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To understand landslide regeneration and provide information necessary for restoration, we sampled seed rain, seed pool, and plant cover on two Ecuadorian landslides. We trapped 1304 seeds and found that, while most seeds were in the family Asteraceae, there was substantial variation in seed rain among plant families. Four hundred and seventy-five seedlings emerged from soil samples, including nonvascular and vascular families; again, species in Asteraceae dominated, with species in Piperaceae also very common. Plant cover, consisting of members of four fern families and 20 vascular plant families—with species in Asteraceae, Melastomataceae and Poaceae most common—was scored as a percentage of the total plot area. Principal components analysis (PCA) showed that, for all three of these plant life stages (seed rain, seed-propagule pool, plant cover), spatial variation was dominated by differences between the two landslides rather than within-landslide plot differences. PCA also showed that plots separated best on axes defined by the families Cecropiaceae, Urticaceae, Melastomataceae, Papilionaceae, Asteraceae, and Araceae with clumping of families in PCA space suggesting common successional strategies. Another multivariate technique, canonical correspondence analysis (CCA), showed that the combined seed rain and seed pool data could predict the percent cover of the family Verbenaceae and that the current plant cover families could predict Asteraceae seeds and seedlings. Finally, we use our past and present landslide data, along with multivariate modeling results, to suggest strategies for successful landslide restoration.
Content may be subject to copyright.
MARCH
1998
Restoration Ecology Vol. 6 No. 1, pp. 35–43
35
©
1998 Society for Ecological Restoration
Seed Inputs to
Microsite Patch
Recovery on Two
Tropandean Landslides
in Ecuador
Randall W. Myster
1
Fausto O. Sarmiento
2
Abstract
To understand landslide regeneration and provide in-
formation necessary for restoration, we sampled seed
rain, seed pool, and plant cover on two Ecuadorian
landslides. We trapped 1304 seeds and found that,
while most seeds were in the family Asteraceae, there
was substantial variation in seed rain among plant
families. Four hundred and seventy-five seedlings
emerged from soil samples, including nonvascular
and vascular families; again, species in Asteraceae
dominated, with species in Piperaceae also very com-
mon. Plant cover, consisting of members of four fern
families and 20 vascular plant families—with species
in Asteraceae, Melastomataceae and Poaceae most
common—was scored as a percentage of the total plot
area. Principal components analysis (PCA) showed
that, for all three of these plant life stages (seed rain,
seed-propagule pool, plant cover), spatial variation
was dominated by differences between the two land-
slides rather than within-landslide plot differences.
PCA also showed that plots separated best on axes de-
fined by the families Cecropiaceae, Urticaceae, Melas-
tomataceae, Papilionaceae, Asteraceae, and Araceae
with clumping of families in PCA space suggesting
common successional strategies. Another multivariate
technique, canonical correspondence analysis (CCA),
showed that the combined seed rain and seed pool
data could predict the percent cover of the family Ver-
benaceae and that the current plant cover families
could predict Asteraceae seeds and seedlings. Finally,
we use our past and present landslide data, along with
multivariate modeling results, to suggest strategies
for successful landslide restoration.
Introduction
L
andslides, caused by gravity but often aided by
torrential rains and earthquakes (Stern 1995), are
one of the most conspicuous landscape features of the
tropical Andes (Sarmiento 1994). Unfortunately, because
landslides are often associated with road construction
and human dwellings (Larsen & Torres-Sanchez 1992),
they can result in property damage and human death
(Sarmiento 1987). Therefore, the research we report in
this paper has direct human utility because it may lead
to intelligent management, restoration, and stabiliza-
tion of Neotropical landslides (Sarmiento 1992) by dis-
covering mechanisms that control natural vegetation
development.
Landslides are unique features of mountain land-
scapes. They (1) remove biomass and soil organic mat-
ter, exposing parent material (Garwood et al. 1979); (2)
produce extreme and complex environmental and bi-
otic spatial gradients (Sousa 1984; Myster & Fernández
1995); (3) contain regeneration sites for rare species
(e.g., various ferns; Walker 1994); (4) have recurrent lo-
calized disturbance—within-landslide resliding and edge
treefall (Hartshorn 1980)—that can slow the recovery
process; and (5) influence material flow, redistribution,
and nutrient cycling (Swanson et al. 1982; Guariguata
1990). In addition, because landslides of various ages
often consist of patches of colonizing vegetation on a
bare matrix (where light regime and mycorrhizal avail-
ability are key elements; Myster & Fernández 1995),
they form a dynamic patch system (Hupp 1983; Pickett
& White 1985).
Because successful land restoration hinges on infor-
mation about plant colonization of disturbed sites (Allen
1988; Dhillion et al. 1994) and because landslides re-
move intact vegetation, seed processes are critical to
landslide restoration and recovery (Flaccus 1959; Melick
& Ashton 1991; Myster 1997). Therefore we sampled,
during the months of July and August 1994, the seed
rain, seed pool, and current vegetation on two Ecuador-
ian landslides at the microhabitat patch scale and asked
these questions:
(1) How do species, families, and numbers of dispersed
seeds of emerged seedlings from the seed pool and
of the vegetation currently established (i.e., percent
cover) vary between two landslides and among mi-
crohabitats within them?
1
Institute for Tropical Ecosystem Studies, University of Puerto
Rico, P. O. Box 363682, San Juan, PR 09036 U.S.A.
2
Institute of Ecology, The University of Georgia, Athens, GA
30602–2022 U.S.A.
Landslide Restoration in Ecuador
36
Restoration Ecology
MARCH
1998
(2) Are there correlations among the three plant life
stages, seeds, seedlings, and current vegetation?
Can seeds or seedlings from a plot be used to pre-
dict its current vegetation? If so, can we guide resto-
ration efforts by modeling these processes?
