La diversidad en los rasgos funcionales relacionados con la fragmentación
mostró un patrón altitudinal opuesto. Los bosques húmedos altoandinos o bosques
húmedos subalpinos (SARF) difieren en las tendencias altitudinales de los rasgos
relacionados con fragmentación, probablemente como consecuencia de los efectos
de borde creado por el viento en el mosaico de bosques enanos de SARF-páramo.
Se concluye que la diversidad funcional de los bosques andinos inalterados en el
área de Guaramacal cambia con la altitud. Por lo tanto, los aumentos de
temperatura global podrían afectar a la funcionalidad de los bosques andinos, pero
no necesariamente de una manera perjudicial.
250
Acknowledgements
_______________________________________________________
ACKNOWLEDGEMENTS
I wish to express my deeply appreciation to all those people that have supported
me during the time I have been doing research in Guaramacal. However, there are
so many people that have helped me in many different ways and occasions that I
am afraid I would fail to mention every one. So, here I will refer mainly to those
people who have been most helpful during the last four years that I have been
working for completing this PhD thesis. For all those people I omitted their names,
receive my apologies for that but also my sincere gratitude for their support.
First, I am deeply grateful to my promotor Prof. Dr. Antoine Cleef and copromotor Dr. Joost Duivenvoorden for their friendship and support, without them,
it had not been possible for me to complete this thesis. Antoine has been a great
tutor in both professional and personal aspects, from the very moment he accepted
to be my promotor, despite of the distance, he has been always available for
communication, attending my questions, reviewing my manuscripts, guiding and
encouraging me with enthusiasm and providing me great and valuable ideas. I
appreciate also his support during my stays in Amsterdam, where he not only
taught me a lot, working with my data, providing me literature and discussing
ideas, but also he has been so thoughtful helping me and my family finding the
best place to live and making us to feel at home. In summary, it has been a
tremendous experience and a great honor working with Antoine. Joost has been
also very supportive and inspiring. He taught me with a great patience and brilliant
skills how to work with some technology tools for analysis ecological data to a
high level of abstraction and to interpret results to obtain meaningful information. I
feel privileged to share his innovative ideas and co-authoring a manuscript with
him.
I thank to my other promotor Prof. Dr. Henry Hooghiemstra for his support and for
reviewing and providing comments that help to improve some chapters of this
thesis. I thank also to the IBED secretaries staff and colleagues for their opportune
collaboration when required. Special thanks to Jody Dos Santos, Ada Hoogendorp
and Mary Parra. Marcela Moscol gave me a friendly support during my stays in
Amsterdam providing me always with useful tips.
My family and I enjoyed and appreciate very much the hospitality and invitations
we received during our stays in The Netherlands. Specially, the exquisite dinners
we shared with Antoine and friends at the Indonesian restaurant. Visiting Harlem
and sharing with Joost and his family in a lovely evening at their home. Also, the
kind and delightful evenings we pass with Paul Maas and Hiltje Maas van de
Kramer at their home in Bunnik. We thank to the Gijs Haverkate family for
lending us their house and making us to feel it as our home.
I am indebted to the UNELLEZ (grants SEI-23195107, SEI-23105102); CONICIT
(grant S1-97001662) and FONACIT (grant PEM-2001002165) which have
supported fieldwork and equipment for this study. UNELLEZ also granted me
permission and financial support for all my visits to Amsterdam to work on my
PhD program. Alberta Mennega Fund (Utrecht University) is acknowledged for
the financial contribution to my stay at IBED, in my visit to Amsterdam in 2007.
251
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INPARQUES and MARN are thanked for the corresponding permits, as well as
the Superintendente of Guaramacal National Park TSU Amilcar Bencomo for his
constant collaboration. I also thank to all the staff of park rangers of Guaramacal
National Park, for being so eager to help when required, among them Ramon
Aldana, Yelitza Briceño, Yilson Camacho, Jorge Rivero, Amabilis Teran. I
specially thank to the park rangers Ramón Caracas and Luis Zambrano† who
collaborated in most of the field trips.
Many helpers assisted in fieldwork including students, park rangers, villagers,
colleagues, brothers and friends. Special thanks to Wilfredo Albarran, Karina
Bastidas, Oscar León, Luis A. Linarez, Pedro Tovar and Máximo Valladares for
their solidarity and recurrent field assistant during the last four years.
I am deeply thankful to the Herbario PORT staffs which have been very supportive
during these past four years. I am grateful to Angelina Licata, who made all profile
vegetation illustrations. Angelina has been as an older sister to me, always very
supportive. She also helped me with species identifications and herbarium
specimens’ curator. Rosalinda Parra and Elida Mendez were very helpful with the
specimens processing and management. During these past four years, Mannelly
Ramírez, José Farreras and Luis Miguel Leonido, have been very collaborative
when required, with my teaching and other related activities at UNELLEZ.
