Phytocoenologia Vol.

45 (2015), Issue 12, 195201

Stuttgart, July 2015
Section Ecoinformatics
Long Database Report

VegPramo, a flora and vegetation database for the Andean pramo

Gwendolyn Peyre*, Henrik Balslev, David Mart, Petr Sklen, Paul Ramsay, Pablo Lozano, Nidia Cuello, Rainer
Bussmann, Omar Cabrera & Xavier Font

Abstract: The pramo region in the northern Andes is very biodiverse, presents high endemism and provides
many ecosystem services. Unfortunately, the pramo is critically threatened by anthropogenic activities and
climate change. Further research and development of efficient conservation strategies are therefore needed for
the region, but they are often limited by the lack of consistent biological data-sources. Here we present Veg-
Pramo (GIVD ID: SA-00-002, http://www.givd.info/ID/SA-00-002), a flora and vegetation database for the
pramo based on phytosociological vegetation plots. VegPramo contains data from 3,000 georeferenced vege-
tation plots with updated nomenclature. The database is accessible through the webportal http://www.
vegparamo.com, from which floristic and vegetation data can be freely consulted and downloaded. This new
tool should make future botanical and ecological pramo studies easier. VegPramo is already geographically
and floristically representative for the pramo region, but we hope it will continue to grow in scientific signifi-
cance via new data addition and revision.
Keywords: Andes; biological database; floristic record; Neotropics; vegetation plot; webportal.
Submitted: 12 March 2015; revised version submitted: 7 May 2015; accepted: 13 May 2015
Co-ordinating Editor: Florian Jansen

GIVD Fact Sheet

GIVD Database ID: SA-00-002 Last update: 2015-06-01
VegPramo Web address: http://www.vegparamo.com
Database manager(s): Gwendolyn Peyre (g.peyre@ub.edu)
Owner: Gwendolyn Peyre; Ut. Botany, Dpt. Plant Biology; Fac. Biology, University of Barcelona
Av. Diagonal 645, 08028 Barcelona, Spain
Scope: Phytosociological data from Andean Pramo ecosystems.
Availability: free online Online upload: no Online search: yes
Database format(s): MySQL Export format(s): plain text file, b-VegAna XML
Plot type(s): normal plots Plot-size range: (1) 10100 (400) m
Non-overlapping plots: Estimate of existing plots: Completeness: Status:
3,000 NA [NA] ongoing capture
Total no. of plot observations: Number of sources (biblioreferences, data collectors): Valid taxa:
3,000 38 2,220
Countries: CO: 52.0%; EC: 33.0%; PE: 3.0%; VE: 13.0%
Forest: 0% Non-forest: aquatic: 0%; semi-aquatic: 0%; arctic-alpine: 100%; natural: 0%; semi-natural: 0%; anthropogenic: 0%
Guilds: all vascular plants: 100%
Environmental data: altitude: 97%; slope aspect: 62%; slope inclination: 72%; surface cover other than plants (open soil, litter, rock etc.): 42%
Performance measure(s): cover: 100%
Geographic localisation: GPS coordinates (precision 25 m or less): 10%; point coordinates less precise than GPS, up to 1 km: 81%; small grid
(not coarser than 10 km): 8%
Sampling periods: 19501959: 1.0%; 19701979: 1.0%; 19801989: 16.0%; 19901999: 16.0%; 20002009: 46.0%; 20102019: 17.0%;
unknown: 5.0%
Information as of 2015-06-01; further details and future updates available from http://www.givd.info/ID/SA-00-002

*Corresponding authors address: Department of Plant Biology, Faculty of Biology, University of Barcelona, Av. Diagonal 645, 08028
Barcelona, Spain; g.peyre@ub.edu. Complete addresses of all authors can be found at the bottom of the paper.

