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8/8/2019 Topographic Controls on the Distribution of Tree Islands in the High Andes of South-western Ecuador
1/13
O R I G I N A LA R T I C L E
Topographic controls on the distribution
of tree islands in the high Andes of
south-western Ecuador
D. Coblentz1* and P. L. Keating2
INT RO DUCT IO N
Explaining the spatial distribution of plant communities has
long been a goal of ecologists (e.g. Forman & Godron, 1986;
Turner et al., 2001). A long history of investigations has
established that the nature of the topographic fabric, typically
evaluated through geomorphometry, or the science concerned
with the geometry of the landscape (Chorley et al., 1957),
1Los Alamos National Laboratory, Los Alamos,
NM, USA and2Department of Geography,
Indiana University, Bloomington, IN, USA
*Correspondence: David Coblentz, MS F665,
Los Alamos National Laboratory, Los Alamos,
NM 87545, USA.
E-mail: [email protected]
ABS T RACT
Aim To evaluate the hypothesis that geomorphometric parameters of upper
montane Andean environments have an important influence on the regional fire
ecology and consequently play a role in the spatial distribution of remnant tree
islands dominated by Polylepis.
Location A glacial landscape located between 3600 and 4400 m elevation in
Cajas National Park, south-western Ecuador.
Methods The eigenvalue ratio method was used to evaluate the regional
geomorphometric parameters of a 30-m digital elevation model for CajasNational Park. The landscape character was evaluated by quantifying the
topographic roughness, organization, and gradient. This information was used to
determine the spatial correlations between terrain characteristics and the
distribution of tree islands in the region.
Results We demonstrate a strong spatial correlation between areas of high
topographic roughness and gradient, and the locations of the major tree islands.
We find that there is a distinctive relationship between the topographic roughness
and organization in the vicinity of the tree islands (e.g. increased upslope
roughness and decreased topographic grain strength) that substantiates the
notion that the tree islands are located in relatively inaccessible topography.
Main conclusions In the northern and central Andes, the location ofPolylepis-dominated forest islands has been shown to be a function of climate, terrain
characteristics, and anthropogenic disturbances. Although the relative importance
of various ecological factors has been debated, it remains clear that fires have
exerted a strong influence on these ecosystems. Other authors have noted that
tree islands are more likely to occur at the base of cliffs, above moist areas, and in
other areas where fires do not burn frequently. Our results corroborate these
observations, and demonstrate that the occurrence ofPolylepis patches is strongly
correlated with specific combinations of terrain features. Although we do not
discount the importance of other factors in determining the spatial position and
areal extent of these forests, we demonstrate strong support for fire-related
hypotheses.
Keywords
Andes, Cajas National Park, fire ecology, geoecology, geomorphometry,
landscape ecology, Polylepis, remote sensing, topographic analysis, tropical
montane forest.
Journal of Biogeography(J. Biogeogr.) (2008) 35, 20262038
2026 www.blackwellpublishing.com/jbi
2008 The Authorsdoi:10.1111/j.1365-2699.2008.01956.x Journal compilation 2008 Blackwell Publishing Ltd
8/8/2019 Topographic Controls on the Distribution of Tree Islands in the High Andes of South-western Ecuador
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plays a fundamental role in the distribution of vegetation and
biodiversity (Coblentz & Riitters, 2004). Despite the important
link between topography and geoecology in montane regions,
most studies of this relationship have been limited to a
primarilyqualitative framework (e.g. Brown & Gibson, 1983).
Digital elevations models (DEMs), rectangular grids of eleva-
tion values, provide an unparalleled tool for the evaluation of
geomorphometric parameters. The current availability of high-
speed computing platforms, high-resolution DEMs now glob-
ally available at a 3-arc-second (about 90 m) resolution, and
high-resolution land cover data sets (e.g. Riitters et al., 2000)make it possible to undertake a rigorous quantitative analysis
that can be used to test the relationships discussed in previous
qualitative investigations.
In the Andean highlands, the spatial distribution of tropical
forests has been shown to be a function of both topographic
features and anthropogenic disturbances (e.g. Troll, 1968,
1973; Young & Keating, 2001). Elevation establishes a complex
set of environmental gradients, including temperature regimes
and precipitation (Troll, 1968; Lauer, 1981). In addition,
upper montane ecosystems are strongly influenced by
topographic features such as slope, aspect, and shape (Troll,
1973; Young, 1998a; Keating, 1999). Given that humans have
been present in the Andes for at least 7000 years (Bruhns,
1994), disturbances have also influenced the composition,
structure and spatial position of forest communities. Agricul-
tural practices, burning, grazing, road construction, and
logging have continually modified many areas of the Andes
(Ellenberg, 1979; Lgaard, 1992; Young, 1994, 1998b; Keating,
1998), leading to a land cover that is characterized by a
complex mosaic of vegetation types (Gade, 1999). Under-
standing the timing and exact nature of landscape transfor-mations is difficult, as each region has a different landscape
history (Young & Zimmerer, 1998). Burning, however, con-
tinues to occur in conjunction with most land-use activities,
especially grazing, and is probably the most significant
disturbance in the Ecuadorian Andes above 3500 m elevation
(Keating, 2007).
