-Darien Global

Drivers of forest cover changes in the Choco

Ecoregion of South America
J. CAMILO FAGUA ,1,2,3,  JACOPO A. BAGGIO ,4,5 AND R. DOUGLAS RAMSEY 1

1
RS/GIS Laboratory, Department of Wildland Resources and the Ecology Center, Utah State University, Logan, Utah 84322 USA
2
School of Informatics, Computing, and Cyber Systems, Northern Arizona University, Flagstaff, Arizona 86011 USA
3
CIAF, Instituto Geografico Agustın Codazzi, Bogota, Distrito Capital, Colombia
4
Department of Political, University of Central Florida, Orlando, Florida 32816 USA
5
Sustainable Coastal Systems Cluster, National Center for Integrated Coastal Research, University of Central Florida, Orlando, Florida
32816 USA

Citation: Fagua, J. C., J. A. Baggio, and R. D. Ramsey. 2019. Drivers of forest cover changes in the Choc

o-Darien Global
Ecoregion of South America. Ecosphere 10(3):e02648. 10.1002/ecs2.2648

Abstract. Tropical rain forests are suffering the highest deforestation and reforestation ever recorded.
Interactions between direct (proximate or direct causes) and indirect (underling or indirect causes) drivers
could cluster these forest cover changes forming hotspots (areas that exhibit significant spatial correlation
of deforestation or reforestation transitions). Using land use–land cover maps and global (I) and local (Ii)
Moran’s tests, we identified these hotspots in the Choco
-Darien Global Ecoregion (CGE) of South America,
a natural region that was declared one of the top 25 hotspots for conservation priorities in the world. Sub-
sequently, we tested and studied the effects and interactions between deforestation and reforestation hot-
spots and their direct and indirect drivers using Bayesian Structural Equation Modeling (Bayesian SEM).
We found that deforestation and reforestation were spatially auto-correlated forming hotspots (I = 0.49,
P = 0.001 for deforestation transitions and I = 0.48, P = 0.001 for reforestation transitions). Also, hotspots
of deforestation and reforestation were auto-correlated within municipality borders (I = 0.5, P = 0.001 for
deforestation transitions; I = 0.49, P = 0.001 for reforestation transitions). Eighteen municipalities located
on the border between Colombia and Ecuador showed significant aggregations of deforestation hotspots,
while thirty-four municipalities in three areas of Colombia and the area between the Colombian and
Ecuadorian border showed significant clustering of reforestation hotspots. Eleven of these municipalities
presented significant clustering of both reforestation and deforestation hotspots. The Bayesian SEM for
deforestation showed that population growth and road density were indirect drivers of deforestation hot-
spots (0.191 and 0.127 standard deviation units). The Bayesian SEM for reforestation found that armed con-
flicts, Gross Domestic Product, and average annual rain were indirect drivers related to reforestation
hotspots (0.228, 0.076, and 0.081 standard deviation units, respectively). Our assessment shows a novel
methodology to study interactions among direct and indirect drivers of forest change and their potential
dissimilar effects on forest transitions.

Key words: Bayesian Structural Equation Modeling; deforestation; hotspots; Moran’s test; reforestation; spatial
autocorrelation.

Received 10 December 2018; revised 1 February 2019; accepted 5 February 2019. Corresponding Editor: Mary
Cadenasso.
Copyright: © 2019 The Authors. This is an open access article under the terms of the Creative Commons Attribution
License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.
  E-mail: Camilo.Fagua@nau.com