2015 Gebrder Borntraeger, 70176 Stuttgart, Germany www.borntraeger-cramer.de

DOI: 10.1127/phyto/2015/0045 0340-269X/2015/0045 $ 3.15

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 196
Introduction
The pramo is a biogeographic region that includes all
natural and semi-natural ecosystems located between the
montane treeline and the permanent snowline (approx.
30005000 m a.s.l.) in the humid Andes of Peru, Ecuador,
Colombia and Venezuela (Luteyn 1999). Its recent ori-
gin, vast extension and multiple influences from the adja-
cent coastal plains and Amazon basin, have led to the
richest tropical alpine flora in the world with over 3,400
species of vascular plants and 1,300 of non-vascular plants
(Sklen et al. 2014). The pramo is considered a fast
evolving biodiversity hotspot (Madrian et al. 2013) with
very high endemism (Luteyn 1999). The pramo vegeta-
tion is also extraordinarily diverse with bunch grasslands,
shrublands, bamboo communities, meadows, rosette
communities, forests, cushion communities and high-al-
titude deserts among others (e.g. Rangel-Churio 2000).
This very diverse region provides numerous ecosystem
services for a large human population, especially regard-
ing water storage and supply. However, the pramo is
ecologically fragile and its homeostasis is critically threat-
ened by the combined effects of anthropogenic activities
at local scale, mostly in form of agriculture and pasture,
and climate change at global scale (Hofstede et al. 2003).
Tropical ecosystems are in general under-studied,
which contrasts with their high biodiversity and often
critical vulnerability that should capture most scientific
attention (Field et al. 2009). Botanical and ecological re-
search on the pramo is relatively developed, however,
most studies are conducted at local scale (up to national
scale), and even though they are of great scientific value,
their results and conclusions are difficult to compare and
cannot be extrapolated to the entire region (Kessler et al.
2011). Therefore, there is a need to increase research
studies that will characterize the pramo as a biogeo-
graphic region. The scarcity of regional studies in the
pramo are due to two main causes, (i) the difficulty to
establish joint international research collaborations,
which implies data and results sharing agreements and (ii)
the limited availability of open access data-sources con-
taining substantial amounts of biological data that can
sustain broad-scale studies. Pramo plant data, in the
form of floristic and vegetation records, are relatively
scarce and scattered, with floristic data contained in mo-
nographs, checklists and smaller floristic works (e.g. Lu-
teyn 1999) and vegetation data contained mostly in ve-
getation classification studies using different sampling
methods (e.g. Cleef 1981). Therefore, there is a need to
compile, homogenize and revise the existing plant data,
and generate new data in the areas with least information.
Biological databases are useful tools for research as
they can contain substantial amounts of information that
is uniform and frequently updated (Dengler et al. 2011).
There are several floristic databases focusing on the trop-
ics worldwide (e.g. TROPICOS, Missouri Botanical
eschweizerbart_xxx
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Gwendolyn Peyre et al.
Garden, http://www.tropicos.org) and some focusing es-
pecially on the Neotropics, providing pictures of herbar-
ium specimens (e.g. Neotropical Herbarium specimens,
Field Museum, http://fm1.fieldmuseum.org) or picture
of live plants (e.g. Neotropical Plants Image Database,
Royal Botanic Gardens Kew, http://www.kew.org/sci-
ence/tropamerica/imagedatabase). In addition, there are
national and smaller scale floristic databases available,
such as the National Colombian Herbarium Collections
(http://www.biovirtual.unal.edu.co). Regarding vegeta-
tion data, the Global Index of Vegetation-Plot Databases
(GIVD) lists 42 tropical vegetation plot databases, 10 of
which are South American but mostly of restricted access
(Dengler et al. 2011). Of particular relevance here is the
recent Colombian Pramo Vegetation Database, which
compiles 1,000 vegetation plots resulting from the con-
siderable field work conducted in the Colombian pramo
in the last decades, but so far it is not publicly accessible
(Rangel-Churio & Pinto-Zrate 2012). Vegetation plot
databases are becoming increasingly attractive to bota-
nists and ecologists (Font et al. 2009) as they provide
both floristic and vegetation data, and can sustain numer-
ous large-scale applications, such as vegetation classifica-
tion, floristic diversity studies and vegetation mapping.
To meet the need for intensifying research and conser-
vation in the northern high Andes, we here present Veg-
Pramo as a significant new botanical data-source for the
pramo. We constructed the database to contain phyto-
sociological data, from which vegetation plots and floris-
tic records can be obtained and we created an open access
webportal for VegPramo.
Methods
Structure of VegPramo
The basic data unit in VegPramo is the vegetation plot,
or relev, sampled with the phytosociological method
that aims at characterizing plant communities by their
representative diagnostic taxa (Braun-Blanquet 1951).
This method has been widely used in Europe but less in
the tropics due to the complex physiognomy of most
tropical plant communities, often including many lianas
and epiphytes; however, it is considered suitable for the
relatively simpler pramo vegetation structure (Cleef
1981). Phytosociological fieldwork involves defining a
plot in a representative area, with standard plot size de-
pending on the vegetation physiognomy and therefore
variable, usually ranging from (1) 10100 (500) m2
(respectively meadows and forests). The environmental
and geographic characteristics are then recorded and the
plant species in each vegetation layer listed. Each species
is assigned a cover index, usually following the scale: +