At elevations above 3000 m, the Ecuadorian Andes are
covered with a spatially heterogeneous assemblage of agricul-
tural lands, paramo (tropical alpine) and forest communities.
Whereas continuous forest covers the outer flanks of both
Figure 1 Topographic map of south-wes-
tern Ecuador with the study area indicated by
the solid box. In the lower figure, the tree
islands of Cajas National Park are designated
by polygons with colours representing four
area designations: red corresponds to tree
islands covering an area < 1 ha; green to
15 ha; blue to 510 ha; and black to
> 10 ha.
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cordilleras up to 4300 m elevation, inter-Andean forest (sensu
Sarmiento, 1995) tends to occur as remnant forest patches in
remote locations (Valencia & Jrgensen, 1992; Stern, 1995;
Young & Keating, 2001). Some continuous forest occurs in the
southern tip of the country (Madsen & llgaard, 1994), but
intensive, long-term land use has reduced most Andean forests
to a series of strips and patches found in gullies, ravines,
volcanic calderas, and remote high-elevation sites. Fragments
of fire-sensitive vegetation are characteristic of many fire-
prone landscapes (Bowman, 2000), and topographic protec-
tion of remnant vegetation communities is well documented in
several tropical ecosystems (e.g. Fensham, 1995).
In many sections of the Andes, including south-western
Ecuador, high-elevation patches ofPolylepis-dominated forests
occur above the timberline separating continuous, closed
forest from paramo (Fig. 1). Polylepis is a taxonomically
complex genus comprising c. 20 species whose centre of
diversity occurs between central Peru and southern Bolivia
(Kessler, 1995; Navarro et al., 2005). Often found at up to
5000 m elevation in Bolivia, these trees occur at the highest
elevation of any angiosperm tree taxa (Navarro et al., 2005).Seven species of Polylepis are found in Ecuador, where they
occur between 2700 and 4300 m a.s.l. (Romoleroux, 1994).
Whereas Polylepis incana may occur as monospecific stands
(Fehse et al., 2002), most Polylepis forest islands in Ecuador
contain other tree taxa, including Columella, Escallonia and
Gynoxys (Fjeldsa, 1992; Romoleroux, 1994).
Several thousand years ago, Polylepis forests probably
covered a far more extensive area than they do at present.
Palaeohistorical evidence suggests that the geographical distri-
butions of these forests fluctuated widely, largely in response to
climatic changes (Simpson, 1979). As a result of the dramatic
landscape transformations described above, however, there has
been a substantial reduction in the area covered. Fires, grazing
and wood-cutting appear to be the principal factors explaining
this change, and these activities continue to restrict the spatial
extent of upper montane forests (e.g. Fjeldsa, 1992; Hensen,
2002; Enrico et al., 2004; Renison et al., 2006). Given the
frequency of these disturbances, these forests now occur
primarily in ravines, on steep rocky terrain, near streams, and
in other protected sites (Lgaard, 1992; Kessler, 1995). Recent
studies indicate that only the upper limits of Polylepis patches
are determined by harsh climatic conditions, which impede the
growth and regeneration of trees (Lgaard, 1992; Cierjacks
et al., 2007); under these conditions, certain sites afford the
seedlings protection from the wind and cold (e.g. Simpson,1986; Ibisch, 1993).
In this article we focus primarily on the notion that fires play
a key role in maintaining the current spatial distribution of
forest patches. Our study is motivated by the hypothesis that
the tree islands in Cajas National Park are located in areas that
experience lower fire frequencies owing to local topographic
variations such as steep terrain. Other issues, such as climatic
factors and disturbance history, are beyond the scope of this
study. To test our hypothesis, we apply the eigenvalue ratio
method (Woodcock, 1977) to quantify the character of the
landscape and to evaluate the terrain where Polylepis patches
occur in a section of south-western Ecuador, and discuss
several salient trends in relation to the presence of fire. As
discussed below, this technique extracts several types of
information about the topographic fabric of the region, and
provides a comprehensive topographic analysis.
M AT E RIALS AND M E T HO DS
Study area
El Cajas National Park covers c. 29,000 hectares of upper
montane ecosystems on the western cordillera of the Ecuado-
rian Andes (Ministerio de Ambiente, 2007). Located c. 35 km
west of Cuenca, the middle of the park is found at 2 50 S,
7915 W (Fig. 1). Elevations within the park range from
c. 3180 to 4500 m. With the area having been glaciated during
the Pleistocene (Hastenrath, 1981; Clapperton, 1993), the
physiography includes more than 230 glacial lakes, intercon-
nected ridges and peaks, and numerous broad hanging valleys.
