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Flora 200 (2005) 416433


Superparamo plant species diversity and phytogeography in Ecuador

Petr Sklenara,, Henrik Balslevb
Department of Botany, Charles University, Benatska 2, 128 01 Prague, Czech Republic
Department of Systematic Botany, Aarhus University, Universitetsparken Bygn. 540, DK8000 Aarhus C., Denmark

Received 14 April 2004; accepted 21 December 2004


In 18 superparamo sites in Ecuador we found 388 species of vascular plants belonging to 146 genera and 52 families,
making the Ecuadorian superparamo ora richer in species than that of Venezuela and that of Colombia which
appears to have fewer species although the number remains uncertain. The most species rich families were Asteraceae
(83 species) and Poaceae (49) which also dominate the grassparamo that surrounds the superparamo. Otherwise the
superparamo is dominated by families that are mostly herbaceous such as Cyperaceae, Brassicaceae, Caryophyllaceae
and Valerianaceae, whereas shrubby families that dominate the subparamo and grassparamo, such as
Melastomataceae, Ericaceae, and Solanaceae, have only few species or are absent in the superparamo. The generic
spectrum was dominated by a suite of species-rich genera (Lachemilla, Gentianella, Valeriana, Calamagrostis, Draba)
with many paramo-endemic species reecting a high level of autochthonous speciation in the (super)paramo. Species
richness varied from 71 to 149 in the individual superparamos surveyed, but species richness was only weakly
correlated to their area and log-transformed area, and negatively correlated to the vertical range of each superparamo.
b-diversity was signicantly correlated to the vertical range (i.e., the number of surveyed altitudinal levels) but it was
not correlated to the area or log-transformed area of each superparamo. Most of the species were narrowly distributed
and 112 (29%) of them were found in a single superparamo, while eight (2%) occurred in all 18 superparamo sites.
Floristic similarity was not correlated to the geographical distance between the sites. Redundance Analysis suggested
that geological origin of the substrate (metamorphic versus volcanic bedrock) was important for the oristic
composition. Occurrence of mountains built from metamorphic rocks, is however, correlated to areas with high
rainfall and the amount of rainfall may be a stronger determinant for species distribution than the presence of volcanic
versus metamorphic bedrocks. TWINSPAN cluster analysis divided the 18 sites into three groups, which corresponded
to dry, humid and very humid superparamos. The groups were also separated along the rst ordination axis in
Correspondence Analysis, while the second axis may correlate to the volcanic history of the area.
r 2005 Elsevier GmbH. All rights reserved.

Keywords: Andes; Habitat islands; Paramo; Phytogeography; Species distribution; Tropical alpine


Corresponding author. The superparamo is the upper vegetation belt of the

E-mail addresses: petr@natur.cuni.cz (P. Sklenar), paramo an essentially aseasonal high elevation grass-
henrik.balslev@biology.au.dk (H. Balslev). and shrub-vegetation of the humid tropical Andes with

0367-2530/$ - see front matter r 2005 Elsevier GmbH. All rights reserved.
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 417

outliers in Costa Rica and Panama. Where more isolated on 25 or more mountains in both the western
seasonal climate reigns further north and south, the and eastern cordilleras along a stretch of ca. 450 km
paramo gives way to other vegetation types above the from the Colombian border in the north to the province
timberline; in Guatemala and Mexico to herbaceous and of Azuay in the south. On most of these mountains the
tussock-grass communities called zacatonales and in superparamo is developed on volcanic bedrocks but on
central Peru and Bolivia to dry, barren grass- and bush- a few mountains in the eastern cordillera it has
steppe called puna (Almeida-Lenero et al., 2004; developed on metamorphic bedrocks. Depending on
Balslev and Luteyn, 1992; Islebe and Cleef, 1995; local climatic conditions and orientation relative to the
Luteyn, 1999; Simpson, 1983; Troll, 1959, 1968). Like trade winds, the superparamo may be dry or humid.
other tropical mountains, the Andes are oristically rich Some areas of superparamo are found on mountains
and often have high endemism, particularly in their that are high enough to have permanent glaciers
upper zones such as the tropical alpine belts above the whereas others do not reach such heights. This setting
timber line (Balslev and Luteyn, 1992; Hedberg, 1957; is an attractive one for comparative studies of species
Smith, 1975). Among the tropical alpine ecosystems, the diversity, taxonomic composition, and distribution of
paramo is the most diverse, mainly because of its large vascular plant species, and phytogeographic relations
geographic extent (Hedberg, 1992; Smith and Cleef, among the superparamo islands. In this paper, we
1988; Vuilleumier and Monasterio, 1986). About 3500 address the following questions: (1) How species rich is
species of vascular plants have been reported from the the Ecuadorian superparamo compared to Colombian
paramo and possibly as many as 60% of them are and Venezuelan superparamos? (2) Is the superparamo
endemic (Luteyn, 1992, 1999). The high concentration ora dominated by the same taxa as the grass paramo
of endemic species is usually attributed to changes of that surrounds it, and are some genera particularly
horizontal and vertical extent of the paramo during diverse, indicating high levels of autochthonous specia-
Pleistocene climatic oscillations and associated pulses of tion? (3) How does species richness and b-diversity vary
geographic isolation (Cuatrecasas, 1986; Simpson and among Ecuadorian superparamos and is the variation
Todzia, 1990; Van der Hammen and Cleef, 1986; Van related to their areas or their vertical ranges? (4) How
der Hammen et al., 1973). are individual species distributed among the super-
Above 41004300 m elevation, the so-called super- paramos and how is endemism structured? (5) How
paramo, which is the subject of this study, replaces the oristically similar are the superparamos and are the
paramo grasslands (or the bamboo paramo in humid observed differences related to ecological or other
areas) and extends to the permanent snow-line at factors?
74800 m. In the superparamo, vegetation cover is
broken up and patches of open soil appear, the tussock
grasses that are characteristic of the grass paramo are
less abundant, and the vegetation is dominated by other Study sites
life-forms such as small herbs, cushion plants, and
sclerophyllous dwarf shrubs. Superparamo vegetation We selected 16 representative areas of superparamo in
has been reported from Venezuela, Colombia, and the eastern and western cordilleras of the Andes in
Ecuador (Berg, 1998; Cleef, 1981; Cuatrecasas, 1968; Ecuador (Fig. 1) stretching over 430 km from Volcan
Luteyn, 1992; Monasterio, 1979), whereas the summits Chiles (01480 N, 771570 W) on the border with Colombia
in Panama and Costa Rica do not reach elevations that to Cajas National Park (21520 S, 791180 W) in southern
are high enough to support superparamo. In northern Ecuador. The superparamos on Chimborazo and
Peru, several summits of the jalca, which is the local Antisana were studied on two opposite sides of the
name for paramo, reach elevations at which super- mountains. These data were treated as independent sets,
paramo may have developed, but so far there are no giving a total of 18 study sites, of which 14 were located
descriptions in the literature that document the existence on active or extinct volcanoes, and four on mountains of
of superparamo vegetation in Peru. On a few occasions non-volcanic origin with metamorphic bedrocks. Nine
the name superparamo has been used for high-elevation of the studied mountains are topped by glaciers today,
vegetation as far south as Bolivia (Cuatrecasas, 1968), but all of them were glaciated at some point during
but since it replaces puna there, super-paramo is an Pleistocene (Hastenrath, 1981). The basic characteristics
inappropriate name. of each site are summarized in Table 1 and further
Since the superparamo is conned to the highest details can be found in Sklenar (2000).
mountains, it occurs scattered as islands separated by In Ecuador, the superparamo forms two distinct
grass paramo, mountain forest, and deep Andean altitudinal belts. The lower superparamo from 4100 to
valleys (Cuatrecasas, 1958; Simpson, 1974, 1975; Van (4400)4500 m is usually rich in species. It is character-
der Hammen and Cleef, 1986). In Ecuador, super- ized by sclerophyllous dwarf shrubs (Loricaria
paramo is particularly well represented and occurs spp., Chuquiraga jussieui, Valeriana microphylla,
418 P. Sklenar, H. Balslev / Flora 200 (2005) 416433