I appreciate the work done by Basil Stergios (UNELLEZ), Laurence Dorr (US)
and Miguel Niño (UNELLEZ), who have also collected plants in Guaramacal and
surrounded mountains, contributing with a valuable herbarium reference collection
for my specimen identifications. G. Davidse (MO) and S. Laegaard (AAU) were
helpful with the identification of some selected grasses. I thank also D. Griffin III
(FLAS), Guido van Rennen (Amsterdam) and Juan Carlos Benavides (Colombia)
for identification of bryophytes and H.J.M. Sipman (B) for lichens identification.
Ross D. Morrison (University of Leicester, UK) kindly corrected and improved the
English text of chapters 2-4. Beryl Simpson (Austin) provided important
comments and language editing on the earlier version of the manuscript of chapter
5.
Finally, but the most important, I thank to my family for their support. My
husband Gerardo Aymard was the first person to push me to pursuit this PhD and
hold me all the time during this last years. Gerardo has been very collaborative
with literature finding and discussing ideas to contribute in one of my thesis’s
chapters, while being also very supportive and affective at home. My daughter
Marianne has been so understanding with her mom’s work, giving me no more
than happiness, pride and satisfaction of having a great behaved, responsible and
excellent scholar teenager, while I have been so busy working on my PhD. My
mother, brothers and sisters have always been very supportive. I specially thank to
my niece Karim Rodriguez Cuello for designing the cover of this thesis.
Working in Guaramacal have been a passion to me, still there are many things I
wish to study there. I thank God for giving me life and the chance to be in this
marvelous and amazing natural place, which I wish to be conserved forever. To all
those people that have helped me in any way to work in Guaramacal, thank you
very much.
252
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Curriculum Vitae
CURRICULUM VITAE
Nidia Lourdes Cuello Alvarado was born on 25 of October of 1964 in
Barquisimeto, Venezuela. She received her BSc degree in Natural Resource
Engineering at the Universidad Nacional Experimental de los Llanos Ezequiel
Zamora, Venezuela, in February 12, 1988, obtaining first place of a promotion of
twelve. Her research project to obtain BSc was entitled “Caracterizacion
florístico-estructural de la vegetación de un sector de la cuenca media del Río
Portuguesa, Edo. Portuguesa, Venezuela”. After that, during two years, she was
based at the Herbario Universitario PORT of UNELLEZ participating as assistant
to a project for inventory of natural resources in the Venezuelan Guayana region
(Proyecto Inventario de los Recursos Naturales de la Región Guayana P.I.R.N.R.G) conducted by Corporación Venezolana de Guayana (C.V.G.TECMIN, C.A.) in the job of collecting, processing and identification of botanical
specimens from the Venezuelan Guayana. During that time she had the
opportunity to stay six months at The Missouri Botanical Garden receiving
training and working on identification of botanical specimens from the Venezuelan
Guayana for the mentioned project. Since then, she got involved as contributor for
preparing floristic manuscript of some legume and Clusiaceae genera for the Flora
of Venezuelan Guayana project. In 1990, she got a position as Instructor professor
in Botany at Universidad Nacional Experimental de los Llanos Ezequiel Zamora,
where she currently works as Titular Professor. She obtained her MSc in Biology
at University of Missouri - St. Louis in May 18, 1997. Her research title for her
MSc Thesis was: Floristic Diversity and Structure of the montane cloud forests of
Cruz Carrillo National Park in the Venezuelan Andes. She also obtained a
Graduate Certificate in Tropical Biology and Conservation at the University of
Missouri-St. Louis in January 12, 1997. Since 1997 to present day, she has been in
charge of the direction of Herbarium PORT of the UNELLEZ, where she has
coordinated grants from FONACIT, Conservation International and The A.W.
Mellon Foundation for herbarium support on collection data basing. At
UNELLEZ, she has also coordinated and teaching the course of Botany for the
Academic Program of Natural Resources Engineering. During her professional life
she has attended and participated in different symposia and meetings, presenting
her works in ten international events and in sixteen national events in Venezuela.
She has been accredited by Venezuelan system of scientific researcher’s promotion
(Programa de Promocion al Investigador -PPI) since 1997. She has also been
awarded with grants from Alberta Mennega Stichting, Elizabeth Bascom (Missouri
Botancial Garden), Smithsonian Institution, CONICIT, FONACIT. UNELLEZ,
Fundación Polar, International Centre for Tropical Ecology (ICTE) of the
University of Missouri-St.Louis. Since 1995 she has been doing research in Ramal
de Guaramacal in the Venezuelan Andes, initially with support from the related
project Flora of Guaramacal, jointly conducted by Basil Stergios (UNELLEZ) and
Laurence Dorr (NMNH of Smithsonian Institution), for her MSc degree Thesis.