Glaciers were absent during the Holocene, and little tephra orpyroclastic materials from volcanoes have fallen on this region
(Rodbell et al., 2002). Soils in El Cajas did not develop from
underlying rocks, but instead are composed of organic and
aeolian materials (Harden, 2007), and are often referred to as
black Andean soils, histosols, or hydric andisols.
The dominant land cover is paramo, a form of tropical
alpine vegetation. Much of the valley bottom is characterized
by bogs or moist grasslands, whereas the slopes are better
drained and covered with tussock grasses. Woody species
occur infrequently in the grasslands, and shrubs are typically
found in isolated patches on the valley floors. Plant commu-
nity patterns are relatively uniform in the tussock-grass
paramo but vary markedly along the valley bottoms (Keating
et al., 2002). The species richness of this ecosystem is probably
as high as in any paramo in the country, and its floristic
composition differs significantly from that of paramos located
to the north of this region (Sklenar & Jrgensen, 1999; Sklenar
& Ramsay, 2001).
Three varieties of forest are found in the park. Continuous,
relatively undisturbed forest occurs in the lower sections of the
park; timberlines usually occur between 3330 and 3365 m a.s.l.
Second, a diverse mixture of secondary forest and tall shrubs
occurs near roads and along some valley bottoms. Finally,
forest patches, which often occur at the base of cliffs or steep
slopes (Fig. 2), are typically found above 3600 m a.s.l. Theseforest islands are often dominated by 510 m tall Polylepis
trees, although the interior portions may contain additional
tree genera, including Clethra, Gynoxys, and Myrsine. Generally
speaking, the relative importance of Polylepis increases above
3800 m (Hansen et al., 2003).
Climatic data have not been collected extensively for this
section of El Cajas, but it has been suggested that the paramos
in this region receive at least 2500 mm of precipitation
annually (Morris, 1985). Most of this falls between January and
June, and a relatively dry season occurs during the rest of the
D. Coblentz and P. L. Keating
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year (Blandn, 1989). Maximum daily temperatures above
3500 m are usually between 3 and 7C when clouds are present
(P.L. Keating, personal observation); minimum temperatures
are usually between 0 and 4C.
The long-term environmental history of these highlands has
been investigated only recently, and much of the disturbance
history is a matter of speculation. After the late glacial period,
which ended 11,000 yr bp, Polylepis increased in importance at
higher elevations. Charcoal evidence suggests that fires also
became more frequent at that time; however, it appears that
this frequency decreased somewhat around 4000 years ago
(Hansen et al., 2003). The role played by humans in modifying
the landscape, however, is not clear; nor is there any
documented spatial correlation between forest communities
and the occurrence of fires.
Although the Ecuadorian highlands have been inhabited by
various indigenous groups during the past several thousand
years, past land-use practices and associated disturbance
regimes in the Cuenca region have received little attention.
Several anthropologists have examined pre-Conquest settle-
ment patterns and material culture (e.g. Collier & Murra, 1943;
Bruhns et al., 1990), and other authors have described more
recent settlement patterns in the region (e.g. Borrero, 1989).
To our knowledge, however, no one has provided detailed
descriptions of human impacts within El Cajas for any periodbefore the 20th century. It is worth noting, however, that early
Spanish explorers described these mountains as lacking forests
in the 1580s (Jimenez de Espada, 1965[1582]). We were unable
to find photographs or other records that document forest
cover change at any time during the past four centuries.
Between 1979 and 1996, this park was referred to as Las
Cajas Recreational Area (Fundacion Natura, 1992). Because
the designation of recreational areas in Ecuador does not
entail strict conservation protection, both burning and grazing
by livestock have precluded the regeneration of woody species
throughout the region. During the dry season, several fires
could occur simultaneously during any given day (Keating
et al., 2002). Many of the burns cover more than several
hundred hectares, so that the paramo represents various states
of recovery after these disturbances. Clearly, forest could not
regenerate under such conditions, and shrub paramo (sensu
Cuatrecasas, 1968), which often serves as a transition com-
munity between forests and grass paramo, rarely occurs. Given
that this region has been under the influence of human activity
for at least 3500 years (White & Maldonado, 1991), it is a
reasonable assumption to treat the forest patches within El
Cajas as relatively stable units. As we illustrate below, the
position and areal extent of these forest islands have probably
not changed during the past several decades. Only in rare cases
did forest margins show any signs of recent burning.