78 W upper limit was dened as where the plant cover

dropped below 5%. At 100 m altitudinal (stratied)
Ecuador levels we randomly set three plots along a 100 m long
transect running parallel with the contours. Most plots
America were 25 m2, but in the upper belt (generally above
4400 m) they were 100 m2 because there we found
4 0
5 increased spatial variability in species composition. In
6 each plot we scored all species using an 8-grade semi-
8 7 quantitative scale combining abundance and cover
9 (Mueller-Dombois and Ellenberg, 1974). Data were
species beta converted to their midpoint values (r 0.05%,
richness diversity
1 Chiles 103 3.2 + 0.5%, 1 2.5%, 2a 10%, 2b 20%,
2 Cotacachi 102 1.8 3 37.5%, 4 62.5%, 5 87.5%) for further analysis.
3 Imbabura 114 1.9
4 Cayambe 138 3 In addition to the quantitative sampling we searched
13 14 12 5 Saraurcu (M) 95 1.8
6 Pichincha 131 2.2
the study sites for additional species outside the sample
7 Antisana - east 129 2.4 plots but this time including also azonal habitats, such
8 Antisana - west 143 2.3
9 Iliniza 143 3.9 as rocky outcrops, cushion mires, and lake or stream
10 Cotopaxi 91 3.6
11 Hermoso (M) 122 1.4
shores, and they were added to the species lists for each
17 12 Tungurahua 71 1.2 site. Species planted occasionally around mountain
13 Chimborazo - west 107 1.6
14 Chimborazo - east 131 2.1 refuges or occurring on ruderal sites around huts were
El Altar
Quilimas (M)
excluded from the phytogeographic analyses. Intro-
17 Yanaurcu (M) 106 0.7 duced species that were naturalized, i.e., they reproduce
18 Cajas 149 1.4
0 50 150 km
and survive independently in natural plant communities,
were included in all analyses. Identication was backed
up with voucher specimens and usually veried by
Fig. 1. Distribution of 18 vegetation study sites in the
specialists for each taxonomic group. Information about
superparamo in Ecuador along a 430 km long stretch of the
Andes in both the western and eastern Cordilleras. Squares
species endemism in Ecuador was extracted from
correspond to Group 1 and circles to Group 2 in Fig. 4. (M) Jrgensen and Leon-Y. (1999) and Valencia et al.
after the name of site indicates metamorphic bedrock. (2000) and was updated from our own knowledge.
b-diversity was calculated from the vegetation sam-
ples for each site as: bT s=a  1, where s is the total
Diplostephium rupestre, Pentacalia spp.), and cushion number of species present in all samples at a site and a is
plants (Plantago rigida, Xenophyllum humile, Azorella the average number of species in the samples from that
spp.), but tussock grasses (Calamagrostis intermedia, site (Whittaker, 1960; Wilson and Shmida, 1984).
Festuca asplundii) are usually also important. The upper Correlation of species richness and b-diversity to
superparamo above (4400)4500 m is characterized by environmental and spatial variables were calculated.
the presence of Senecio nivalis, S. canescens, Arenaria The explanatory variables were superparamo area
dicranoides, and Cerastium floccosum. The vegetation is (measured as plane area estimates from the IGM
poor in species and conned to favorable habitats. 1:50,000 maps; if present, the extension of glaciers was
Tussock grasses and sclerophyllous shrubs are mostly excluded), vertical range of the superparamo (i.e., for
absent, and the most common growth-forms are short- species richness vertical distance between the base and
stem grasses, prostrate subshrubs and herbs, acaulescent the top of the superparamo or the position of the
rosettes, and cushion plants (Sklenar, 2000). permanent snow-line, but for b-diversity number of
surveyed altitudinal levels), presence/absence of glacier,
volcanic/metamorphic bedrock, volcanic activity yes/no,
Methods position in western or eastern cordilleras.
Floristic relationship between pairs of sites was
Field work was carried out from May through measured as the difference between observed and
December 1995, June through September 1997, and expected numbers of shared species (Connor and
June through July 1999. The vegetation was sampled in Simberloff, 1978). Expected numbers were estimated
a stratied random design (Mueller-Dombois and by a random pair-wise sampling from the entire species
Ellenberg, 1974). This sampling was limited to the zonal list in which each species had an adjusted probability of
vegetation, avoiding azonal patches of cushions mires, being sampled, for species found in the vegetation plots
stream-sides, etc. The lower limit of the superparamo the expected numbers were the frequency of occurrence
was here dened as the transition to the grass paramo, and for species found only outside the vegetation
i.e., where a continuous grass cover begins, while the plots the expected numbers were the number of site
Table 1. Baseline data for the 18 study sites.