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INTRODUCTION
Tropical rain forests consist of evergreen vege-
tation that reaches 30 m in height and are located
in the lowlands of the wet tropical climatic zone
of the planet (Primack and Corlett 2009). Differ-
ent studies have documented extensive defor-
estation and minor reforestation (regrowth of
secondary vegetation) in these forests during the
first ten years of the 21st century. Consequently,
the tropical rain forest domains have exhibited
significant and constant forest loss (Sanchez-
Cuervo et al. 2012, Aide et al. 2013, Hansen et al.
2013, Keenan et al. 2015, Qin et al. 2017). Defor-
estation and reforestation dynamics play a key
role in global environmental changes, degrading
or protecting ecosystem services that are funda-
mental to human development, such as climate
and soil regulation, water supply, carbon storage,
and biodiversity (Lambin et al. 2003b, Leblois
et al. 2017). Identifying the causes that lead to
reforestation and deforestation is key to develop-
ing future policies that address deforestation and
provide incentives for reforestation in order to
protect these essential ecosystem services.
Causes of forest change can be classified into
two types: (1) direct drivers (proximate or direct
causes) which are immediate actions that directly
result in forest loss or gain; in other words, direct
drivers refer to the land cover that replaces forest
or converts back to forest; (2) indirect drivers
(underling or indirect causes) are interactions of
social, economic, political, cultural, or other pro-
cesses that indirectly affect forest changes (Geist
and Lambin 2001, 2002). The previous statement
has been the basis for a wide array of literature
focused on studying direct (Roberts et al. 2002,
Flamenco-Sandoval et al. 2007, Houghton 2012,
Mueller et al. 2012, Tng et al. 2012, Alejandra
Chadid et al. 2015, Armenteras et al. 2017) or
indirect drivers (Armenteras et al. 2006, 2013,
2017, Hosonuma et al. 2012, Kissinger et al.
2012, Mueller et al. 2013, Leblois et al. 2017) to
assess forest cover change at different spatial
scales throughout the tropical rain forest domain.
Studies that include both direct and indirect dri-
vers as well as their interactions have been less
prevalent (Davalos et al. 2014, Richards 2015).
In the Neotropics, the tropical terrestrial ecore-
gions of Central and South America, the loss of
forest cover is mainly a product of farming
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FAGUA ET AL.
expansion, logging, mining, and, more generally,
by natural resource extractive activities (Lambin
et al. 2003b, DeFries et al. 2010). However, farm-
ing is estimated to be the largest specific direct
driver of deforestation (Hosonuma et al. 2012,
Kissinger et al. 2012). On the other hand, regio-
nal analyses of indirect drivers of deforestation
in the Neotropics have found that deforestation
is related to socioeconomic (e.g., National Gross
Domestic Product) and demographic indicators
followed by accessibility (e.g., existence of roads,
rivers; Rudel and Roper 1997, Aide et al. 2013,
Armenteras et al. 2017). More localized studies
of deforestation also highlight the importance of
socioeconomic as well as demographic drivers
(Armenteras et al. 2013, Sanchez-Cuervo and
Aide 2013), as well as topography, soil fertility,
and climate variables (Laurance et al. 2001, Stei-
ninger et al. 2001, Sanchez-Cuervo and Aide
2013).
With respect to reforestation, previous litera-
ture has identified abandonment of farming
areas as the main cause of reforestation (Aide
et al. 2013). Other documented causes of land
abandonment include the following: (1) the
migration of rural populations to urban areas
(Aide and Grau 2004); (2) modernization of agri-
cultural systems that increases production while
using less land (Grau and Aide 2008); (3) the shift
from agriculture to manufacturing in some Car-
ibbean islands (Grau et al. 2003); and (4) armed
conflicts in the case of Colombia (Sanchez-
Cuervo and Aide 2013). Since the migration of
rural populations to urban areas was common in
Central and South America during the 21st cen-
tury, it is also expected that countries with recent
armed conflicts have higher land abandoned and
reforestation trends. Reforestation has also been
indirectly related to other regional and local fac-
tors (e.g., population density, socioeconomic
indicators, climate, topography, soil fertility, and
accessibility; Rudel et al. 2002, Grau et al. 2003,
Lambin et al. 2003a, Sanchez-Cuervo et al. 2012,
Aide et al. 2013).
Direct and indirect drivers of deforestation
and reforestation could vary spatially promoting
or restricting forest transitions in natural land-
scapes (Grau et al. 2003, Sanchez-Cuervo and
Aide 2013). In this case, deforestation and refor-
estation may exhibit statistically significant spa-
tial clustering or “hotspots” (Ferreira et al. 2007,
2 March 2019 ❖ Volume 10(3) ❖ Article e02648