Analysis of vegetative cover
The vegetative cover in the study area was quantified through
an analysis of a 15 15 km subset of a Landsat Thematic
Mapper (TM) image (30-m cell size) acquired on 26 March1987. Although more recent imaging would be desirable,
Landsat, SPOT and Aster images collected subsequent to this
date have included substantially more cloud coverage and thus
are of limited utility for mapping land cover. Atmospheric
correction was performed with a technique described in Colby
& Keating (1998). The subscene was rectified using 23 ground
control points, and a root mean squared error (RMSE) of
0.230 was obtained. A nearest neighbour resampling algorithm
was used, and the rectified image was subset to the coordinates
given below. The normalized difference vegetation index
(NDVI) was calculated to distinguish the forested areas from
other land-cover types. Calculated as near infrared ) red/near
infrared + red, or TM bands 4 ) 3/4 + 3, this index provides
an indication of leafy green biomass (Sader et al., 1989; Mausel
et al., 1993). Because the forested areas contained significantly
more leafy biomass than the paramo vegetation, a density slice
technique (Jensen, 2005) was utilized to map the forest islands
and secondary forest. After the range of NDVI values was
determined for each cover type, a threshold was established to
separate trees from other land-cover types; aerial photographs
acquired on 9 October 1976 were used to assist in this
classification process.
To determine the characteristics and diversity of vegetation
that occurs within Cajas National Park, extensive field
reconnaissance was performed during June 1990 and October1991. Initially, a 9 12 km2 area was selected that included
representative physiographic features, numerous forest
patches, and several hundred hectares of secondary
forest located near roads and in valleys; continuous, primary
forest was not included in the study area. UTM coordinates
(Zone 17 S) for the upper left corner of the study site are
695000 E, 9696000 N, and those for the lower right corner are
707000 E, 9687000 N. Further ground verification was per-
formed during June and July 1995, and aerial photographs
acquired on 12 December 1995 were also used to check the
Figure 2 A typical tree island located in the south-west portion
of the study area at c. 3900 m in Cajas National Park. Dominated
by Polylepis, the forest patch covers c. 10 ha and is located in a
boulder-strewn area at the base of a cliff. Note that the tussock-
grass paramo surrounding this site has been burned within the
past year.
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accuracy of the mapping process. Comparison of photographs
from both dates revealed that little, if any, change in forest
cover has occurred during the past 20 years. Approximately
15% of the scene was covered with clouds or cloud shadows,
and these areas were not included in the analysis.
Evaluation of geomorphometric parameters
Geomorphometry, the study of landscape geometry, provides a
way to quantify the relationships between topographic vari-
ables and vegetative land cover. Within biogeography, there
has been a growing recognition that topographic parameters
exert a strong influence on patterns of vegetative cover (see
discussion in Coblentz & Riitters, 2004), and are valuable in
wildlife habitat modelling (Sappington et al., 2007). In the
following analysis we employ a quantitative analysis of
topography using the evaluation of geomorphometric param-
eters (see review in Mark, 1975). Specifically, we use the
eigenvalue ratio method (see discussion in Appendix S1 and
Fig. S1) to quantify topographic roughness, organization and
gradient across the landscape (Chapman, 1952; Watson, 1966;
Woodcock, 1977; Guth, 1999; McKean & Roering, 2003). As
7915'W 7912'W 7909'W
248'S
245'S
3500 4000
Topography (m)
7915'W 7912'W 7909'W
248'S
245'S
0.0 0.2 0.4 0.6 0.8 1.0
Roughness
7915'W 7912'W 7909'W
248'S
245'S
0 1 2 3 4 5
Organization
7915'W 7912'W 7909'W
248'S
245'S
0.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4
Gradient
(a) (b)
(c) (d)
Figure 3 The spatial relationship between the (a) topography, (b) topographic roughness, (c) topographic organization, and (d) topo-
graphic gradient for the study area shown in Fig. 1 computed using the eigenvalue ratio method. The white-filled polygons designate
the spatial extent of the tree islands, which are generally located near regions of high topographic roughness, high gradient and low
organization, suggesting a correlation between the location of the tree islands in Cajas National Park and the rough topographic features in
the landscape.
D. Coblentz and P. L. Keating
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discussed above, our study is motivated by the hypothesis that
the tree islands in Cajas National Park are located in areas that
experience lower fire frequencies owing to local topographic
variations such as steep terrain. We test this hypothesis by
determining spatial patterns of geomorphometric parameters
and relating them to the locations of the tree islands. In
addition, through the use of histograms and other graphical
techniques, we demonstrate relationships among topographic
variables, elevation, and size of forest patches. A summary of
the topographic analysis methodology used to quantify various
geomorphometic parameters is provided in Appendix S1.