Later, she developed her own research project with grants from UNELLEZ and
FONACIT for geobotanical exploration and floristic surveys of the vegetation
types occurring in Guaramacal area. Results of this work have been used in part
for the completion of her PhD thesis.
253
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Flora,
vegetation and ecology in the Venezuelan Andes
Publications
Duivenvoorden, J. F. and Cuello, N. (submitted). Functional diversity changes
with altitude in Andean forests in Venezuela. Global Ecology and Biogeography.
Cuello, N., Cleef, A.M. & Aymard G. (submitted). Phytogeography of the vascular
páramo flora of Ramal de Guaramacal, Andes, Venezuela. Anales del Jardín
Botánico de Madrid.
Cuello, N. & Cleef, A. M. 2009. The páramo vegetation of Ramal de Guaramacal,
Trujillo state, Venezuela. 2. Azonal vegetation. – Phytocoenologia 39 (4): 389409.
Cuello, N. & Cleef, A. M. 2009. The páramo vegetation of Ramal de Guaramacal,
Trujillo state, Venezuela. 1. Zonal vegetation. Phytocoenologia 39 (3):295-329.
Cuello, N. & Cleef, A. M. 2009. The forest vegetation of Ramal de Guaramacal in
the Venezuelan Andes. Phytocoenologia 39(1):109-156.
Cuello, N. & G. Aymard. 2008. Ilex guaramacalensis, a new species
(Aquifoliaceae) from the Ramal de Guaramacal in the Venezuelan Andes. Novon
18: 319-324.
Solórzano, N., F. Romero y N. Cuello. 2006 (2003). Potencial forrajero de los
bosques de Mesa de Cavacas, estado Portuguesa, Venezuela. Rev. Unellez Ciencia
y Tecnologia 21:1-17.
Cuello, N. 2004. A New Vining Species of Swartzia (Fabaceae) from Venezuelan
Amazon. Novon 14:420-423.
Cuello, N. 2004. Los bosques del Parque Nacional Guaramacal, estado Trujillo,
Venezuela: Testigos del desarrollo sostenible dentro de la región Andina y
Llanera. Memorias del IV Simposio Internacional de Desarrollo Sustentable en los
Andes, AMA-Mérida 2001. La Estrategia Andina para el Siglo XXI (CD-Rom).
sesión IV. Taller sobre selvas y bosques nublados.
Aymard, G. & N. Cuello. 2004. Two new species of Aegiphila (Verbenaceae) from
Venezuela and Brazil. Novon 14: 20-24.
Cuello, N. 2003. A New Species of Tovomita (Clusiaceae) from the Venezuelan
and Peruvian Amazon Region. Novon 13:34-36.
Cuello, N. 2002. Altitudinal Changes in Forest Diversity and Composition in the
Ramal de Guaramacal in the Venezuelan Andes. Ecotropicos 15(2):160-176.
Cuello, N. 2001. Swartzia humboldtiana (Fabaceae), una nueva especie del Rio
Casiquiare, estado Amazonas, Venezuela. BioLlania, Ed. Esp. N° 7.
Dorr, L., B. Stergios, A. Smith & N. Cuello. 2000.[2001] Catalogue of the
Vascular Plants of Guaramacal National Park, Portuguesa and Trujillo States,
Venezuela. Contributions from the United States National Herbarium 40:1-155.
Washington, DC. 155 pp.
Cuello, N. 1998 [2000]. Caracterización de los bosques montanos del Parque
Nacional Cruz Carrillo en los Andes de Venezuela. Memorias del IV Congreso
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Publications
Latinoamericano de Ecología: Ecología y Desarrollo Sostenible: Reto de America
Latina para el Tercer Milenio. Arequipa, Perú. Pp.127-130.
Cuello, N. (ed.) 1999 [2000]. Parque Nacional Guaramacal. Unellez - Fundación
Polar, Caracas, Venezuela. 245 pp., figs., fotos a color. ISBN: 980-248-099-I, 980379-003-X.
Cuello, N. y F. Romero. 1999 [2000]. Introducción. pp. 1-6. En: Parque Nacional
Guaramacal, N. Cuello (ed.). Unellez - Fundación Polar, Caracas, Venezuela.
Cuello, N. y O. Barbera. 1999 [2000]. Aspectos Climáticos del Parque Nacional
Guaramacal. pp. 47-49. En: Parque Nacional Guaramacal, N. Cuello (ed.).