Name of site Province Maximum Coordinates Estimated superparamo Vertical Recent Geological Recent volcanic
elevation (m) latitude longitude area (km2) range (m) glacier origin activity

Chiles Carchi 4768 01480 N 771570 W 6.4 564 No Volcanic No

Cotacachi Imbabura 4939 01220 N 781200 W 6.7 737 Yes Volcanic No

P. Sklenar, H. Balslev / Flora 200 (2005) 416433

Imbabura Imbabura 4609 01160 N 781100 W 1.6 330 No Volcanic No
Cayambe Pichincha 5790 01020 N 771590 W 43.3 600 Yes Volcanic No
01060 S 771550 W

Saraurcu Pichincha 4676 12.6 576 Yes Metamorphic
Pichincha Pichincha 4698 01100 S 781330 W 6.5 398 No Volcanic Yes
Antisana-west Napo 5704 01300 S 781100 W 29 500 Yes Volcanic Yes
Antisana-east 01270 S 781070 W 29 500
Iliniza Cotopaxi 5263 01400 S 781420 W 26.9 600 Yes Volcanic No
Cotopaxi Cotopaxi 5897 01400 S 781260 W 65.5 600 Yes Volcanic Yes
Hermoso Tungurahua 4639 11100 S 781120 W 7.2 539 No Metamorphic
Tungurahua Tungurahua 5016 11270 S 781260 W 11.1 400 Yes Volcanic Yes
Chimborazo-west Chimborazo 6310 11280 S 781520 W 62.2 700 Yes Volcanic No
Chimborazo-east 11280 S 781460 W 62.2 500
El Altar Chimborazo 5319 11410 S 781240 W 44.3 500 Yes Volcanic No
Quilimas Chimborazo 4730 11460 S 781240 W 47.2 630 No Metamorphic
Yanaurcu Chimborazo 4580 21150 S 781300 W 15.4 380 No Metamorphic
Cajas Azuay 4451 21520 S 791180 W 17.7 151 No Volcanic No

The vertical range of the superparamo is the vertical distance in meters between the lowest sampling plot and the top of the mountain or the transition to the permanent snow-line, coordinates
indicate the approximate location of the transects, estimated superparamo area is a plane estimate obtained from the IGM maps (excluding glaciers where present).

420 P. Sklenar, H. Balslev / Flora 200 (2005) 416433

occurrences (null hypothesis II of Connor and Simberloff, Species diversity was unevenly distributed among
1978). The size of the random sample was equal to the taxonomic groups (Fig. 2). Asteraceae and Poaceae
actual number of species for any given site. The expected entirely dominated the ora and together they com-
numbers of shared species, independent of sample size prised one-third of the total species diversity. Other
(Connor and Simberloff, 1978; Simberloff et al., 1981), families were conspicuously less diverse and there were
were estimated repeatedly 999 times via computer 16 families with a single species. There were a suite of
simulation (Sklenar, 2000). The chi-square (w2 ) distance species rich genera, but the majority of the 146 genera
between observed and expected numbers was used as a present in the Ecuadorian superparamo, i.e., 84 genera,
standardized measure of oristic relationships between were represented by a single species.
the mountain pairs. The w2 values were tested against
geographical distances between the sites by means of
Variation in species richness and b-diversity among
Mantel test, part of the R-Package (Legendre and
Vaudor, 1991); the null hypothesis of the test was that
there would be no linear relationship between the
Species richness differed considerably among the
geographical distance and the oristic similarity.
superparamo sites (Fig. 1). Cajas was richest in species
Two-way indicator species analysis (TWINSPAN;
(149 species) even if it is one of the lowest mountains
Hill, 1979) was used to group the 18 sites according to
(Table 1). The lowest number of species was found in
oristic and vegetation similarities with presenceab-
Tungurahua (71) and Cotopaxi (91), which are the
sence data for all species and semi-quantitative abun-
mountains that have the most recent volcanic activity.
dance estimates for species present in the plots based on
Species richness correlated weakly with the estimated
average values of each species in the plots at each site.
area of superparamo at each site (Spearman rank
For the abundance data we used CA (Correspondence
correlation coefcient r 0:173, p 0:49), or the log-
Analysis, part of CANOCO package; Ter Braak and
transformed area (r 0:264, p 0:29), and correlation
Smilauer, 1998) to demonstrate the relative position of
with the vertical range of the superparamo was negative
study sites in the multivariate space; data were square-
(r 0:191, p 0:45; n 18 in all cases). When the
root transformed and rare species were down-weighted
correlation analyses were performed independently for
prior the analysis. For the presenceabsence data we
each of the two groups of mountains resulting from the
used RDA (Redundancy Analysis) to test the correla-
cluster analysis (see below), the results became signi-
tion between species distribution among the sites and six
cant for the area and log-area of the 13 mountains of
environmental and spatial variables.
Group 2 (r 0:664, p 0:01) but not for the ve
mountains of Group 1 (r 0:667, p 0:22). The
correlation did not change for the vertical range of the
Results superparamo (Group 1: r 0:395, p 0:51; Group 2:
r 0:088, p 0:77).
A considerable spatial pattern of b-diversity in the
Species richness and taxonomic composition
superparamo vegetation (Fig. 1) was signicantly
correlated with the number of altitudinal levels surveyed
There were 383 species, 146 genera and 52 families of
at each site (r 0:59, p 0:01), whereas its correlation
vascular plants in the 18 Ecuadorian superparamo sites
with other variables (i.e., area, vertical range of super-
surveyed. There were two subspecies in Eudema nubigena
paramo) was weak and not signicant (p40:05). This
and Valeriana alypifolia, two varieties in Potentilla
signicant correlation to the altitudinal levels remained
dombeyi, and two hybrids in Gentianella, which we, for
within the cluster Group 2 (r 0:605, p 0:03) but not
simplicity treated in the same way as species in the
within the Group 1 (r 0:051, p 0:93), whereas the
analyses, so the number of taxa (subsequently referred to
other correlations remained not signicant.
as species) was 388. Of these, 374 were recorded within
the continuous superparamo, i.e., above the transition with
the grass- or bamboo paramo at (4000) 41004200 m. The Species distributions
remaining 14 species were found within 100 m below this
limit, in isolated patches of superparamo vegetation, such The majority of species were narrowly distributed; 112
as rock crevices, surroundings of boulders, lake shores, or species (29% of the total) were found only at one single
eroded land, and we therefore expect that they will site, and 274 or over two-thirds of the species occurred
eventually be found in the continuous superparamo. Of at fewer than seven sites (Fig. 3). Only eight species (e.g.,
the 388 species, ve (Aira caryophyllea, Anthoxanthum Calamagrostis intermedia, Gentiana sedifolia, Luzula
odoratum, Poa annua, Rumex acetosella, and Sagina racemosa, Poa cucullata) were found at all 18 sites.
procumbens) are introduced to Ecuador. A full listing of Species restricted to either dry or humid sites (Groups 1
the species is provided by Sklenar (2000). and 2 in Fig. 4, resulting from the rst step of the cluster
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 421


80 Asteraceae (26 endemic)



Number of Species

Poaceae (12 endemic)


Brassicaceae (12 endemic)

Gentianaceae (11 endemic), Rosaceae

20 Scrophulariaceae, Caryophyllaceae, Cyperaceae



Lachemilla (2 endemic), Gentianella (10 endemic)

Valeriana (5 endemic)
Calamagrostis (3 endemic)

Draba (8 endemic)
Number of Species


Festuca (5 endemic), Geranium (3 endemic), S enecio


genera 63-146

1 3 5 7 9 11 13 15 17 19 21 23 25 27 29 31 33 35 37 39 41 43 45 47 49 51 53 55 57 59 61 63

Fig. 2. Ranked species numbers of families and genera encountered in 18 study sites in the superparamo in Ecuador.