Because the objectives of our topographic analysis require
precise measurements, we constructed a 30-m DEM. Althoughthis resolution gives a threefold improvement in the topo-
graphic resolution over commonly available DEMs (e.g. SRTM
data), we acknowledge that it is still too coarse for a reliable
evaluation of the metre-scale topographic details that would be
required for a study of individual tree islands. The conclusions
of our study are therefore limited to topographic features with
a wavelength of about several tens of metres, and should be
considered a first-order analysis of the broad-scale topographic
characters that contribute to the existence and distribution of
the tree islands in Cajas National Park. A more detailed
geomorphometric analysis of individual tree island locales is
left for future research efforts when higher-resolution DEMs
(< 10 m) become available.
RE S ULT S
The study area, shown in Fig. 3a, covers an area of c. 90 km2
and has considerable topographic relief, with elevations
ranging from 3361 to 4406 m a.s.l., with a mean and standard
deviation of 3892 and 115 m, respectively. In general, the tree
islands (white polygons in Fig. 3) are located in close
proximity to areas of high topographic roughness (red regions
in Fig. 3b) and low topographic organization (Fig. 3c). Areasof high roughness generally correspond to high topographic
gradients (Fig. 3d). The correlation between tree island
locations and steep/rough topography is stronger for the
larger tree island areas (> 10 ha), which also tend to be located
at high elevations (at an average of 3900 m vs. about 3850 m
for the smaller tree island patches). As shown in Fig. 4, the
elevation distributions for the larger tree islands are also
skewed towards higher elevations with a smaller standard
deviation for the distribution (c. 95 m vs. a standard deviation
of > 120 m for the tree island sites covering smaller areas).
0
2
4
6
8
10
%
3000 3500 4000 4500
Elevation (metres)
0
2
4
6
8
10
%
3000 3500 4000 4500
Elevation (metres)
0
5
10
15
%
3000 3500 4000 4500
Elevation (metres)
0
5
10
15
%
3000 3500 4000 4500
Elevation (metres)
0
5
10
15
20
25
%
3000 3500 4000 4500Elevation (metres)
Random 10 ha
N: 570 mean: 3892 m N: 2010
mean: 3873 m
N: 6659 mean: 3879 m N: 4289 mean: 3855 m
N: 2785
mean: 3909 mFigure 4 Histograms of the elevation dis-
tribution for tree island sites with areal
extents of < 1 ha, 15 ha, 510 ha, and
> 10 ha. For reference, the elevation distri-
bution for 570 random sites in the study area
is also shown. Although the average elevation
of the various tree island sites is similar to the
average elevation of the random sites, there
are significant differences in the topographic
character (Figs 5 & 6).
Topographic controls on tree islands
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Variations in the character of the topographic roughness
and topographic organization between the tree island sites and
a reference set of 481 sites (randomly selected within the study
area) is shown in Fig. 5, with mean and standard deviation
values listed in Table 1. Several generalizations can be made on
the basis of these distributions. The low topographic organi-
zation for the highest tree island sites (blue) is indicative of a
high degree of scatter in the topographic roughness distribu-
tion (quite similar to the case for a randomly oriented
topographic distribution) possibly reflecting the geomorphic
processes of Pleistocene-era glaciation. Moreover, within the
highest-elevation belt, grass height and coverage are typically
lower, and other life forms such as cushion plants increase in
importance. Fires not only are less likely to be ignited at highelevations, but are less likely to consume this relatively
heterogeneous vegetation.
The topographic character of the smallest size class (< 1 ha)
is closest to that of the random sites. This suggests that, given
the spatial limitation of the DEM used in this analysis (cell size
of 0.09 ha), we may not have detected terrain features that
influence the locations of these islands. Tree islands at lower
elevations (red distributions) have the most variable and
highest organization, and consistently higher roughness values
which is expected given the mechanism by which glaciers
carve landscapes. We also note that the difference between the
lower (red) and higher (blue) tree island sites increases with
forest island area in both the roughness and organization
distributions (Fig. 5). Sites with high values of both organi-
zation and roughness (e.g. lower-elevation tree islands with
spatial extent greater than 10 ha) span regions with greater
topographic variation, and the higher coherence in the surface-
normal vectors (which define the surface orientation) in the
flatter regions increases the organization values compared with
those in the rougher regions. As expected, this effect is more
pronounced for the larger tree island sites.