Methods
Study Site
The study site was the Maquipucuna Reserve (0
8
05
9
N,
78
8
37
9
W) in northwestern Ecuador near the town of
Nanegal. The site occupies the Andean piedmont at the
southern limit of the Chocó biotic province, one of My-
ers’ hot spots for biodiversity (Sarmiento 1995
c
) and a
typical location of a Tropandean landscape (Sarmiento
1994, 1995
a
). The reserve stretches from 1150 m above
sea level at the Umachaca River bridge to 1970 m above
sea level at the summit of Mount Montecristi; hence it
includes the lower montane, the transitional, and the
upper montane forest communities in the Upper Guayl-
labamba River basin. The reserve was established in
1987 to protect these representative areas of the land-
scape in the Tropandean region (Sarmiento 1995
b
). This
basin is the site of several development-aid projects:
Sustainable Agriculture and Natural Resources Man-
agement (SANREM), Sustainable Use of Biological Re-
sources (SUBIR), and Integrated Rural Development of
the Northwestern Pichincha Province (UDRI). Old-growth
lower montane wet forest is restricted to the upper
reaches of the Maquipucuna range, whereas secondary
growth and abandoned fields and pastures are located
nearer the rivers (Sarmiento 1995
a
). The soil developed
from volcanic ash deposits. It is high in organic matter,
medium-textured, and moderately fertile.
The Maquipucuna Reserve has dominant overstory
vegetation in the families Lauraceae, Rubiaceae, Mora-
ceae, Myristicaceae, Araliaceae, Meliaceae, and Myrtaceae.
The understory vegetation includes vascular plants in
the families Euphorbiaceae, Solanaceae, Acanthaceae,
Piperaceae, Melastomataceae, and Asteraceae and tree
ferns with a forb layer consisting of epiphytes, orchids,
“huaicundos” (Bromeliaceae:
Guzmania
spp.), and thick
layers of moss (Sarmiento 1995
a
). The two landslide
study sites are located in the Rio Tulambi watershed
(Fig. 1) and are both 10 years old. Landslide 1 (1475 m
elevation; 0
8
06
9
46
99
N, 78
8
37
9
53
99
W) is 850 m
2
(0.085 ha)
in area and west-facing, with a 45% slope surrounded
by old-growth forest. Landslide 2 (1795 m elevation;
0
8
06
9
38
99
N, 78
8
37
9
61
99
W) is 2100 m
2
(0.21 ha) in area and
west-southwest facing, with a 70% slope surrounded by
old-growth forest at the headwaters of a steep brook.
Sampling Design
We established 12 standarized 1-m
2
sample plots on or
next to each of the two landslides on areas that were a
mixture of bare soil and vegetation. These plots were on
three parallel transects: transect 1 at 10 m from the top of
the landslide, transect 2 at 20 m, and transect 3 at 30 m.
Along each transect, we placed one 1-m
2
plot in each of
four microhabitats: the forest next to the landslide (F) 10
m from the landslide, the forest/landslide border just
outside the lip (FB), the landslide/forest border just in-
side the lip (LB), and the center of the landslide (C). The
center microsite was 10 m from the forest edge for land-
slide 1 and 25 m from the forest edge for landslide 2.
First, at each microsite we placed seed traps of 1-m
2
cloth staked parallel to and 1 m above the ground, cov-
ered with a piece of hardware cloth. Each stake was
covered with tangelfoot, a sticky resin designed to ex-
clude crawling insect seed predators (Myster 1997).
These were in place for 24 days in July and August
1994, after which we collected the seeds. Next, we took
small soil samples (100 g of top soil taken by insertion
of a 10-cm-diameter jar at a depth of 2 cm; Myster 1993)
from each microsite, incubated the samples in pots at
the site for 1 year, and recorded the microsite and spe-
cies of each seedling that emerged. Finally, in August
1994, we sampled the plant cover of each plot by visu-
ally estimating the percentage of the area of the plot at
each microsite covered by each plant species. Because
the vegetation is layered (Myster & Fernández 1995), in
a few instances the plant cover exceeded 100% either for
individual species or in total. In those cases the values
were not normalized to total cover in order to reflect the
Figure 1. Map of a section of the Maquipucuna Reserve show-
ing forest, pasture, oldfields previously in agriculture, and hu-
man dwellings (clear triangles). The study landslides are indi-
cated as landslide 1 (lower elevation) and landslide 2 (higher
elevation).
Landslide Restoration in Ecuador
MARCH
1998
Restoration Ecology
37
known vegetative structure. Nomenclature follows that
of Gentry (1993), Webster (1993), and Sarmiento (1995
c
).
Statistical Analysis
Principle components analysis (PCA; SAS Institute, Inc.
1985) and canonical correspondence analysis (CCA;
SAS Institute, Inc. 1995) were performed on the data set
as statistical tests of relationships among the plots, spe-
cies, and life stages needed to maximize restoration suc-
cess. These methods are preferable to simple similarity
indices (Pielou 1984) that were not calculated. Because
our focus is on individual species and how to restore
them, community parameters such as richness or total
cover were also not calculated. Due to the small number
of species found in all three samplings (seed rain, seed
pool, vegetation data), we grouped abundances at the
family level for the multivariate analyses. Six families
(Asteraceae, Melastomataceae, Verbenaceae, Pappaver-
aceae, Solanaceae, Poaceae) were represented in all
three life-stage categories, and an additional four fami-
lies (Urticaceae, Bombacaceae, Cecropiaceae, Passiflora-
ceae) were represented in the percent cover data as
seeds or seedlings. Therefore, CCA used a matrix with
the 24 plots as rows (12 microsites for each of the two
landslides) and these 10 most common plant families—
once for the pooled seed rain and seed pool data and
once for the vegetation cover—defining the columns (10
families
3
2 [seed rain, seed-propagule pool]
5
20 total
columns). The PCA used a matrix with the same rows
as CCA but with the 24 most common plant families
(those with greater than 9% total cover) as the columns.
PCA was performed on the plant cover data alone in or-
der to minimize the number of matrix zeros that can
lead to invalid results (Pielou 1984; SAS Institute Inc.
1985). Both an R-type and Q-type PCA (after matrix
transposition; Pielou 1984) were performed.