analyses, see below) were also very narrowly distributed all of them (Fig. 5A). The 13 humid sites had a total of
within these groups, and only few of them occurred at 323 species, of which 137 were restricted to them, and
many sites (Fig. 3). The ve dry sites had a total of 241 there were no species occurring in all of them. The most
species, of which 64 were restricted to them, and only widespread were Geranium sibbaldioides (Fig. 5H) and
Festuca vaginalis and Plantago nubigena were present in Luzula gigantea (Fig. 5L) which occurred in 11, and F.
422 P. Sklenar, H. Balslev / Flora 200 (2005) 416433



All superpramo sites

Number of Species

60 Humid superpramo sites

Dry superpramo sites



1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

Number of sites (N)

Fig. 3. Pattern of geographic distribution of all species among the 18 study sites in the superparamo in Ecuador (open bars), and
pattern of distribution of species restricted to the 13 humid superparamos (hatched bars) and to the ve dry superparamos (black

asplundii (Fig. 5J) and Lachemilla holosericea (Fig. 5I) sence data. Tungurahua was consistently separated from
which occurred in 10 humid sites, respectively. the remaining nine mountains because of absence of
Azorella aretioides. The rest formed two sub-groups; the
southern superparamo sites (Quilimas, Yanaurcu, Altar,
Floristic similarity among superparamos Hermoso) were separated from the north-central ones
(Antisana-east, Saraurcu, Chiles, Cotacachi), due to the
Grouping of the superparamo sites was similar in occurrence of Calamagrostis guamanensis (Fig. 5O) in the
TWINSPAN clustering using either presenceabsence latter. This species was recorded also on Hermoso but
data of species from the superparamo sites (from both only outside vegetation plots and therefore this site
zonal and azonal habitats) or abundance data of the appeared in either the southern or the north-central sub-
zonal vegetation, and the differences were essentially group in the two analyses.
only in the nal divisions. This shows that adding the The rst CA ordination axis using the abundance data
species from the azonal vegetation patches (presen- captured the gradient expressed by the rst division step
ceabsence) does not dramatically change the picture of TWINSPAN clustering; sites from the Group 1 occur
obtained on the basis of the survey that included only on the right side while sites from the Group 2 on the left
the zonal vegetation (abundance). The analysis based on side of the ordination diagram (Fig. 4A). Cajas appears
abundance generally had higher eigenvalues and only a very distinct site separated from the remainder by the
this result is presented (Fig. 4B). second ordination axis.
The rst division step separated Chimborazo-west, A total of 374 species formed the common species
Antisana-west, Iliniza, Cotopaxi, and Pichincha (Group pool, which was used for estimations of the expected
1), due to the occurrence of Plantago nubigena and numbers of shared species (Appendix 1). Six sites
Festuca vaginalis (Fig. 5A). In the second group resulting (Chiles, Cotacachi, Imbabura, Cayambe, Saraurcu,
from the rst division (Group 2), Imbabura, Cajas, Antisana-east) had positive w2 values between each pair
Cayambe, and Chimborazo-east were separated next due of them (the only exception being ImbaburaAntisana-
to occurrence of Pentacalia peruviana; Cotacachi was east), mostly positive or low negative values with
added to this group in the analysis using presenceab- Chimborazo-east, Hermoso, Altar, Quilimas, and Ya-
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 423

(A) Chimborazo - west (B)
! 0.473
Antisana - west

Group 1
Cotopaxi 0.395

! 0.325
Chimborazo - east
Chimborazo-west 0.276

Group 2
Hermoso (M)

Altar Iliniza Cotopaxi Cotacachi

Cotacachi 0.495
Yanaurcu (M) Antisana-west Chiles 0.353
Saraurcu (M) Chimborazo-east
Imbabura Antisana - east
Chiles Quilimas (M) Pichincha 0.398
Cayambe Saraurcu (M)
Antisana-east Hermoso (M)
Tungurahua El Altar
Yanaurcu (M) ! 0.328
-1.0 Quilimas (M)
-1.0 0 1.5

Fig. 4. Correspondence Analysis of oristic abundance data from 18 superparamo sites in Ecuador (l1 0:316, l2 0:138, total
inertia 1.537); squares (&) indicate sites of Group 1 and circles (J) indicate sites of Group 2 resulting from the cluster analysis (A)
and cluster (TWINSPAN) analysis of abundance data from the 18 sites with the rst division separating two groups based on the
presence (Group 1) or absence (Group 2) of Festuca vaginalis and Plantago nubigena; eigenvalues for each division step provided, !
indicate divisions with higher residual than the tolerance given by the algorithm (B); (M) indicates site with metamorphic bedrock.

naurcu, and high negative values with Chimborazo- the bedrock variable accounted for only a small portion
west, Cotopaxi, Antisana-west, and Iliniza (Appendix (10.7%) of total data variability.
2). The latter four sites had positive values between each
pair of them. Three sites (Pichincha, Tungurahua,
Cajas), did not appear to demonstrate a clear pattern.
Pichincha showed mostly small values (negative or Discussion
positive), the highest in combination with Cotacachi,
Cayambe, and Iliniza, and the lowest with Hermoso and Species richness
Cajas. Tungurahua had small negative values with most
of the mountains, only with Chimborazo-west the value Most of the sites were visited only once, but Iliniza
is remarkably low (w2 24:56). Cajas was unusual since and Chimborazo were explored repeatedly. The rst 5-
all but one of its values were negative, the lowest one days survey of Iliniza yielded 127 species, but 16
with Chimborazo-west. additional species (13% of the total) were found during
The overall spatial pattern of oristic similarities the second 7-days intensive research of the mountain
between the mountains was not correlated with the three months later. In 1999, the transect area on
geographic distance between the sites; a Mantel test Chimborazo-west was revisited during a 1-day trip.
between the two matrices (Appendices 1 and 2) gives Four additional species were found (3.5% of the total),
r 0:034, p 0:38, using 999 permutations. The but three of them occurred above the limit surveyed in
RDA showed that of the six variables tested (estimated 1995. Some gaps in our species distributions may
area, vertical range of superparamo, presence/absence of therefore result from sampling error. However, if our
a recent glacier, position in western/eastern cordillera, experience at Iliniza is representative, we have recorded
presence/absence of recent volcanic activity, and volca- 8090% of all species at each superparamo site.
nic/metamorphic bedrocks), only the latter (bedrock The remarkable diversity of neotropical paramos,
volcanic/metamorphic) was signicantly correlated to previously highlighted by Smith and Cleef (1988) and
the observed species composition of the sites (forward Luteyn (1999), is underlined by our record of 388 species
manual selection, po0:05, 199 permutations). However, from the superparamo in Ecuador. This number
424 P. Sklenar, H. Balslev / Flora 200 (2005) 416433