We applied the KolmogorovSmirnov test (KS-test) to
determine if the tree island histograms for each elevation range
and size subset differed significantly from the random sampledistribution; this test has the advantage that it is not based on
any underlying distribution of data. The D-statistic listed in
Table 1 is a measure of the difference between the randomly
selected distribution and the distribution at the listed elevation
and size subset. The confidence level that the histograms are
sampling different distributions is reflected in the significance
level probability. For reference, identical distributions would
return values of 0.0 and 0% confidence for D and the
confidence probability, respectively. The null hypothesis that
the histograms are sampling the same distribution can be
0
5
10
15
0.0 0.5 1.0
Roughness
Random
0.5 1.0 0.5 1.0 0.5 1.0 0.5 1.0
0
5
10
Organization
Random
0 4 8 0 4 8 0 4 8 0 4 8 0 4 8
(a)
(b)
Figure 5 (a) Histograms showing the elevation dependence on the topographic roughness for the tree island sites with areal extents of
< 1 ha, 15 ha, 510 ha and > 10 ha, and a reference set of 481 sites (Random). (b) Histograms showing the elevation dependence on the
topographic organization for the tree island sites with areal extents of < 1 ha, 15 ha, 510 ha and > 10 ha, and a reference set of 481 sites
(Random). The histograms are colour-coded to correspond to three elevation ranges: red (35003700 m), green (37003900 m) and blue
(39004100 m); the number of samples for each of the elevation ranges is also colour-coded. Averages, standard deviations, n-values and the
results of the KolmogorovSmirnov test applied to the distributions are listed in Table 1.
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rejected at a > 99% confidence level for nearly all classes, with
the exception of the high-elevation subregion for 15 ha
patches and the lowest-elevation subregion for 510 ha tree
island patches although we note that the KS-test confidence
level for these two regions is still in excess of 75%. These results
support our a priori assumption that the geomorphological
parameters of the tree island locations are significantly
different from those of the region (as measured by a random
sampling).
The trade-off between topographic roughness and topo-
graphic organization can be graphically illustrated given the
fact that only two of the three eigenvalues are independent.
The flatness [ln(S1/S2)] (the reciprocal of roughness) can be
plotted against organization [ln(S2/S3)] to eliminate one of the
dependent variables and to provide a way to describe thepattern of vector orientations as clusters (unimodal distribu-
tion of the vectors) or girdles (multimodal distribution of the
vectors) in K-space (cf. Fig. S2 and Woodcock, 1977). The
K-space distributions computed from the flatness and organ-
ization values for Cajas National Park are shown in Fig. 6, in
which much of the information present in the histogram
distributions is distilled graphically. In general, the lower tree
island sites (black symbols) have greater organization (reflect-
ing greater coherence in the surface-normal vector distribu-
tions) but lower flatness (higher roughness) values. In contrast,
the largest tree island sites (inverted triangles) are the least
organized, with significantly greater flatness values. Again, this
reflects the fact that the larger tree island sites include large
portions of the flatter down-slope landscape at the base of
high-relief topography. We also note that lower tree island sites
lie closer to the cluster-girdle threshold defined by K = 1,
whereas the higher tree island locations have distributions
more characteristic of clusters (see Fig. S2 for an explanation
of K-space).
DIS CUS S IO N
Numerous authors have expressed concern about the ongoing
degradation of upper montane tropical ecosystems (e.g.
Millones, 1982; Hess, 1990). Polylepis forests are clearlyamong South Americas most endangered ecosystems (Ro-
moleroux, 1994; Navarro et al., 2005); however, here we argue
that, in the case of Cajas National Park, most of the forest
removal probably occurred in the distant past. As a result of a
long history of several land-use practices, especially wood
gathering and grazing (Ellenberg, 1979; Lgaard, 1992), this
ecosystem is currently restricted to sites where burning is
strongly inhibited. Extensive field observations within our
study area suggest that burning rarely, if ever, occurs within
the boulder-strewn areas at the bases of cliffs, near road cuts,
Table 1 Summary of roughness and
organization statistics for the histogram
distributions shown in Fig. 5 and the results
of the KolmogorovSmirnov test. Elevation
range n
Average 1 standard
deviation
KolmogorovSmirnov test
(D-statistic, significance level
probability [%])*
Roughness Organization Roughness Organization
Random
35003700 m 98 0.33 0.88 1.77 0.96
37003900 m 160 0.29 0.85 1.23 0.79
39004100 m 223 0.30 0.97 1.08 0.61
< 1 ha
35003700 m 121 0.36 0.62 1.90 0.71 0.21, 98.82 0.22, 99.30
37003900 m 749 0.30 0.80 1.34 0.70 0.13, 98.64 0.18, >99.99
39004100 m 851 0.31 0.77 0.93 0.57 0.14, 99.98 0.15, >99.99
15 ha
35003700 m 978 0.41 0.77 2.26 0.86 0.36, >99.99 0.36, >99.99
37003900 m 1658 0.31 0.91 1.52 0.84 0.12, 98.11 0.18, >99.99
39004100 m 2704 0.30 0.98 1.12 0.62 0.09, 95.6 0.07, 76.1
510 ha
35003700 m 908 0.38 0.65 1.75 1.02 0.25, >99.99 0.11, 79.8
37003900 m 1016 0.43 0.64 2.06 0.55 0.55, >99.99 0.53, >99.99
39004100 m 1987 0.35 0.65 1.41 0.68 0.34, >99.99 0.26, >99.99
> 10 ha
35003700 m 950 0.41 0.54 2.00 0.94 0.37, >99.99 0.20, 99.82
37003900 m 668 0.22 0.74 0.80 0.27 0.51, >99.99 0.38, >99.99
39004100 m 1109 0.24 0.72 0.73 0.36 0.37, >99.99 0.32, >99.99
*The KolmogorovSmirnov test (KS-test) results for each elevation range and area subset are
based on a comparison with the random distribution. The KS-test parameters listed are first the
D-statistic (a measure of the difference between the two distributions) and second the confidence
probability that the data sets are drawn from the different distributions. For reference, identical
distributions would return values of 0.0 and 0% for D and the confidence probability,
respectively.