Results
The complete sampled data set is given in Table 1. For
many of the species, this is the first ecological informa-
tion ever published. The text focuses on key species, pre-
sented in the same order to ease comparison and to
identify families of species in Table 1. Lacking evidence
to the contrary, we assume that the importance of spe-
cies for landslide restoration is directly proportional to
their abundance. Hence, we give spatial details for the
10 most abundant species in each of the three life stages
by dividing the total number of seeds, seedlings, or
cover values into groups based on the location of indi-
vidual plots (landslide versus transect versus micro-
site). In each case, the ten species exceeded 90% of the
total sample.
Seed Rain
A total of 1304 seeds from 25 families was collected in
the seed traps (Table 1). Most seed came from species in
the family Asteraceae (
Vernonia patens
,
Andropogon
spp.,
Hieracium
spp.,
Baccharis latifolia
), where between-land-
slide differences (landslide 1 had much more seed than
landslide 2) dominated the within-landslide variation
(transects versus microsites). For
Cecropia monostachya
(Cecropiaceae), landslide 2 seed rain dominated over
that of landslide 1.
Vaccinium
spp. (Ericaceae) seeds
were distributed evenly over all microsites.
Miconia
spp. (Melastomataceae) seeds were found predomi-
nantly on the landslide border microsites of the second
and third transects.
Digitaria sanguinalis
(Poaceae) seeds
were found mainly in landslide 1. Finally, seeds of
Boconia
frutescens
(Papaveraceae),
Solanum
spp. (Solanceae), and
Prestonia
spp. (Apocynaceae) were found mainly in the
forest microsite on the first transect.
Seed-Propagule Pool
There were 23 lichen, 148 moss, 2
Sellaginella diffusa
, 15
fern individuals (all mainly from landslide 2), and 287
seedlings from the 38 vascular families that emerged
from the soil samples (Table 1). Again, species in the
family Asteraceae dominated, and between-landslide
differences dominated over transect and microsite dif-
ferences. Those Asteraceae species were
Elephantopus
mollis
, whose seeds fell mainly on landslide 2;
Vernonia
patens
and
Pseudelephantopus spiralis
, whose seeds fell
mainly on landslide 1; and
Baccharis latifolia
, whose
seeds fell mainly on the center and landslide border mi-
crosites in both landslides. The other six most common
species showed most dominance between landslides as
well:
Piper aduncum
(Piperaceae) seedlings on landslide
2,
Althernantera
spp. (Amaranthaceae) on landslide 1,
Digitaria sanguinalis
(Poaceae) on landslide 1,
Verbena
litoralis
(Verbenaceae) on landslide 2, and both
Asple-
nium
spp. (Aspleniaceae) and
Desmodium canum
(Fa-
baceae) on landslide 1.
Plant Cover
Liverworts covered 2% of only one plot, and moss cov-
ered 170% over a number of plots (mainly on landslide
2). We found for plant cover, as for seeds and seedlings,
that (1) the greatest total cover came from a species in
the Asteraceae (
Vernonia patens
); and (2) for Asteraceae,
between-landslide variation (mainly on landslide 1 in
this case) dominated over within-landslide variation.
Dominance of between-landslide variation was also
found for both
Urera baccifera
(Urticaceae) and
Miconia
spp. (Melastomataceae), which were found only on
Landslide Restoration in Ecuador
38
Restoration Ecology
MARCH
1998
Table 1.
Number of seeds and emerged seedlings from the soil samples and plant cover of all sampled species.*
Species Common name Lifeform Seeds Seedlings Cover (%)
Vernonia patens
pinchulán T 683 15 225
Bidens
sp. amorseco FO 8
Andropogon
sp. sigse G 67
Hieracium
sp. hierba de sal FO 151
Baccharis latifolia
chilca S 50 9
Cecropia monostachya
guarumo de monte T 61 123
Vaccinium
sp. sacha mortiño S 49
Miconia
sp. colca S 45 3 261
Digitaria sanguinalis
chirimbilla G 32 18
Setaria
sp. pasto miel G 2
Boconia frutescens
chandor S 26 1 1
Solanum
sp. tomatillo S 26 2 93
Prestonia
sp. laca FO 26
Plantago major
lengua de suegra FO 22
Asclepias
sp. palomitas FO 21
Verbena
sp. verbena FO 16
Passiflora
sp. sacha taxo FO 14 40
Pilea
sp. llausa S 13
Clematis
sp. savaleta FO 13
Ipomoea
sp. sacha papa FO 12
Rubus
sp. mora FO 12
Galium
sp. cebollín FO 7
Gonzalagunia
sp. aretes de deslaves T 2
Amaranthus
sp. ataco FO 9
Oxalis
sp. lechero rastrero FO 8
Monnina
sp. ushmayuyo S 8
Cyperus
sp. hierba de perro FO 7 1
Muehlenbeckia
sp. cacaotillo belludo FO 3
Ochroma pyramidale
balza T 3 — 12
Desmodium
sp. uña de gato FO 1 7
Nectandra
sp. pagche T 1
Elephantopus mollis
orejuela FO — 65
Pseudelephantopus spiralis
verdeo FO — 6
Mickania cordifolia
pisco lulo S 4
Ageratina pichinchensis
rastrera FO — 2
Ageratum
sp. hierba de sapo FO 1
Bidens pilosa
amorseco FO — 1
Sabicea urticifolia
ortiguillo S — 1
Piper aduncum
cordoncillo S — 63 22
Piper pytolaccaefolium
matico S — 1
Althernantera
sp. futula S 33
Sporobolus poiretii hierba mansa G 3
Paspalum conjugatum paja G — 1
Verbena litoralis verbena FO — 18 65
Solanum cayannensis tomatillo S — 5
Asplenium sp. F 6
Desmodium canum una de gato FO 6
Cuphea cartagenensis llausa S — 4
Hydrocotyle leucocephala orejitas FO — 4
Cyperus hemafroditus hierba de perro FO 2
Phyllanthus miruri sacha oliva S 2
Tripogandra serrulata hierba dentada FO 2
Anagallis minima chiquita FO — 1
Begonia parviflora siguemesiguama S 1
Justicia amata infiel S — 1
Oxypetalum coreifolium hierba mala FO 1
Psidium guajaba guayava T — 1
Sida rhombifolia escobilla S — 1
Stachytarpheta cayannensis hierba mora FO 1
Asplenium sp. F 200
Nephrolepis sp. suro guaga F 6
Landslide Restoration in Ecuador
MARCH 1998 Restoration Ecology 39
landslide 2. In contrast, Asplenium spp. (Aspleniaceae)
were most common on center microsites. Trichipterix pi-
lossisima (Cyatheaceae) was found mainly on the first
transect in landslide 1, and Cyathea spp. (also in
Cyatheaceae) was found only on landslide 2 in the third
transect. Finally, Chusquea spp. (Poaceae) was found
only on landslide 1, Anthurium ssp. (Araceae) was found
in equal abundance on both landslides, and landslide
2 showed the most cover of both Philodendrom spp.