Unellez - Fundación Polar, Caracas, Venezuela.
Cuello, N. 1999 [2000]. Observaciones sobre la vegetación del Parque Nacional
Guaramacal. pp. 105-117. En: Parque Nacional Guaramacal, N. Cuello (ed.).
Unellez - Fundación Polar, Caracas, Venezuela.
Cuello, N. 1999 [2000]. La Unellez en el Parque Nacional Guaramacal. pp. 183191. En: Parque Nacional Guaramacal, N. Cuello (ed.). Unellez - Fundación
Polar, Caracas, Venezuela. (recopilación).
Aymard, G., L. Dorr & N. Cuello. 1999. Rugea tayloriae (Rubiaceae) a new
species from montane forests of Guaramacal, Trujillo, Venezuela. Novon 9:315317.
Cuello, N. 1999. Two new distinctively large-leaved species of Tovomita
(Clusiaceae) from the Venezuelan and Peruvian amazonian region. Novon 9:150152..
Aymard, G., N. Cuello, & R. Schargel. 1998. Floristic composition, structure, and
diversity in moist forest communities along the Casiquiare Channel, amazonas
state, Venezuela. Pp. 499-510. In: F. Dallmeier & J. Comisky (eds.), Proceedings
of the Smithsonian Institution/Man and the Biosphere. Symposium on Measuring
and Monitoring Forest Biological Diversity. Whashington, DC.
Pennington, R. B., G. Aymard & N. Cuello. 1997. A new species of Andira
(Leguminosae, Papilionideae) from the Venezuelan Guayana. Novon 7:72-74.
Cuello, N. 1997. Floristic Diversity and Structure of the montane cloud forests of
Cruz Carrillo National Park in the Venezuelan Andes. Master Thesis, University of
Missouri-St. Louis, U.S.A.
Aymard, G. & N. Cuello. 1995. Two new species of the genus Sterigamapetalum
(Rhyzophoraceae) from Venezuela and Brazil. Novon 5:223-226.
Cuello, N. 1994. Lectotipificación y nuevo estatus de Desmodium orinocense
(DC.) Cuello (Leg.- Pap.). Novon 4(2):98-99.
Aymard, G. & N. Cuello. 1994. Meliosma gentryi Aymard & Cuello (Sabiaceae)
una nueva especie para la flora de la Guayana venezolana, BioLlania 10:30-35.
255
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Flora,
vegetation and ecology in the Venezuelan Andes
Aymard, G. y N, Cuello. 1991. Catalogo y adiciones a las especies Neotropicales
del género Canavalia. (Leg.-Pap.). En: Memorias del Semimario-Taller de Trabajo
Internacional sobre Canavalia:1-62. Ceniap-Fonaiap-UCV. Maracay. Venezuela.
Aymard, G. y N. Cuello. 1991. Dos nuevas especies neotropicales del género
Canavalia (Leguminosae - Papilonaceae - Phaseoleae - Diocleinae). BioLlania
8:87-92.
Cuello, N. y G. Aymard. 1991. Contribuciones a la Flora del estado Portuguesa,
Venezuela: El género Desmodium (Leguminosae-Papilionoideae). Biollania 8:4759.
Cuello, N. y G. Aymard, 1991. Rinoreocarpus ulei (Melchior) Ducke. (Violaceae),
un genero y especie nuevo para la Flora de Venezuela. BioLlania 8:111-115.
Cuello, N., T. Killeen y C. Antezana. 1991. "Linea de Intercepción" (Lineintercept), una metodología apropiada para el estudio de sabanas tropicales. In:
Miranda L. C. and E. Castellano (eds.) Memoria del I Curso Internacional sobre
vegetación y ecología tropical con énfasis en los métodos de estudio de la
vegetación. TCA. Bolivia. 15 p.
Aymard, G. y N. Cuello. 1990. Revisión del género Canavalia (Leg.-Pap.) para
Venezuela. Recursos Tropicales para la alimentación animal. Vol. 1(4). Facultad
de Agronomía. UCV, Venezuela. 40 p.
Aymard, G., N. Cuello & A. Fernández. 1990. Observaciones sobre el hallazgo de
Cinchona amazonica Standl. (Rubiaceae) en la Guayana venezolana. BioLlania
7:125-130.
Cuello, N., G. Aymard & B. Stergios. 1988. Observaciones sobre la vegetación de
un sector de la cuenca media del Río Portuguesa. Edo. Portuguesa. Venezuela.
BioLlania 6:163-193.
Aditionally, 32 contributions to floras and catalogues, 1 book review, 20 technical
reports.
256