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or within very moist areas. Moreover, fires are relatively
unlikely to consume vegetation located above moist ravines or
depressions, where soil moisture levels are very high (Keating
et al., 2002).
Although the works cited above mention similar factors
concerning Polylepis distribution, this is, to our knowledge,
the first article to quantify the correlation between the
occurrence of Polylepis forests and terrain characteristics.
Clearly, additional studies involving extensive fieldwork will
be required to elucidate the fine-scale processes that govern
fire movement and the mortality of vegetation. Whereas
numerous models have been developed to simulate fire
behaviour in temperate forests (Keane et al., 2004), no such
modelling efforts, to our knowledge, have been applied to
high-elevation grasslands. The results of fire simulationefforts are likely to confirm the observations made in El
Cajas and elsewhere, but one should bear in mind that the
impacts of fire on vegetation vary with many factors,
including time of year, combinations of disturbances (espe-
cially grazing and burning), dominant species present, and
elevation (Verweij & Budde, 1992; Horn, 1998; Keating,
1998). One should also note that the interactions between
fires, grazing and other anthropogenic disturbances are very
complex and certainly merit further research (Renison et al.,
2006; Cierjacks et al., 2007).
The tropical Andes are well known for their species
diversity, endemism, and habitat diversity (Gentry, 1995;
Jrgensen & Leon-Yanez, 1999; Luteyn, 1999; Young et al.,
2002). Given both the high rates of land-cover conversion
and the growing interest in conservation issues, the impor-
tance of mapping and analysing tropical montane ecosystems
at the landscape scale is obvious. Most digital mapping
projects for such areas have focused on either Central
America or Mexico (e.g. Sader et al., 1989; Helmer et al.,
2000; Nagendra et al., 2004; Aguilar, 2005). A growing
number of mapping projects for the tropical Andes have
been performed at the regional to subcontinental scale (e.g.
Sierra et al., 1999), but thus far relatively few studies have
involved mapping tropical Andean vegetation at the land-
scape scale (e.g. Colby & Keating, 1998; Echavarria, 1998).
Even fewer have included GIS analyses or modelling to
explain how vegetation is influenced by disturbance or terrain
features (e.g. Keating, 1997; Vanacker et al., 2003; Kintz et al.,
2006). In part this trend stems from the scarcity of cloud-free
images for these regions, as well as from the difficulties
associated with performing the necessary fieldwork. For agrowing number of areas, however, digital data sets are
becoming available, and we recommend that upper montane
ecosystems be studied with geospatial techniques when
possible.
Here we have taken advantage of DEMs to evaluate
quantitatively the geomorphometric parameters of the land-
forms in Cajas National Park and to evaluate correlations
between landforms and the occurrence of tree islands. We
reiterate and readily acknowledge that it is an oversimplifica-
tion to claim that the distribution of tree islands is controlled
solely by the topographic character. Our working hypothesis,
however, is that certain topographic features inhibit the
movement of fire and therefore influence the current spatial
distribution of tree islands in the Cajas region. The close
proximity of the tree islands in Cajas National Park to areas of
high topographic roughness, high gradient and low organiza-
tion supports the notion that rough terrain encourages the
persistence of tree islands in this area. Forest fragments are also
highly threatened by burning in some lowland tropical
ecosystems (Gascon et al., 2000; Cochrane, 2001), but, even
in these environments, topographic features or rocky sub-
strates may afford some protection (e.g. Fordyce et al., 1997).