(Araceae) and Cecropia monostachya (Cecropiaceae).
Statistical Analysis
The canonical correspondence analysis found moder-
ate correlations between the seeds of Solanaceae and
Table 1. Continued.
Species Common name Lifeform Seeds Seedlings Cover (%)
Trichipterix pilosissima helecho arboreo F 103
Cyathea sp. helecho arboreo F 115
Diplazium lindbergii dos caras F 75
Polybotria sp. milojas F 36
Baccharis sp. chilca S — — 5
Gurania macrophyla zambo do zanja FO 22
Pilea apurascensis ortiguillo liso S 51
Boehmeria sp. campanilla FO 57
Urera baccifera ortiguillo S — — 120
Blakea sp. FO 55
Erythrina megistophyla porotón T — — 60
Swartzia sp. T 70
Begonia glabra siguemesigueme S — — 13
Chusquea sp. suro G — — 265
Neurolepis sp. G 96
Agrostis sp. hierba G 5
Anthurium sp. pucsia FO 113
Pepperomia sp. S 62
Acalypha pladichephalus pigua FO — 76
Bomarea sp. FO 25
Philodendrom sp. cartucho FO 180
Xanthosoma sagittifolia toa FO — 81
Cyphomandra hartwegii tomate de arbol S 12
Witheringia sp. S 25
Hyptis sp. FO 51
Heliconia sp. plantanillo FO 71
Costus sp. sacha caña S 80
Rumex sp. lengua de vaca FO 1
Anthurium sp. FO 2
Oxalis microcarpa FO — 2
Phytolacca rivinoides FO — 4
Pothomorphe peltata S—— 5
Xanthosoma sagittifolium FO — 1
Cyphomandra hartwegii T—— 2
Cordia sp. T 1
Cyclanthera sp. FO 1
Epidendrum sp. FO — 1
Nephrolepis sp. F — — 1
Guzmania sp. huaicundo FO — 5
Masdevalia sp. orquídea FO — 1
Beilschmiedia sp. FO 1
Caladium sp. FO — 1
Hedyosum sp. motilón T — — 5
Scheflera sp. chiflera T 2
*Species are presented in the same order as in the text. Spatial variation of seeds, seedlings, and cover is presented in the text, as are the
family names. Lifeforms are fern (F); tree (T); shrub (S); forb (FO); and graminoid (G). Species present with less than 5% of total cover
were in the families Polygonaceae (Rumex sp.), Papaveraceae (Boconia frutescens), Araceae (Anthurium sp.), Oxalidaceae (Oxalis micro-
carpa), Phytolaccaceae (Phytolacca rivinoides), Piperaceae (Pothomorphe peltata), Araceae (Xanthosoma sagittifolium), Solanaceae (Cyphoman-
dra hartwegii), Boraginaceae (Cordia sp.), Cucurbitaceae (Cyclanthera sp.), Orchidaceae (Epidendrum sp.), Davalliaceae (Nephrolepis sp.),
Bromeliaceae (Guzmania sp.), Orchidaceae (Masdevalia sp.), Lauraceae (Beilschmiedia sp.), Araceae (Caladium sp.), Chloranthaceae (Hedyo-
sum sp.), and Araliaceae (Scheflera sp.).
Landslide Restoration in Ecuador
40 Restoration Ecology MARCH 1998
Passifloraceae (10.6338), between the cover of Melasto-
mataceae and Cecropiaceae (10.6659), between the seeds
of Urticaceae and the cover of Asteraceae (10.6426), and
between the seeds of Asteraceae and the cover of Ver-
benaceae (10.6251). In addition, the standardized ca-
nonical coefficients (SCC) showed that the seeds grouped
by families were best described by the cover of Aster-
aceae (10.8159) and by the absence of cover of Passiflo-
raceae (20.5045), and that the percent cover by families
was best described by the seeds of Verbenaceae (10.9746).
There was some evidence, however, that Passifloraceae
was a suppressor enhancing the SCC relationships (SAS
Institute, Inc. 1985). Finally, the canonical redundancy
analysis of the CCA showed that neither the seed fami-
lies nor the cover families were good predictors of the
other, with only a 0.1015 cumulative proportion of the
variance explained. But the percent cover of all the fam-
ilies could predict the seed input of Asteraceae (10.4596),
and the seed input of all the families could predict the
cover of Verbenaceae (10.8430).
Principle components analysis on the plant cover
data showed that the first three axes defined a majority
(61%) of the variance, suggesting that the current vege-
tation defined relationships among the plots. This was a
Q-type ordination (Pielou 1984), with plots defined in
family space and the resulting PCA axes defined by
these significant correlations with plant families: PCA
axis I had positive correlations with Melastomataceae,
Papilionaceae, Araceae, and Cecropiaceae and negative
correlations with Urticaceae, Asteraceae, and Poaceae;
PCA axis II had positive correlations with Urticaceae,
Melastomataceae, Poaceae, and Araceae and a negative
correlation with the fern family Aspleniaceae; and PCA
axis III had positive correlations with Bombacaceae,
Asteraceae, and the fern family Cyatheaceae.