Fig. 5. Examples of distributions of species among 18 study sites in the superparamo in Ecuador ( species not found at study site;
J species found at study site); AF species characteristic of dry sites, GL species characteristic of humid sites, MR species
characteristic of very humid sites, ST species negatively characterizing very humid sites, UV narrowly distributed species with
trans-Andean distribution, WX narrowly distributed species with and cis-Andean distribution.
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 425

includes some species that typically occur in the grass (Luteyn, 1999). Their importance is further underlined by
paramo at lower elevations (e.g., Hieracium frigidum, the fact that nearly half of the 40 most important
Stachys elliptica) or bamboo paramo (e.g., Gentianella (combining frequency and abundance) species of the
splendens), and only marginally enter the lower parts of Ecuadorian superparamo belong to these two families
the superparamo belt. On the other hand, we are aware (Sklenar, 2000). Orchidaceae, especially epiphytic ones,
of several species, such as Azorella ecuadorensis, Eudema and shrubby families, such as Melastomataceae,
rupestris, and Geranium chimborazense, previously col- Ericaceae, and Solanaceae rank among the most im-
lected in the superparamo in Ecuador, that were not portant paramo groups (Luteyn, 1999), but they are poor
encountered in this survey. Those species may either be in species or even absent in the superparamo ora. In
very rare, or restricted to sites that were not covered by contrast, several herbaceous families (e.g., Cyperaceae,
our study (e.g., Xenophyllum roseum from the central Brassicaceae, Caryophyllaceae, Valerianaceae) are
part of the Cajas National Park), or possibly they were among the most species-rich groups in the superparamo
not recognized in the eld (e.g., sterile specimens of ora whereas they are (relatively) less important when the
Carex are very hard to identify to species level). whole paramo belt is considered.
With 388 species of vascular plants the Ecuadorian The most species-rich genera, usually with numerous
superparamo ora seems to be richer in species than country-endemic species, are Lachemilla, Gentianella,
other superparamo oras. In the superparamo in Valeriana, and Draba (Fig. 2), and they are diverse also
Venezuela Ricardi et al. (1997) reported 234 species in in the paramos of Colombia and Venezuela (Gaviria,
119 genera and Berg (1998) found 168 species in 147 1997; Santana Castaneda, 1994; van der Hammen and
vegetation samples. In the Colombian eastern cordillera Cleef, 1986; Xena de Enrech, 1992). In Ecuador,
Van der Hammen and Cleef (1986) found 112 species in especially Gentianella and Draba are very conspicuous
69 genera, mostly in the lower superparamo belt (Cleef, superparamo genera, contributing probably with the
1981). A transect study in the Parque Los Nevados in highest number of endemic species (Jrgensen and
the central cordillera of Colombia, revealed 170 species Leon-Y, 1999; Leon-Y, 2000; Pringle, 1995). However,
growing above 4000 m (Cleef et al., 1983; Rangel et al., if Senecio were considered in a broad sense, including
1983). In this study we surveyed representative super- Dorobaea and Pentacalia, its total number of species
paramo sites throughout Ecuador on 16 mountains, but would be 17 and the genus would then be the richest in
at least ten distinct superparamo areas were not the superparamo. At the other extreme, 57% of the
explored (e.g., Carihuairazo, Quilindana, Sincholagua). genera have only one species. Such unequal distribution
In Venezuela, superparamo is conned to only three of species richness among genera was also found in the
areas in the surroundings of Merida (Monasterio, 1979), paramo ora of the eastern cordillera in Colombia by
and the low number of superparamo species reported Simpson and Todzia (1990), who argued that presence
from that country may therefore reect the actual in the paramo of many species-rich genera reected a
situation well. In Colombia, in contrast, there are many high degree of autochthonous speciation. The skewed
areas that reach the superparamo belt (Cleef, 1981), but taxonomic composition towards a few speciose genera
their total area may not surpass that of the Ecuadorian documented here for the Ecuadorian (super)paramo
superparamo (Hofstede et al., 2003). Different total would agree with that scenario.
areas of superparamo in Ecuador and Colombia may
determine the differences in species richness between
their oras. Nevertheless the number of known Colom- Variation in species richness among superparamos
bian superparamo species will certainly grow when more
sites are explored (A. M. Cleef, pers. comm.; P. Pedraza, We expected to nd a strong positive correlation
pers. comm.) so denitive conclusions concerning the between the area of the surveyed superparamo and its
relative number of species in the three countries cannot species richness (Simpson, 1974), but instead we found
be made until the species lists are more complete. that the correlation was weak and not signicant.
However, when the correlations were calculated for the
ecologically more homogeneous groups of mountains
Taxonomic composition separately, the expected positive correlation pattern did
appear, at least for the group of humid mountain sites
It is well known that Asteraceae and Poaceae dominate (Group 2 in Fig. 4). We also expected to nd a positive
the paramo ora (Luteyn, 1999; Luteyn et al., 1992; correlation between the vertical range of the superparamo
Ramsay, 1992; Ricardi et al., 1997) and this study and its species richness (Simpson, 1974), but instead we
conrms that these two families are the most species rich found a negative correlation. These unexpected correlation
also in the superparamo ora. Together they account for patterns reect the great ecological variability among the
one third of all superparamo species in Ecuador, which is superparamo sites. Small mountains with only species-rich
roughly the same proportion as in the paramo in general lower superparamo, such as Cajas, are more diverse than
426 P. Sklenar, H. Balslev / Flora 200 (2005) 416433