Our twin goals were to demonstrate the utility of
performing a quantitative topographic analysis and to evaluate
to what degree topography controls the tree island distribu-tion. Other influences are certainly important, but quantifica-
tion of their role is beyond the scope of this study. These
factors include grazing intensity, variance in precipitation,
bedrock geology of the region, hydrology, and other topo-
graphic parameters. We also acknowledge that micro-climatic
factors are very important in governing the position of
treelines. Recent work demonstrates unequivocally that the
relationships between microclimate and fire ecology are very
complex (Bader, 2007). Regardless of the relative importance
of other factors that we did not measure, this article
0
2
4
6
Flatnessln(S1/S2)
0 2 4 6
Organization ln(S2/S3)
Clusters
Girdles
Streng
th
Figure 6 The location in K-space [a plot of organization (ln(S2/
S3) vs. flatness (ln(S1/S2)] for the tree island locations in three
elevation ranges, 35003700 m (black), 37003900 m (grey),
39004100 m (white), and for areal extents of < 1 ha, 15 ha, 5
10 ha, and > 10 ha. Also shown are the locations for the random
sites in the study area. As discussed in Appendix S1, the variation
in the K-space location for the various subregions provides a way
to quantify the topographic differences among the various regions.
The n-values for the distributions are listed in Table 1.
D. Coblentz and P. L. Keating
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demonstrates that the position of forest patches in El Cajas is
indeed consistent with what is currently understood about fire
behaviour. These results should motivate a more detailed study
of the small-scale (< 10 m resolution) topography of individ-
ual tree islands as such data become available (most likely
requiring 1-m resolution LiDAR data).
CO NCLUS IO NS
In this article we have tested the hypothesis that the spatial
distribution of high-elevation tropical forest patches in Cajas
National Park is controlled by the landscape-scale topographic
character by evaluating the correlation between the tree island
distribution and a number of geomorphometric parameters
including organization, roughness and spectral power. Within
the assumptions and resolution limitations of our study, we
draw the following conclusions regarding the correlation
between features in the topography and the location of the
tree islands:
(1) A quantitative topographic analysis is a valuable tool for
understanding the distribution of land cover, particularly inregions where insular mountain ranges dominate the topo-
graphic distribution.
(2) Whereas climatic factors may have determined the original
distribution ofPolylepis-dominated forests, it appears that fires
exert a strong influence on the present distribution of these
forests. Our quantitative analyses demonstrate that areas at the
base of cliffs or above moist valleys are among the few places
where these ecosystems persist in south-western Ecuador.
(3) The geomorphometric analysis of the topography indicates
that the tree islands of Cajas National Park are found in close
proximity to rough topography with strong topographic
gradients. Overall, we find that the topographic roughness
exerts the primary control on the topographic character of the
landscape in the immediate area surrounding the tree islands.
(4) Climate, terrain, and disturbances all exert strong influ-
ences on the spatial position and areal extent of upper
montane tropical forests. The interrelationships among these
factors merit far more attention than they have previously been
accorded. As technology improves, scientists ability to relate
the position of plant communities to numerous variables will
increase further. In the future, extensive field data could be
coupled with model simulations to enhance the understanding
of tropical forest dynamics.
ACK NO W LE DGE M E NT S
Research for this article was funded by the Fulbright
Commission of Ecuador, the National Geographic Society,
Indiana University, and NSF Grant EAR-9422423 awarded to
B. Hansen, B. Leon, D. Rodbell, G. Seltzer, and K. R. Young.
The Instituto Geografico Militar in Quito provided topo-
graphic maps and aerial photographs, and LAND INFO
Worldwide Mapping, LLC created the DEM utilized in this
study. We are grateful to INEFAN for providing a permit to
conduct research within the park, to the herbarium staff at the
Catholic University in Quito for providing facilities and
logistical support, and to Stephanie Dickenson and Tom Evans
for providing statistical consulting. The program GMT (Wessel
& Smith, 1991) was used for a large part of the analysis as well
as for the figures in this paper.
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S UP P O RT ING INFO RM AT IO N
Additional Supporting Information may be found in the
online version of this article:
Appendix S1 Topographic analysis and the eigenvalue ratio
method.
Figure S1 Example surface-normal vectors illustrating topo-
graphic surface roughness.
Figure S2 Illustration ofK-space (topographic flatness plotted
vs. the topographic organization) and the location of several
example topographic distributions.
Please note: Wiley-Blackwell is not responsible for the content
or functionality of any supporting materials supplied by the
authors. Any queries (other than missing material) should be
directed to the corresponding author for the article.
B I O S K E T C H E S
David Coblentz is a solid Earth geophysicist with a joint
appointment at Los Alamos National Laboratory and the
University of New Mexico who, through speculative geody-
namics research, explores the common ground between
geodynamics and geoecology.
Phil Keating has spent 17 years investigating the influences
of anthropogenic disturbances on vegetation in the Ecuado-
rian Andes. He often combines field-based ecological studies
with remote sensing and GIS to determine how different fire
regimes modify floristic composition and landscape structure
in both paramo and timberline forest communities. DrKeating is currently a research associate at Indiana University.
Editor: Jorge Crisci
D. Coblentz and P. L. Keating
2038 Journal of Biogeography 35, 20262038 2008 The Authors. Journal compilation 2008 Blackwell Publishing Ltd