(less than 1%), 1 (up to 5%), 2 (up to 25%), 3 (up to
50%), 4 (up to 75%), or 5 (up to 100%), a categorized
07.07.15 12:10
 VegPramo, a flora and vegetation database for the Andean pramo
variable that requires careful manipulation in statistics
but is flexible in transformations (e.g. abundance, pres-
ence/absence).
We programmed VegPramo in SQL and Java follow-
ing the Iberian and Macaronesian Vegetation Informa-
tion System model (Font et al. 2009). The database con-
tains six main tables interconnected via one primary key:
the central Plot data table, the secondary Georeferences
and Attributes tables, the Plot source table, as well as two
checklists, the Taxon list and the Syntaxon list. The Plot
data table refers to the original plot species list and their
cover index. The Georeferences table contains the UTM
coordinates (mostly at 1 km 1 km scale) and precise lo-
cality (toponymy, municipality, province and country).
The Attributes table contains the information on vegeta-
tion physiognomy, or layers, in terms of height (in cm)
and cover (in %), but also other environmental character-
istics such as slope, orientation, soil, etc. The Plot source
table contains the published or unpublished references in
which the plot was first displayed. The two checklists
Taxon list and Syntaxon list, are lists of codified names for
plant taxa and plant communities (referred to as syntaxa
in phytosociology), with their updated validity status
(accepted, rejected, dubious, etc.). Both files check the
original names used in the VegPramo data and update
them automatically to their accepted form based on the
most recent source. The taxon checklist contains names
of vascular plants recorded in the pramo region for
about 188 families, 1,700 genera and about 15,000 species,
infra-specific taxa included, of which half are accepted
names. This exhaustive list is based on several sources
from which the information was obtained after definition
of the geographical area (country, province and elevation
above 2800 m a.s.l.): the Pramo Checklist (Luteyn 1999),
the Catalogue of the Ecuadorian Vascular Plants (Jr-
gensen & Len-Ynez 1999), the Catalogue of the Flow-
ering Plants and Gymnosperms of Peru (Brako & Zaruc-
chi 1993) and the Catalogue of the Flowering Plants of
the Venezuelan Pramos: Dicots and Monocots (Briceo
& Morillo 2011a, 2011b). Due to the selection process,
the taxon checklist also includes taxa from the upper
montane ecotone in certain areas and taxa from the south-
ern transition in northern Peru. We also consulted addi-
tional smaller sources and added their information for
specific taxa and we gave priority to the most recent
source in case of synonymy. We then revised and updated
the VegPramo names using TROPICOS and The Plant
List (http://www.plantlist.org). The syntaxon checklist
contains more than 500 names of plant communities de-
scribed in the literature, mostly using the phytosociolog-
ical syntaxonomic classification of plant communities
into classes, orders, alliances and associations, the latter
being the basic unit similar to species in taxonomy. The
syntaxonomic affiliation of the plots if only provisional
as there is no actual global syntaxonomic revision of the
pramos and synonymy will be provided in the future.
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phyto_45_1_2_195_202_peyre_0045.indd 197
197
Origin of the data
The 3000 vegetation plots contained in VegPramo come
from multiple sources. We retrieved 2,700 plots from the
existing literature on pramo vegetation in South Amer-
ica including published bibliography, thesis and scientific
reports covering the period 19552010. We took into ac-
count all plots obtained with the phytosociological
method and sampled in both zonal and azonal vegetation.
Most data from Colombia come from the extensive phy-
tosociological fieldwork carried out in the pramos by
local and international botanists within major research
projects such as ECOANDES (e.g. van der Hammen
2008). In Ecuador and Venezuela, even though there is a
long tradition of phytogeographical, geobotanical and
ecological studies in pramos, the interest in the phytoso-
ciological method is more recent and relatively few re-
searchers have used it in the pramo (e.g. Ramsay 1992).
We found no data for Peru, as the method has not yet
been used in the pramo, where biological studies remain
scarce. After mapping the distribution of the literature
plots in the region, we added our own 300 plots obtained
between 20112013, to cover the less sampled pramo ar-
eas in Peru, Ecuador and Venezuela.
Results and discussion
Data contents in VegPramo
The VegPramo data are contained in 170 UTM quadrats
of 100 km2, spread throughout the estimated 35,000 km2
potential pramo area (Fig. 1). The distribution of the
plots is not uniform because pramo fieldwork expedi-
tions were mostly driven by floristic interest and facility
of access. Therefore, some pramos are over-sampled,
like Chingaza in central Colombia, whereas others are
under-sampled, such as the Peruvian pramos. The Veg-
Pramo data are unequally spread over the four Andean
countries; however, they should be representative of their
respective pramo area and flora (Table 1). In Venezuela,
the pramo is geographically limited and has the lowest
floristic diversity, partly due to the desertic conditions of
the high pramos in the Cordillera de Mrida (Monaste-
rio & Reyes 1980), and its data currently accounts for
13% of the VegPramo plots. Colombia hosts the richest
pramo flora (Rangel-Churio 2000) and represents 52%
of the VegPramo data. Ecuador has the largest pramo
extension covering 30% of its territory and it is also nu-
merically and geographically well represented in our
data. Although geographically limited to the Amotape-
Huancabamba zone in northern Peru, the southernmost
pramos are very diverse ecosystems and remain rela-
tively preserved (Weigend 2002), however VegPramo is
quite incomplete for Peru, represented by a mere 3% of
all plots.
07.07.15 12:10
 198 Gwendolyn Peyre et al.