some of the bigger mountains with an overall impover- Species distributions

ished ora, such as Cotopaxi, Cotacachi, or Chimborazo-
west, despite their larger superparamo area and/or larger The superparamo ora is distinct from the grass
vertical range. Apparently, the exceptional superparamos paramo and subparamo belts below (Cleef, 1981;
of Cotopaxi and Chimborazo-west inuenced the overall Cuatrecasas, 1968; Ramsay and Oxley, 1997). Species
correlation pattern of species richness with area and also endemism is generally high in the superparamo (Berg,
produced the strong negative albeit not signicant 1998; Luteyn, 1992) although it may be even higher in
correlation among the ve dry sites (Group 1 in Fig. 4B). the montane forest (Balslev, 1988; Sklenar and Jrgen-
sen, 1999; but cf. Young et al., 2002). The proportion of
species endemic to Ecuador in our sample was estimated
at 23%, which is consistent with previous reports for the
Variation in b-diversity among superparamos Ecuadorian paramo (Leon-Y, 2000; Sklenar and Jr-
gensen, 1999). Two genera, Floscaldasia and Raouliopsis
We expected, and found a positive correlation (Asteraceae), seem to be restricted to superparamo
between b-diversity and the vertical range of the vegetation, but only the former reaches to Ecuador
superparamo (measured as number of surveyed altitu- (Cuatrecasas, 1979; Luteyn, 1999; Sklenar and Robin-
dinal levels), i.e., the longer the gradient of the super- son, 2000; Van der Hammen and Cleef, 1986).
paramo, the greater its b-diversity. As previously Almost one-third (29%) of the Ecuadorian super-
demonstrated, species composition and vegetation paramo species are restricted to a single mountain and
structure change considerably from the lower to the almost one-half of them (47%) are distributed in less
upper superparamo belts (Cleef, 1981; Sklenar, 2000; than four superparamo sites. This is consistent with
Sklenar and Ramsay, 2001), which would give greater b- ndings of Ramsay (1992) and Sklenar and Jrgensen
diversity on higher mountains (e.g., Cayambe and (1999), who reported a high number of narrowly
Iliniza mts.). The Cotopaxi superparamo is among the distributed species in the Ecuadorian paramo. The
most b-diverse in our sample despite the fact that it is number of very narrowly distributed superparamo
among the least species rich (Fig. 1). This paradox may species may be partly inated by the occasional
be related to the occurrence of a pioneer vegetation in occurrence of grass paramo and bamboo paramo
the Cotopaxi superparamo in addition to grass paramo species in the superparamo belt at one or a few
as documented by Stern and Guerrero (1997) and mountains. In addition some species reach Ecuador
Sklenar (2000). Lower mountains, where only lower only marginally. That is true for Valeriana henrici and
superparamo is developed (e.g., Yanaurcu, Cajas mts.), Lysipomia multiflora that are found in the Cajas region
have lower b-diversity values, even though the overall but are otherwise distributed in Peru (Ayers, 1999;
species richness may be high. Eriksen, 1989). Nevertheless, many Ecuadorian super-
Among the highest Ecuadorian mountains (Antisana, paramo species have a truly limited geographical
Cayambe, Cotopaxi, Iliniza, and Chimborazo) and distribution, e.g., Gentianella sulphurea (Fig. 5W) and
consequently those with the largest superparamos Loricaria antisanensis (Fig. 5X), and some are restricted
(Table 1), the Chimborazo-west has rather low b- to a single mountain. Other examples of such steno-
diversity. This superparamo is among the least diverse endemic species are Senecio ferrugineus at Chimborazo-
in Ecuador, but it is conspicuous because of its east (Sklenar, 2001b), Aphanactis antisanensis from
vegetation structure and presence of several central/ Antisana (Robinson, 1997), Oritrophium llanganatense
southern Andean oristic elements (e.g., Werneria from Hermoso (Sklenar and Robinson, 2000), and
apiculata, Stipa spp., Viola polycephala) and due to this several species from the Cajas region, e.g., Gentianella
it is sometimes compared to puna which is the dry longibarbata, G. hirculus (Pringle, 1995), Valeriana
paramo-equivalent in Peru (Acosta-Sols, 1985). secunda (Eriksen, 1989), and Draba steyermarkii.
Through the combined effects of climate (rain-shadow Mountain ridges and deep valleys acted as major
of the mountain) and human activities (burning and barriers to migrations of high-Andean plants during
grazing), this area has been altered into patchy, desert- glacial and interglacial periods (Jrgensen and Ulloa,
like superparamo (Sklenar and Lgaard, 2003) with 1994; Jrgensen et al., 1995). Many paramo species are
reduced species richness and b-diversity. Smaller moun- wind-dispersed or epizoochorous (Frantzen and Bou-
tains seem to exhibit consistently lower b-diversity, with man, 1989; Melcher et al., 2000, 2004; Simpson and
the exception of Chiles. Plant communities on Chiles in Todzia, 1990), which would make them capable of
general have lower a-diversity (less species per samples) dispersal over long distances. Superparamo species do
than communities at other sites of similar size but there tend to occur over broad vertical ranges which generally
is a high species turnover along the altitudinal gradient correlate with the species geographic distribution
in the rocky habitats (Sklenar, 2001a) which produce (Sklenar and Jrgensen, 1999). In our data, about
high b-diversity at this particular site. 60% of the species have a trans-Andean distribution,
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 427

i.e., they occur in both cordilleras and are distributed Due to its scattered occurrence along the Andes, the
across the deep inter-Andean valley. This is somewhat paramo has often been treated as an insular system
less than 72% of trans-Andean species previously (Simpson, 1974, 1975; Vuilleumier, 1970). In one
reported for the highlands of Ecuador (Balslev, 1988), classical study the extent of the paramo during glacial
but our number is certainly affected by the single- periods was found to be more important for explaining
mountain species. Trans-Andean distributions are found the patterns of paramo plant species diversity than the
in many species, e.g., Senecio nivalis, Nototriche spp., current extent of the paramo (Simpson, 1974). In
Pentacalia microdon (Fig. 5V), Saxifraga magellanica, Ecuador, species richness is only weakly correlated to
Arenaria dicranoides, and Xenophyllum rigidum (Fig. the present size of the individual superparamos. On the
5U), from the highest elevations, that should be the other side the extent of superparamo during glacial
most isolated from each other by the inter-Andean periods is not meaningful given the fact that the
valley. This may be a relictual occurrence of species assumed downward movement of vegetation belts would
which were widespread during glacial periods or it may merge almost all Ecuadorian areas of superparamo to
have resulted from a post-glacial over-land dispersal of one large conuent area. Although geographic distance
propagules along and across the Andes. Nevertheless, at various scales is (negatively) correlated to oristic
the pattern suggests that the distribution of (super)- similarity in the Andes (e.g., Luteyn, 2002; Ricardi et al.,
paramo plants is less affected by topographical barriers 1997; Simpson and Todzia, 1990) it does not explain the
than are the distributions of montane forest species observed general pattern of oristic similarity in the
where only about half of species have trans-Andean Ecuadorian superparamos. This nding of poor expla-
distribution (Balslev, 1988). natory power of geographic distance conrms previous
Although the species richness of separate super- results obtained from a sample of the high-paramo ora
paramo areas seems to be higher in Ecuador, one- (Sklenar and Jrgensen, 1999).
mountain endemics are much more common in the The presence of volcanic versus metamorphic bedrock
eastern cordillera of Colombia. In the Sierra Nevada del is the only signicant variable in the RDA, but it
Cocuy, for instance, 20% of the 107 superparamo accounts for only a small portion of the variability.
species are endemic (Van der Hammen and Cleef, 1986). Nevertheless, in three of the sites (Hermoso, Saraurcu,
We found that only about 3% and 2% of the species Quilimas), we thoroughly examined the boundary
found at Cajas and Chimborazo, respectively, were between volcanic and metamorphic basements, and we
endemic, and most other mountains have no steno- failed to discern any obvious changes in species
endemic species at all. Unlike the metamorphic eastern composition or vegetation structure when crossing the
cordillera of Colombia, the major Ecuadorian super- boundary, although Vargas et al. (2000) reported
paramos occur on volcanoes, several of which are still distinct changes in species composition for the Llanga-
active. Repeated destruction of vegetation through natis region where Hermoso is located. The cluster and
volcanic events followed by recolonization of the new indirect ordination analyses (Fig. 4) give a similar result;
substrate may not provide sufcient time between the superparamos of mountains built from metamorphic
successive eruptions for a speciation. It is interesting rocks are not clustered together but are grouped with
to note in this context that the paramo ora of the superparamos of nearby volcanoes.
volcanic central cordillera of Colombia is poorer in The four superparamo sites developed on meta-
species than that of metamorphic eastern cordillera morphic bedrocks all occur in the eastern cordillera
(Salamanca, 1992; Van der Hammen and Cleef, 1986). where they face the Amazon lowlands and therefore
An additional factor may be the size of individual receive the highest amounts of precipitation among the
superparamos; possibly the larger area of the Sierra studied superparamo sites (Bendix and Lauer, 1992;
Nevada del Cocuy compared to Ecuadorian paramo Hastenrath, 1981). We believe that the signicance of
areas (Hofstede et al., 2003) also contributed to the high geology (volcanic versus metamorphic bedrocks) for the
proportion of endemic species found there. An addi- species distributions (Fig. 5) in the RDA is an artifact
tional contributing factor could be the varying degree of and that this variable acts as a substitute for climate,
isolation of the areas today and in the past. While especially humidity on the sites. Unfortunately, there are
downward shifts of vegetation belts by 10001500 m no climatic data of the necessary spatial resolution
during glacial periods (Simpson, 1974; Van der Ham- available for the highest elevations of the Ecuadorian
men, 1974; Van der Hammen and Cleef, 1986) would Andes for a rigorous test of this hypothesis. Some
merge most areas of Ecuadorian superparamos and support for it, however, comes from data concerning the
facilitate species migrations between the now separated effect of a rain-shadow on species composition of two
superparamos, the Colombian Cocuy superparamo area Ecuadorian superparamos, Antisana and Chimborazo
would remain isolated from other areas of superparamo (Sklenar and Lgaard, 2003), that was based on
allowing longer time for speciation. Phylogeographic published precipitation measurements. Other circum-
studies of wide-spread species could test this hypothesis. stantial evidence for the effect of humidity on the
428 P. Sklenar, H. Balslev / Flora 200 (2005) 416433