Of the three alternative ways to label the plots using
the Q-type PCA ordination axes loadings with percent
cover data, labeling by landslide (Fig. 2) shows plot dif-
ferences (supporting the visual data inspection related
earlier in the results) better than labeling by transect
(Fig. 3) or microsite (Fig. 4; although there was some
clumping shown here). Further, landslide 1 tended to
have negative values on PCA axis I, defined by abun-
dances of species in the families Urticaceae, Aster-
aceae, and Poaceae; and landslide II tended to have
positive values on PCA axis I, defined by abundances
of species in the families Melastomataceae, Papilion-
aceae, Araceae, and Cecropiaceae. We also performed
an R-type PCA ordination (Pielou 1984) on the same
data in which the 24 plant families were separated in
plot space. It showed a distinct clumping of the families
Cucurbitaceae, Bombacaceae, Davalliaceae, Begoniaceae,
Amaryllidaceae, and Solanaceae, with Poaceae, Melas-
tomaceae, and Labiatae defining the extremes of the
R-type plot space.
Discussion
Because intact vegetation is lost after severe distur-
bance, the correspondence among these three life stages
and the possible effect of seed processes (such as seed
Figure 2. Principal components analysis scatterplot of the 24
plots in plant family space using percent cover labeled by
landslide number: 1 versus 2.
Figure 3. Same as Figure 2 but with labeling by transect: 1
versus 2 versus 3.
Landslide Restoration in Ecuador
MARCH 1998 Restoration Ecology 41
rain and seed pool) on the standing vegetation should
be of critical importance in the recovery and restoration
of landslides (Myster & Fernández 1995). We found that
Asteraceae (Vernonia patens and Baccharis latifolia) was
the most common family in both the seed rain and the
seed pool and one of the most common families of the
current vegetation. In addition, Melastomataceae (Mico-
nia spp.) and Verbenaceae (Verbena spp.) were well rep-
resented in all three plant stages, and Papaveraceae
(Bocconia frutescens) was just present in all three. It was
difficult, however, to find the same species or family
from all three life stages in the same plot (also see
Tsuyuzaki & Kanda 1996). Vernonia patens was the only
species detected in all three, and some species in the
families Solanaceae and Poaceae were present in seed
rain, seed pool, and plant cover but in different plots.
Principal components analysis suggested that the
plots were best grouped by landslide and that, in gen-
eral, the families—whether seeds, seedlings, or plant
cover—that differed mainly by landslide rather than by
transect or microsite were also the most abundant. For
example, Asteraceae, with the most seeds, seedlings,
and plant cover of any family, differed mainly by land-
slide in each of the three life stages. Although there was
agreement between the raw data and the PCA analysis,
the individual species did not always match, and some
relative abundances shifted from one landslide to an-
other depending on life stage. Finally, the R-type PCA
analysis showed that species in certain families tended
to occur in the same plots and that those species could
have similar life-history or successional roles (Gitay et
al. 1992).
There were interesting comparisons with other Neo-
tropical landslide studies. Our numbers of dispersed
seeds (on average 2.25 seeds/m2/day) were greater
than those of landslide studies conducted in Puerto
Rico (0.24 seeds/m2/day; Walker & Neris 1993; 31
seeds/m2 over 2 months; Myster & Fernández 1995)
and Costa Rica (0.64 seeds/m2/day; Myster 1993), per-
haps due to the more diverse surrounding rain forest in
Ecuador. But this study agreed with those in Puerto
Rico (Walker & Neris 1993; Myster & Fernández 1995)
and Costa Rica (Myster 1993) in that the differences be-
tween landslides were greater than the differences in lo-
cations within a landslide for seeds, for saplings, and
for percent cover. The only seed genus that achieved
any substantial landslide seed load in this Ecuadorean
study, in Costa Rica (Myster 1993), and in Puerto Rico
(Walker & Neris 1993) was Cecropia, but Gonzalagunia
seeds were also present at all three locations.
We suggest that the large moss cover on these land-
slides can enhance germination both by compensating
for the loss of water after a landslide and by facilitating
scarification of seed coats. This may help to explain
why the number of seedlings that emerged from our
Figure 4. Same as Figure 2 but with labeling by microhabitat:
forest (F) versus forest border (FB) versus landslide border
(LB) versus center (C).
Table 2. A comparison between common tree species
sampled as plant cover on Puerto Rican, Costa Rican, and
Ecuadoran landslides.*
Puerto Rico Costa Rica Ecuador
Alchornea latifolia A. latifolia
Cecropia schreberiana C. polyphlebia C. monostachya
Cyathea arborea C. caracasana C. brunnescens
Gleichenia bifida G. bifida
Gonzalagunia rosea G. spicata G. dependens
Miconia mirabilis M. tonduzii M. aeruginosa
Miconia racemosa
Miconia prasina
Miconia tetrandra
Miconia impetiolaris
Ocotea sintenisii O. pittieri O. sp.
Ocotea leucoxylon
Ocotea spathulata
Palicourea riparia P. lasiorrhachis
P. standleyana
Piper aduncum P. aduncum
Piper hispidum P. lanceaefolium P. pytolaccaefolium
Piper glabrescens
Piper amalago
Psychotria brachiata P. elata
Urera baccifera U. caracasana U. baccifera
*Puerto Rican landslides: more than 100 plots 2 3 5 m sampled 10 times over
a 5-year period (Myster & Walker 1997); Costa Rican landslides: 18 plots 3 3 5
m sampled once (Myster 1993); Ecuadorian landslides: 24 1 3 1 m plots sam-
pled once (this study). All landslides were 5–10 years old and surrounded by
lower montane wet forest. Life forms are trees (Alchornea spp., Cecropia spp.,
Gonzalagunia spp., Ocotea spp.), shrubs (Miconia spp., Palicourea spp., Piper
spp., Psychotria spp., Urera spp.), or ferns (Cyathea spp., Gleichenia spp.).