Table 2. Comparison of the relative importance (in %) of the

!!
((
!
(
(!
!( 10 main plant families in the pramo in terms of species be-
!!
((
tween this study and the regional checklist (Luteyn 1999).
(
!
(
!

!!
((
(
!
!
( VegPramo Luteyn (1999)
(
!(!
!(( !
(!
! (
(
!
!
(
(
!
!
(
Asteraceae 21 Asteraceae 25
!
(
(!
!(
Venezuela Poaceae 7 Poaceae 7
!!
((
(
!
Orchidaceae 5 Orchidaceae 4
(!
!(
(
! !
(
(!
!(
(
!
((
!
(
!
!!
( !
((
! Melastomataceae 4 Scrophulariaceae 4
(!
!(!
(
(!
! (
(
!
!
(
!
!
(
!
(!
((
(!
!
(
(
! Rosaceae 4 Melastomataceae 3
!!
((
(
!(
! (
!
(!
!(
!
(!
!
(!
!
(!
(
(
!(
!
(!
(
( (
!
(!
(!
!
(!
!
( Cyperaceae 3 Gentianaceae 3
(!
!
(
!(
!
((
(
!
(
!
Scrophulariaceae 3 Ericaceae 2
!!
((
Bromeliaceae 2 Bromeliaceae 2
!!
((
(
!
(
!
Gentianaceae 2 Rosaceae 2
Colombia
!!
( ( !
(
(!
!
((
(
!
!
Apiaceae 2 Fabaceae 2
(!
! ( (
!
(!
!
(( !
!
(!
((
!! (!( (!
(! (
(!
! (!
(
!
!
(
!
(
(!
!(
(
!
(!
! (!
!(
(!
!( (
(
! Ecuador
(
!
!
(
(!
!((
!
(!
!
(
!
(
! of the main families, in terms of order and values, generally
(
!
(
!(( !
! (
(!
(
!
!( (!
!(!
(
( !
! ( agrees with the regional floristic checklist compiled by Lu-
(
! Peru
(
!
teyn (1999) (Table 2). The expression of plant species
(
! N plots
(
!
(
! < 50
within the pramo area covered by VegPramo plots fol-
(
!
(
!
!
(
(
! 51-100 lows the traditional logarithmic series distribution (Fig. 2),
(
!
(
! ( 101-200
!
(!
!
(
!(
( > 200
!
with most genera represented by less than five species and
(
!
(
!
!
( Elevation (m) only 5% are represented by more than 10 species. Among
(!
! (
(
!
(!
!(
!
(
(
! 0 50 100 200 Km
< 3000 the richest genera are Diplostephium (Asteraceae), Miconia
> 3000
(Melastomataceae), Huperzia (Lycopodiaceae) and
Elaphoglossum (Dryopteridaceae). Regarding species, very
Fig. 1. Distribution of the VegPramo plots in the northern few are widely distributed, like Pernettya prostrata (Cav.)
Andes (potential distribution above 3000 m a.s.l. highlighted DC. (Ericaceae) present in 851 plots and Calamagrostis ef-
in grey).
fusa (Kunth) Steud. (Poaceae) present in 586 plots, and
most species are only represented in a few plots with 450
of them only in one plot.
The VegPramo data represent 123 vascular plants fam-
ilies, 504 genera and 2,220 species. While at family and ge-
nus level, VegPramo is rather complete, it remains incom- The VegPramo webportal
plete at species level representing a little less than 50% of
the total pramo flora estimate by Rangel-Churio (2000) VegPramo (GIVD: SA-00-002) is a free, open access
and 65% of Luteyns estimate (1999). In VegPramo, biological database, accessible from the webportal http://
Asteraceae, Poaceae and Orchidaceae are the richest fami- www.vegparamo.com in English and Spanish. The data-
lies in species, and we observe that the relative importance base is part of the BIODIVER databases complex devel-
oped by the Biosystematics and Vegetal Biodiversity Re-
search Group at the University of Barcelona that includes
among others the Information System for Iberian and
Table 1. Pramo area (Josse et al. 2009), flora (Sklen et al.
Macaronesian Vegetation (SIVIM), the Information Sys-
2005) and plot numbers in VegPramo for Venezuela, Co-
lombia, Ecuador and Peru. tem for Andorran biodiversity (SIBA) or the Atlas of the
Pyrenean Flora (POCTEFA). From the webportal, flo-
Venezuela Colombia Ecuador Peru ristic data (taxonomic record) and vegetation data (vege-
Area (km2) 2,405 14,086 18,435 953 tation plots) can be consulted via online searches and
downloaded. The webportal is designed for online
Plant 848 1,927 1,524 1,035
searches by taxon name (genus, species), plant commu-
species
nity (syntaxon name or combination of species) or geo-
Plots in 391 1,562 971 77 graphic area (UTM quadrat or locality) and search results
VegPramo will give the selected record with its geographical distri-