distribution of superparamo species comes from the Trisetum spicatum (Figs. 5S and T). The remaining four
observation that some species that are mostly restricted mountains (Cayambe, Imbabura, Chimborazo-east, and
to the humid superparamos from the eastern slopes Cajas) basically differ from the very humid superparamo
(e.g., Distichia muscoides, Caltha sagittata, Cuatrecasa- by the absence of the above species rather than by
siella isernii), can be found in wet sites along springs or forming a homogeneous group among themselves.
small depressions in the terrain that occasionally occur However, in agreement with previous studies that
in the dry superparamos (Sklenar, 2000; Ceron, 1994). pointed to the peculiarity of the Cajas region (e.g.,
Sklenar and Jrgensen, 1999) this site occurred well
separated from the other sites in the Correspondence
Floristic similarity among superparamos Analysis. Volcanic activity terminated during the
Tertiary in the southern Ecuadorian Andes, including
Based on the distribution of species from the montane Cajas (Hall, 1977), whereas the northern paramos all
forests and paramos, two oristic regions have been experienced volcanic activity up till present time. The
proposed for the high Andes of northern and central different volcanic histories may be reected in the
Ecuador, with the divide being the inter-Andean valley second CA ordination axis.
(Jrgensen and Ulloa, 1994). Our data suggest two Simpson (1983) pointed to the importance of the
somewhat different regions. Although the climatic data ecological similarity between areas to explain the
is very limited, we attribute this pattern to differences in composition of different Andean oras. At the geo-
humidity based on our eld observations of vegetation graphic scale studied here of tens to hundreds of
physiognomy. Most mountains of the eastern cordillera, kilometers, we suggest that ecological similarity, con-
northern Ecuador (Chiles, Cotacachi, Imbabura), Chim- sidered mainly as humidity, is more important for
borazo-east (in the western cordillera), and Cajas in the distribution of superparamo species than other factors,
south belong to humid superparamos (Group 2 in Fig. such as geographic distance or topographic separation
4B), which are characterized by the presence of by valleys. Our ndings support previous works that
Jamesonia spp., Geranium sibbaldioides, Lachemilla stressed the importance of precipitation for vegetation
holosericea, F. asplundii, Carex sect. Acicularis, and composition by distinguishing humid and dry super-
Luzula gigantea among others (Figs. 5GL). The dry paramos in the Parque Los Nevados, Cordillera Central
superparamos (Group 1 in Fig. 4B), are distinguished by of Colombia (Perez and Van der Hammen, 1983), and
the occurrence of Festuca vaginalis, Plantago nubigena, others who recognized rain-shadow desert paramo on
Astragalus geminiflorus, Bidens andicola, Conyza carda- the western side of Chimborazo (Ramsay, 1992).
minifolia, Calamagrostis mollis, Cerastium imbricatum, Several interesting questions remain for further ex-
and Silene thysanodes (Figs. 5AF; although the latter ploration in the high paramo. Some of these are: How
two were also found in some more humid super- can we explain the smaller number of one-mountain
paramos), and comprise Iliniza, Pichincha, Cotopaxi, endemic species in Ecuador compared to Colombia? At
and the Chimborazo-west and Antisana-west. what geographic scale do island biogeographical factors
Among the dry superparamo sites, Chimborazo-west become more important than ecological factors? Do grass
stands out from the other sites (see above). Among the paramo and subparamo species conform to the pattern as
humid superparamo sites, two groups are apparent. observed in the superparamo ora?
Sites from the eastern cordillera (except Cayambe),
Chiles and possibly also Cotacachi from the western
cordillera represent very humid superparamos that
receive high amounts of precipitation originating in
the Amazon basin. Among them, Tungurahua is Acknowledgements
exceptional, being an active volcano on which the
vegetation is constantly recovering from the inuence Petr Sklenar thanks the Danish Research Academy
of volcanic events (Sklenar, 2000). These very humid and the Grant Agency of the Czech Republic (Grant
superparamo sites are characterized by the occurrence No. 206/97/1198) and Henrik Balslev thanks the Danish
or abundance of species, such as Huperzia rufescens, Natural Science Research Council (Grant No. 21-01-
Nertera granadensis, Loricaria complanata, Calamagros- 0617) for support. Two anonymous reviewers are
tis guamanensis, C. ecuadoriensis, Draba spruceana, and thanked for suggestions to improve the quality of the
Xenophyllum sotarense (Figs. 5MR). These species paper. Both authors acknowledge Renato Valencia,
often have a strictly cis-Andean distribution, but usually director of QCA herbarium, P.U.C.E., Quito, for
do not occur on all of those mountains. These very research facilities, INEFAN, Quito, for research and
humid sites are also characterized by the absence of a export permits. Veronika Kosteckova is thanked for
group of species that occur only in the dry or moderately assistance in the eld and Flemming Nrgaard for
humid superparamo sites, such as Perezia pungens and producing gures.
Appendix 1

Matrix of OBSERVED (above diagonal) and EXPECTED numbers of species held in common (below diagonal) between pairs of sites, diagonal indicates
the number of species for each site used in the analysis.