Landslide Restoration in Ecuador
42 Restoration Ecology MARCH 1998
soil samples—12 per 100-g sample—after one year was
larger than that from landslide soil samples of Guari-
guata (1990) and Myster and Fernández (1995), both of
which had 2 per 100-g sample in Pureto Rico, and of
Myster (1993), which had 5 per 100-g sample in Costa
Rica even when standardized for time period. But the
Costa Rican study (Myster 1993) did have many seed-
lings from Asteraceae and Melastomaceae in common
with this study.
Taxa of the extant vegetation (i.e., plant cover) were
shared with these other landslide studies much more
often than the sampled seeds and seedlings. For exam-
ple, taxa such as Cecropia (Cecropiaceae), Urera (Urtice-
aeae), Witheringa (Solanaceae), and Chusquea (Poaceae)
were also common in the Costa Rican study (Myster
1993), with the first three families and genera also com-
mon in Puerto Rico (Myster & Fernández 1995). Table 2
compares the plant cover of species in landslides from
Puerto Rico and Costa Rica (Myster 1993) and shows a
close similarity in plant genera, between the Puerto
Rican and Costa Rican (Lugo 1987; Myster 1997) land-
slide sites, with two of the same species (Alchornea latifo-
lia, Gleichenia bifida) found at both sites. This relation-
ship is weaker on the site in Ecuador, however, with
few genera or families in common with either Puerto
Rican or Costa Rican sites. For example, Chusquea spp.
(Bamboo; Tsuyuzaki & Kanda 1996) is very common in
Ecuadorian landslides but is not found in either Costa
Rica or Puerto Rico.
Although the rehabilitation of montane tropical rain
forests is important for attaining sustainability, preserv-
ing the rich biodiversity in the tropics, and combating
the global effects of deforestation, such as greenhouse
gas emissions and climate change (Brown & Lugo
1994), landslide restoration may also save lives and
property. We have now studied landslide succession at
three different Neotropical sites, each surrounded by
lower montane wet forest and located so as to capture
much of the natural variation found in the Neotropics.
For example, Ecuador is a mainland site at 08 latitude,
Costa Rica (Myster 1993) is a mainland site at 108 N lati-
tude, and Puerto Rico (Myster & Fernández 1995; Myster
1997) is an island site at 188 N latitude. Based on these
studies we suggest that landslide restoration should fo-
cus on (1) addition of seeds and seedlings that match
those microsites where the species occur naturally (for
example, through use of the spatial detail given here);
(2) addition of both nutrients and vesicular-arbuscular
mycorrhizal spores using forest ground-level soil; (3)
addition of soil in areas where unweathered substrate is
exposed; (4) layering of the exposed surface with moss
to provide germination sites for seeds; (5) planting of
trees as recruitment foci or nurse trees (McDonnell &
Stiles 1983; Myster & Pickett 1992); and (6) proactive re-
moval of exotic species (Sarmiento in press).
Although canonical components analysis showed
that whole plant life-stage groups did not predict one
another well—seed load in a given plot did not do a
good job predicting the plant cover in that plot—it did
identify several key plant families that could be used
both for successful landslide restoration efforts and fu-
ture modeling. These families had predictive power to-
ward (1) whole life stages (e.g., seeds in the family Ver-
benaceae could predict plant cover); (2) families within
the same life stage (e.g., seed load in the family Solan-
aceae could predict the seed load in the family Passiflo-
raceae); and (3) families in other life stages (e.g., seed load
in the family Urticaceae could predict the plant cover in
the family Asteraceae). We suggest that predictive mod-
els such as canonical correspondence analysis that relate
seed and seedling inputs and success to the current vege-
tation in a plot-wise or microsite-specific manner will
lead to the most successful restoration efforts.
Acknowledgments
We thank the staff of the Maquipucuna Reserve, espe-
cially manger B. Castro and field assistant L. Pozo, for
help with this research. We also thank E. Freire and T.
Muñoz of the National Herbarium of Ecuador and A.
Mariscal and C. Cerón from Universidad Central del Ec-
uador for their help with species identification. Finally,
we thank D. Gorchov, D. Schaefer, and J. Thomlinson for
helpful comments on previous versions of the manu-
script. This research was performed under National Sci-
ence Foundation grant BSR-8811902 to the Institute for
Tropical Ecosystem Studies, University of Puerto Rico,
and the International Institute of Tropical Forestry as
part of the Long-term Ecological Research Program in
the Luquillo Experimental Forest. Additional support
was provided by the U.S. Forest Service and the Univer-
sity of Puerto Rico. F.O.S. was supported by a grant from
the MacArthur Foundation to the Institute of Ecology,
University of Georgia, as part of the Below-ground Ecol-
ogy Project. Additional support was provided by the
Scott Neotropic Fund of the Lincoln Park Zoological So-
ciety.
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... Gradient terminology contains not just systemic variation, potentials/dissipation, and directional issues but also characteristics such as steepness, steadiness, dynamics (Whittaker 1975), and even fractals (Mandelbrot 1983) if the gradients reoccur hierarchically at larger and larger spatial, temporal, or spatiotemporal scales, but terminology is lacking to describe interaction among abiotic and biotic factors located at different coordinates on the same gradient or on different gradients, which may include mechanistic influence, i.e., gradients may "overlap" in space and time and affect each other. Examples of this in mountain ecosystems include (1) the spatial gradient of increasing tree seedling abundance from an old-field into the surrounding mountain forest ecosystem overlapping with multiple temporal gradients of the same tree seedling increase at various places within the old-field (Myster 2008), and (2) multiple temporal gradients of recovering landslides overlapping across the spatial elevational gradients in tropical mountains (Myster and Sarimento 1998). These examples show how landslides and conversion of forest to agriculture in mountain ecosystems can "interrupt" spatial gradients. ...