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phyto_45_1_2_195_202_peyre_0045.indd 198 07.07.15 12:10

 VegPramo, a flora and vegetation database for the Andean pramo
a) 



1JHQHUD






        
1VSHFLHV
Fig. 2. Floristic contents in VegPramo a) number of species per plant genus and b) number of VegPramo occurrences per
species.
bution, taxonomic/syntaxonomic contents and descrip-
tion if available. Names on the portal are always checked
by either the taxon or the syntaxon checklist so to allow
the user to work with updated valid names. Finally, the
search results can be downloaded in TXT or b-VegAna
XML format, making their export to data edition/analy-
sis programs easy.
Future perspectives
The pramo ecosystem is unique and ecologically threat-
ened, which make further research and conservation
measures on this singular neotropical environment a ne-
cessity. Towards this goal, VegPramo provides a sub-
stantial botanical data-source for the pramo, with data
well distributed and geographically representative of the
pramo region. Nonetheless, the floristic contents of the
database and especially the large amount of low-occur-
rence species indicate that VegPramo still needs to grow
in floristic representativeness.
The VegPramo data can be used for many research
studies involving flora and vegetation. The most com-
mon use of large phytosociological datasets is the classifi-
cation of vegetation into plant communities, or types (e.g.
Peyre & Font 2011). In addition, diversity studies can be
conducted on a taxon (e.g. Ericaceae), a specific area (e.g.
southern Ecuador), a plant community (e.g. Polylepis
forests) or comparing units (e.g. comparison of the
Calamagrostis grasslands). Mapping the current distribu-
tion of a taxon or a plant community can be carried out
easily, using a sub-dataset of VegPramo. Moreover,
modeling the actual or future distributions of pramo
species can be done using many predicting techniques
thanks to the versatility of the phytosociological data.
Conservation plans and strategies rely on data analysis
from research carried out in research or management-
oriented institutions. On one hand, research institutions
massively produce useful publications, but the data is of-
eschweizerbart_xxx
phyto_45_1_2_195_202_peyre_0045.indd 199
199
b) 