P. Sklenar, H. Balslev / Flora 200 (2005) 416433

Chil Cota Imba Caya Sara EAnti WAnti Pich Ilin Coto WChim EChim Herm Tung Altar Quil Yana Cajas

Chiles 100 62 60 67 67 70 44 51 49 26 30 56 64 34 69 67 55 61
Cotacachi 53.8* 102 77 78 59 69 61 76 69 39 41 64 62 43 72 67 59 71
Imbabura 57.9 59.4*** 114 92 55 66 66 74 72 42 43 71 59 42 68 64 54 69
Cayambe 66 67.4** 73.4** 137 69 84 85 92 91 50 53 93 72 46 82 77 63 83
Saraurcu 49.6*** 50.8* 54.6 61.7* 93 77 45 49 47 22 26 60 74 39 76 76 63 60
Antisana-east 63.2* 64.5 70.1 80.6 59.4*** 129 67 66 62 35 33 77 84 48 84 84 66 73
Antisana-west 66.7*** 68.4* 74.4* 85.8 63.1*** 82.1*** 140 87 102 66 69 87 53 39 62 72 49 77
Pichincha 63.5*** 65.1** 70.7 81.1** 59.7** 77.8** 82.4 130 95 62 61 83 54 45 68 67 55 71
Iliniza 68.5*** 70.2 76.2 87.9 64.5*** 84.1*** 89.4** 84.4** 143 74 73 83 48 42 67 67 53 79
Cotopaxi 48.7*** 49.6*** 53.6** 60.7** 46*** 58.5*** 62 58.8 63.3** 91 54 46 30 29 34 34 30 47
Chimborazo-west 53.6*** 55*** 59.3*** 67.4*** 50.8*** 64.9*** 68.5 65.1 69.8 49.9 103 53 29 27 44 41 35 53
Chimborazo-east 63.3* 64.9 70.2 80.8** 59.6 77.3 81.9 77.7 84.1 58.6*** 64.6** 129 58 45 73 76 61 78
Hermoso 59.1 60.5 65.2 74.8 55.6*** 71.5*** 76*** 72.1*** 77.7*** 54.7*** 60.3*** 71.6*** 117 48 74 83 65 68
Tungurahua 39.6* 40.6 43.7 49.1 37.6 47 49.8*** 47.5 50.6** 37.1** 40.6*** 47.1 44.3 71 45 46 43 39
El Altar 63.2 64.7* 70.1 80.7 59.6*** 77.3 82.2** 77.7** 83.9*** 58.6*** 64.6*** 77 71.4 47.1 130 92 87 82
Quilimas 65.2 66.9 72.5 83.5 61.3*** 79.9 84.8** 80.2*** 87.1*** 60.3*** 66.8*** 80 73.9* 48.5 80*** 135 80 83
Yanaurcu 54.9 56.2 60.6 69.2 51.8*** 66.3 70.2*** 66.5*** 71.9*** 50.8*** 56*** 66.1 61.9 41.5 66*** 68.3** 106 65
Cajas 69* 70.7 77* 88.8 64.9 84.8** 90.4** 85.6*** 92.8*** 63.6*** 70.9*** 85 78.6** 51*** 84.7 87.7 72.4* 147
Species pool 374, signicant differences marked with asterisks (0.05Xp40.01*, 0.01Xp40.001**, pp0.001***).

Appendix 2

Matrix of geographic distances between the sites in kilometers (above diagonal) and matrix of oristic dissimilarities measured by w2 distance (below
diagonal) which were used in the Mantel test.

P. Sklenar, H. Balslev / Flora 200 (2005) 416433

Chil Cota Imba Caya Sara E Anti WAnti Pich Ilin Coto WChim EChim Herm Tung Altar Quil Yana Cajas

Chiles 0 64 64 85 100 140 147 126 183 172 272 268 220 256 281 290 345 434

Cotacachi 1.25 0 22 54 70 94 98 64 122 115 212 209 171 202 228 237 292 375
Imbabura 0.076 5.215 0 33 49 80 85 64 120 108 208 204 159 193 218 228 282 370
Cayambe 0.015 1.667 4.713 0 17 56 63 67 111 93 193 188 136 172 196 205 260 354
Saraurcu 6.104 1.324 0.003 0.864 0 45 52 71 107 85 185 179 123 161 184 193 248 344
Antisana-east 0.732 0.314 0.24 0.143 5.215 0 8 58 69 43 140 134 80 117 141 150 205 299
Antisana-west 7.73 0.8 0.95 0.01 5.19 2.78 0 56 62 35 133 126 74 110 134 143 198 292
Pichincha 2.46 1.825 0.154 1.465 1.92 1.79 0.257 0 58 57 149 147 118 143 169 179 232 312
Iliniza 5.55 0.02 0.23 0.109 4.75 5.81 1.776 1.331 0 30 91 89 79 92 118 127 177 254
Cotopaxi 10.6 2.27 2.51 1.89 12.5 9.44 0.258 0.174 1.809 0 101 96 61 87 113 122 176 263
Chimborazo-west 10.4 3.56 4.48 3.08 12.1 15.7 0.004 0.26 0.147 0.337 0 11 81 48 57 62 96 163
Chimborazo-east 0.84 0.01 0.009 1.842 0.003 0.001 0.318 0.362 0.01 2.71 2.08 0 71 37 47 53 92 167
Hermoso .406 0.037 0.59 0.1 6.089 2.185 6.96 4.54 11.4 11.2 16.2 2.58 0 41 62 70 125 225
Tungurahua 0.79 0.142 0.07 0.2 0.052 0.021 2.34 0.13 1.46 1.77 4.56 0.09 0.309 0 26 35 89 185
El Altar 0.532 0.824 0.06 0.021 4.513 0.581 4.96 1.21 3.4 10.3 6.57 0.21 0.095 0.09 0 9 64 165
Quilimas 0.05 0.0001 1 0.51 3.525 0.21 1.93 2.17 4.64 11.5 9.96 0.2 1.121 0.13 1.8 0 55 158
Yanaurcu 0.0002 0.14 0.72 0.56 2.422 0.001 6.4 1.99 4.97 8.52 7.88 0.39 0.155 0.054 6.682 2.004 0 112
Cajas 0.93 0.001 0.83 0.38 0.37 1.64 1.99 2.49 2.05 4.33 4.52 0.58 1.43 2.82 0.09 0.25 0.76 0
The positive or negative signs with the w2 values indicate respectively whether or not the observed number of shared species is higher or lower than the expected number.
P. Sklenar, H. Balslev / Flora 200 (2005) 416433 431

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