Book
This book is the second volume in a series on montology dedicated to the transdisciplinary reflection of mountain research, considering the diversity of views on mountains and their problemata in the context of rapid technological development and unprecedented accumulation and dissemination of information around the world. The necessity for a new orderly and structured lexicon arose from the need to critically reassess the colonial past in the development of mountain territories, the development of a new and alternative understanding of mountain topics in the light of decolonized epistemology. The creation of coordinated and ordered terms for the main parts of mountain research creates the basis for an unorthodox understanding of the ontology of mountains and helps to better understand the complex cultural and natural essence of mountain socio-ecological systems. At the same time, a local episteme of mountains, considering local values, small scales, and vernacular visions are of particular importance, which must be taken into account in the current terminology. The purpose of the book is to provide methodological support for montology as a convergent and transdisciplinary science of mountains, based on the harmonization of its terminological base. The book pays special attention to onomastics, toponymy, standardization and other nuances of terms used in mountain research. According to this goal, three dozen articles in a relatively small format (about 3 pages) vividly, attractively and innovatively reflect the modern view of one or more related terms. Articles include definition(s) of the term, description of etymology, onomastics or toponymy used, examples of local characteristics compared to traditional sources, possible vernacular terms. Articles are grouped into four main areas: 1) Basic glossary of montology terminology, 2) Towards mountain socio-ecological systems, 3) Innovative disciplinary systemic realm, 4) Mountain classifications, onomastics, critical toponomy and rediscovery of meaning. The authors of the articles are leading experts in the field of mountain research from around the world. The book is intended for scientists, experts and teachers. It is provided with an annotated list of the most important montology terms. Keywords: interdisciplinary studies of mountain areas toponymy Montology decolonized epistemology socioecological systems onomastics
... One of the most significant socioecological gradients of the EDGs is the differential resource use preference that increases in the mid-elevation mountains as a result of terracing, building of irrigation channels, and other transportation networks. In fact, most of the successional dynamics of the cloud forest is a direct consequence of road construction and talud breakage, hence providing for landslides (e.g., Myster and Sarmiento 1998), rockslides, or the feared "waiku"or a destructive mudflow that washes downstream-bound catastrophes. Huge floods of the piedmont are directly related to the changes in forest cover and resource use of the headwaters. ...
Chapter
There is consensus to advance science with unorthodox narratives generated with new discoveries, different perspectives, or challenging innovation altogether. However, it is also consensual that these mountain narratives, like the waves in fluid water or air, move along the time scales with different dynamics and distinctive rhythms, generating a symphony of knowledge, which can only be integrated with the crosscutting ability of montology as a convergent science (Sarmiento 2020). Indeed, applied montology is the appropriate avenue for developing an environmental awareness of the whole mountainscape. With the wise trend of consilience (Wilson 1998) and the untested hype of noetic science (Nickell 2010), we contribute this chapter with the objective of increasing our epistemology of mountains, including them as socioecological landscapes and not as mere ecosystems.
... Post-landslide habitat degradation poses several chronic stresses for plant colonisation, such as excessive solar radiation and soil erosion associated with vegetation removal and soil nutrients loss due to topsoil removal (Wilcke et al. 2003, Lin et al. 2006, Walker & Shiels 2013. The longterm colonisation of persistent pioneer plants can further retard the establishment of late-successional species and, in turn, arrest vegetation succession (Guariguata 1990, Myster & Sarmiento 1998, Royo & Carson 2006. Through intercepting strong sunlight, providing materials for soil organic matter enrichment, and suppressing the growth of persistent pioneer plants, large trees in or near landslide scarps can exert positive effects on the growth and survival of recruited and existing small trees and ultimately promote vegetation recovery (Walker & Shiels 2013, Chen et al. 2014. ...
... One of the most significant socioecological gradients of the EDGs is the differential resource use preference that increases in the mid-elevation mountains as a result of terracing, building of irrigation channels, and other transportation networks. In fact, most of the successional dynamics of the cloud forest is a direct consequence of road construction and talud breakage, hence providing for landslides (e.g., Myster and Sarmiento 1998), rockslides, or the feared "waiku"or a destructive mudflow that washes downstream-bound catastrophes. Huge floods of the piedmont are directly related to the changes in forest cover and resource use of the headwaters. ...
Book
The importance of the Neotropics to the world's climate, biogeochemical cycling and biodiversity cannot be questioned. This book suggests that gradients are key to understanding both these issues and Neotropical ecosystem structure, function and dynamics in general. Those gradients are either spatial, temporal or spatio-temporal, where many temporal and spatio-temporal gradients are initiated by disturbances (e.g., tree-fall, landslide, cultivation). And in particular for the Neotropics, three large spatial gradients - latitude, longitude, altitude (elevation) - are of critical importance. The editor has over 30 years of experience investigating Neotropical gradients in Costa Rica, Puerto Rico, Peru and Ecuador, and has published 5 previous books on different aspects of the Neotropics. Once again he has assembled top-shelf Neotropical scientists and researchers, here to focus on gradients: their nature, interactions and how they structure ecosystems.
... One of the most significant socioecological gradients of the EDGs is the differential resource use preference that increase in the mid-elevation mountains as a result of terracing, building of irrigation channels, and other transportation networks. In fact, most of the successional dynamics of the cloud forest is a direct consequence of road construction and talud breakage, hence providing for landslides (e.g., Myster and Sarmiento 1998), rockslides, or the feared "waiku"or destructive mudflow that washes downstream-bound catastrophes. Huge floods of the piedmont are directly related to the changes in forest cover and resource use of the headwaters. ...
Chapter
Full-text available
There is consensus to advance science with unorthodox narratives generated with new discoveries, different perspectives, or challenging innovation altogether. However, it is also consensual that these mountain narratives, like the waves in fluid water or air, move along the time scales with different dynamics and distinctive rhythms, generating a symphony of knowledge, which can only be integrated with the crosscutting ability of montology as a convergent science (Sarmiento 2020). Indeed, applied montology is the appropriate avenue for developing an environmental awareness of the whole mountainscape. With the wise trend of consilience (Wilson 1998) and the untested hype of noetic science (Nickell 2010), we contribute this chapter with the objective of increasing our epistemology of mountains to include them as socioecological landscapes and not as mere ecosystems.
... Landslides can be devastating agents leading to destruction processes, responsible for losses to the built (Schuster & Highland, 2001) and the natural environment (Geertsema et al., 2009). Landslides play an important role in a variety of ecosystem functions (Myster & Sarmiento, 1998;Walker & Shiels, 2012) and can have enduring legacies. Geertsema and Pojar (2007) argue that landslides are important agents of site, soil, and habitat diversity. ...