1VSHFLHV




   !
1RFFXUUHQFHV
ten kept away, especially in the tropics, and the results are
not always applicable or easy to synthesize regularly for
management planners (Guisan et al. 2013). On the other
hand, research carried out in management-oriented insti-
tutions is more easily applicable to case studies for con-
servation but sometimes miss a more global context and
it is often limited by the lack of substantial amounts of
quality biological data (Cayuela et al. 2009). In this way,
VegPramo provides a novel approach in terms of quan-
tity and quality of data, despite the multiple data sources,
thanks to the overall taxonomic revision and the georef-
erencing of the data. Moreover, the vegetation plot for-
mat is especially useful as it gives abundance as well as
presence/absence data, which are both important for
studying the geographic distribution of endangered spe-
cies. Conservation strategies are increasing and improv-
ing their organization in the pramo region, but given the
enormous task, efforts must be joined and collaborations
developed, nationally as well as internationally (Hofstede
et al. 2003). We hope the opportunity of using and im-
proving VegPramos data will help increase the activity
of conservationists and ease the communication with re-
searchers.
We encourage further sampling effort on pramo ve-
getation to improve VegPramo with new data, especially
from narrowly distributed and very biodiverse under-
sampled pramo areas such as the Peruvian pramos. We
are developing new statistical tools for the webportal (di-
versity, fidelity index, etc.) that we believe will be useful
for exploratory analyses online and also new features of
ecological interest, such as the IUCN status of species if
available, which is valuable information for conservation
prioritization not only of a specific taxon but also of hab-
itats. Finally, we invite data revision, comments, updates
and references through the interactive feedback option
online that will allow any user to participate in the ad-
vances of VegPramo.
07.07.15 12:10
 200
Author contribution
G.P. led the writing while all authors critically revised the
manuscript.
Acknowledgments
We would like to thank the Agency for Administration of
University and Research Grants (AGAUR) from the Gen-
eralitat de Catalonia (Spain) for the PhD grant to the first
author. Many thanks to our data and fieldwork collabora-
tors: Roberto Carrillo, Javier Castillo, Antoine Cleef, Jos
Campos de la Cruz, Raiza Garcia, Rene Lpez, Gilberto
Morillo, David Rivera, Katya Romoleroux, Roberto Rueda,
Dionys Snchez, Isidoro Snchez-Vega, Francisco Santiago,
Walter Vargas, Renato Valencia and the staff of the Aarhus
University Herbarium, CPUN Herbarium at the National
University of Cajamarca, the Herbarium at the Royal Bo-
tanic Gardens Kew, the Herbaria MERC and MERF at the
University of the Andes, the Missouri Botanical Garden
Herbarium, the New York Botanical Garden Herbarium,
the PORT Herbarium at the National Experimental Univer-
sity of the Llanos Ezequiel Zamora, the QCA Herbarium at
the Pontifical Catholic University of Ecuador and the San
Marcos University Herbarium. We also thank the expert
taxonomists who helped with the determination of our field
samples: Benito Briceo, Mats Gustafsson, Nicholas Hind,
Simon Lgaard, Ulf Molau, Robbin Moran, Benjamin ll-
gaard, Paola Pedraza and Carmen Ulloa. Finally, we would
like to thank Rafael Quadrada for helping with the pro-
gramming of VegPramo.
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Author addresses

Peyre, Gwendolyn (Corresponding author, g.peyre@ub.edu)1,2, Balslev, Henrik (henrik.balslev@bios.au.dk)2, Mart, David
(utopicmail@gmail.com)1, Sklen, Petr (petr@natur.cuni.cz)3, Ramsay, Paul (paul.ramsay@plymouth.ac.uk)4, Lozano, Pablo
(pablo_lozanoc@yahoo.com)5, Cuello, Nidia (nidia.cuello@gmail.com)6, Bussmann, Rainer (rainer.bussmann@mobot.org)7
Cabrera, Omar (hocabrera@utpl.edu.ec)8 & Font, Xavier (xfont@ub.edu)1
1
Department of Plant Biology, Faculty of Biology, University of Barcelona, Av. Diagonal 645, 08028 Barcelona, Spain
2Ecoinformatics and Biodiversity, Department of Bioscience, Aarhus University, Ny Munkegade 114, 8000 Aarhus C, Denmark
3
Department of Botany, Charles University, 128 01, Prague, Czech Republic
4Department of Biological Sciences, University of Plymouth, Drake Circus, Plymouth, Devon, PL4 8AA, UK
5Amazonian State University, Cristobal Colon y Alvaro Valladares, Puyo, Pastaza, Ecuador
6UNELLEZ-Guanare, Natural Renewable Resources Program, University Herbarium (PORT), Mesa de Caracas, Portuguesa

3323, Venezuela
7William L. Brown Center, Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, USA
8Institute of Ecology, 23 Technical Particular University of Loja, San Cayetano Alto, Marcelino Champagnat, Loja, Ecuador

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