Book
This book introduces an innovative approach to sustainable and regenerative mountain development. Transdisciplinary to biophysical and biocultural scales, it provides answers to the "what, when, how, why, and where" that researchers question on mountains, including the most challenging: So What! Forwarding thinking in its treatment of core subjects, this decolonial, non-hegemonic volume inaugurates the Series with contributions of seasoned montologists, and invites the reader to an engaging excursion to ascend the rugged topography of paradigms, with the scaffolding hike of ambitious curiosity typical of mountain explorers. Chapter 8 is available open access under a Creative Commons Attribution 4.0 International License via link.springer.com.
... Landslides can be devastating agents leading to destruction processes, responsible for losses to the built (Schuster & Highland, 2001) and the natural environment (Geertsema et al., 2009). Landslides play an important role in a variety of ecosystem functions (Myster & Sarmiento, 1998;Walker & Shiels, 2012) and can have enduring legacies. Geertsema and Pojar (2007) argue that landslides are important agents of site, soil, and habitat diversity. ...
Chapter
This chapter begins by giving a brief overview of the forces involved in the geodynamics of mountains and mountain ranges, including the processes needed for the generation of mass movement processes. In the remaining parts of this chapter, the following issues associated with mountain landslides are addressed: the anatomy of landslides, common landslide materials, and landslide movement types, along with landslide causes and triggers. The purpose of the final section of this chapter is to reflect on the extent to which the increasing intensity of human activities on mountainscapes, particularly climate change and urbanization, has magnified potential disaster risk for downslope settlements.
Chapter
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To compare revegetation patterns between native and exotic species on abandoned pastures in northern Japan, we surveyed the vegetation structure and the seedbank using a flotation technique in 140 1 1 m plots. Previously introduced grasses such as Poa pratensis, Phleum pratense, and Dactylis glomerata were abundant 20 yr after the pasture abandonment, while Sasa senanensis, native shrub species, regenerated from propagation that had spread from the surrounding forests. S. senanensis shrublands and P. pratensis/P. pratense grasslands established on deep soils while D. glomerata grasslands established on shallow soils. Trees rarely became established on abandoned pastures. The seed density in the seedbank, representing 19 species, ranged from 542 to 2957 seeds/m². The dominant species in the vegetation (P. pratensis, Trifolium repens, and Rumex acetosella) were also common in the seedbank, whereas Cerastium holosteoides var. angustifolium, Chenopodium album var. centrorubrum, and Erigeron canadensis were widespread in the seedbank but did not occur in the extant vegetation. S. senanensis regenerated by vegetative propagation, and P. pratensis and P. pratense developed a seedbank. We concluded that for native species, particularly S. senanensis, vegetative reproduction has an important role on revegetation rather than regeneration from the seedbank, and the dwarf bamboo may be a keystone species in the ecosystems.
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Inappropriate land use practices followed by overexploitation of forest resources have exacerbated environmental degradation to a point that restoration ecology is being considered as a valuable option for conservation and developmeent of the equatorial Andes. -from Author
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The fate of tropandean landscapes in Ecuador is of special concern because of the ongoing biological impoverishment and habitat fragmentation. Ecuador hosts a great variety of ecosystems, ranging from the lush Amazonian forests to snow-capped mountains and from the Andean vergent to the semi-arid Galapagos islands. The topographic complexity of the Andes mountains produces a large number of bioclimatic zones, each with different land use regimes. Ecuador is one of the most biologically diverse nations per unit area (Steinitz-Kannan, Colinvaux, and Kannan 1983) in the neotropics. In addition to the great biodiversity, Ecuador has a rich variety of indigenous cultures, which have lived there for millennia. Post-Colombian anthropogenic pressures have had severe impacts on the land and biotic resource base, with feedbacks on the traditional lifestyles of indigenous peoples. Currently, Ecuador stands at an important environmental crossroad. Therefore, protection of the fragile and fragmented natural tropandean ecosystems is an immediate concern for Ecuadorian land use planning.
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Examines, in a variety of contexts, a number of theoretical and empirical relationships between disturbance (environmental fluctuations and destructive events, whether predictable and/or cyclical or not) and patch dynamics (where discrete spatial patterns possess internal characteristics and also inter-relate with surrounding patch and non-patch areas). The main sections are on: patch dynamics in nature; adaptations of plants and animals in a patch dynamic setting; and implications of patch dynamics for the organisation of communities and the functioning of ecosystems. A final chapter moves towards a general theory of disturbance. All 21 chapters are abstracted separately. -P.J.Jarvis
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Ecologists are aware of the importance of natural dynamics in ecosystems. Historically, the focus has been on the development in succession of equilibrium communities, which has generated an understanding of the composition and functioning of ecosystems. Recently, many have focused on the processes of disturbances and the evolutionary significance of such events. This shifted emphasis has inspired studies in diverse systems. The phrase "patch dynamics" (Thompson, 1978) describes their common focus. The Ecology of Natural Disturbance and Patch Dynamics brings together the findings and ideas of those studying varied systems, presenting a synthesis of diverse individual contributions.
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The recovery of the vegetation in lower montane forest (1440 m elevation) was studied on landslides resulting from the earthquakes and unusually high rainfall in March 1987. The study site was located on the Quijos River watershed, Napo Province, NE Ecuador. Number of individuals of Tessaria integrifolia was negatively correlated with distance from the river, indicating its importance in the newly established riparian habitat. The first herbaceous plants (eg Blechnum, Equisetum) and shrubs (eg Piper, Baccharis, Senecio, Miconia) appeared early in the debris fan at the base of the landslides. Protected gullies and remnant patches of vegetation were favorable microhabitats for establishment of colonizers. A climbing bamboo Chusquea cf. exasperata) appeared on a landslide site two years post-earthquake. Proximity to a pool of potential colonizers is important to the initial species composition of the vegetation on the steepest zone. -from Author