Reddy et al. 2016, Harris et al. 2017). The spatial
identification of deforestation and reforestation
hotspots could be an important tool for the
enforcement of environmental laws and regula-
tions and for the preservation and management
of forest areas. Deforestation hotspots may indi-
cate massive, and improper logging operations,
suggesting insufficient enforcement of environ-
mental laws (Ferreira et al. 2007), as well as iden-
tifying areas where forest protection strategies
should be implemented (Sanchez-Cuervo and
Aide 2013). Whereas reforestation hotspots indi-
cate areas where forest restoration programs
might be established (Sanchez-Cuervo and Aide
2013). In the Neotropics, these hotspots are tradi-
tionally estimated based on forest change trends
between administrative subdivisions (e.g., coun-
tries, states, municipalities; Myers 1993, Sanchez-
Cuervo and Aide 2013, Reddy et al. 2016).
However, both types of transitions can exhibit
statistically significant spatial clustering within
the same administrative division (Hansen et al.
2013, Harris et al. 2017). Proximity or overlap
between areas identified as deforestation and
reforestation hotspots could suggest that farming
activities are not established permanently after
forest removal. Conversely, remoteness between
deforestation and reforestation would show per-
manent use of the land after deforestation.
Increasing the scale of analysis by identifying
hotspots at the spatial scale of land use–land
cover (LULC) maps would allow more accurate
estimations.
Using Bayesian Structural Equation Modeling
(Bayesian SEM), we estimated the relationships
between deforestation and reforestation hotspots
and their direct and indirect drivers in the
Choco
-Darien Global Ecoregion (CGE) of South
America. Here, we focus on the effects of indirect
drivers on the appearance of deforestation and
reforestation hotspots. The CGE is a natural
region that was declared as one of the top 25 hot-
spots for conservation priorities in the world
(Myers et al. 2000, Olson et al. 2001, WWF 2016).
Our two main objectives were (1) to build a
model to quantify and compare the effect of
direct and indirect drivers on the appearance of
deforestation hotspots using a method where all
these drivers could interact, and (2) to build an
analogous model to compare direct and indirect
drivers of reforestation and their relationship
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FAGUA ET AL.
with the appearance of reforestation hotspots. To
construct the models, we first assessed if defor-
estation transitions were spatially clustering to
form deforestation hotspots between 2002 and
2015 at the original spatial scale (250 m nominal
pixel size) of LULC maps developed using
MODIS imagery. We selected this time period to
build the LULC maps due to the availability of
MODIS imagery. Likewise, a similar assessment
was performed to test whether reforestation tran-
sitions were spatially clustered. Finally, we
assessed whether deforestation and reforestation
hotspots were spatially correlated with the delin-
eation of different municipalities in the region.
Following our analysis of spatial correlations, we
addressed direct and indirect drivers of defor-
estation and reforestation at the municipality
level to explain the emergence of hotspots.
METHODS
Study area
The CGE is located along the Pacific Coast
from southern Panam
a to northeastern Ecuador.
CGE also includes the lowlands of the Mag-
dalena River Valley between the central and
western branches of the Colombian Andes
(Fig. 1A; Olson et al. 2001, WWF 2017). Past geo-
logic events have turned the CGE into one of the
most diverse and endemic tropical rain forest
areas on the planet (Gentry 1986, Gregory-Wod-
zicki 2000, WWF 2017). The CGE is also the raini-
est area on Earth (Poveda and Mesa 2000);
consequently, LULC analyses using high spatial
resolution remote sensing are limited due to
cloud cover. Although there are few studies on
forest change within the territories of the coun-
tries that share the CGE (Gonz
alez et al. 2011,
Ministerio del Ambiente del Ecuador 2012, San-
chez-Cuervo et al. 2012) and other similar regio-
nal and global analyses (Sanchez-Cuervo et al.
2012, Aide et al. 2013, Hansen et al. 2013,
Keenan et al. 2015, Qin et al. 2017), the CGE lack
of a spatial analysis of LULC change focused on
its land.
Classification and mapping of LULC
Based on previous regional LULC studies that
included areas in the CGE (Friedl et al. 2010,
Gonz
alez et al. 2011, Ministerio del Ambiente
del Ecuador 2012, Sanchez-Cuervo and Aide
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 FAGUA ET AL.

-Darien Global Ecoregion (CGE): (A) estimated historical extent of Tropical Rain Forest (TRF)
Fig. 1. The Choco
in South America: TRF-CGE (estimated TRF in CGE), TRF-Amz (estimated TRF in Amazon basin), and TRF-BrAt
(estimated TRF in Brazilian Atlantic Forest). Land use and land cover maps for the CGE at 2002 (B) and 2015 (C).
(These maps are available at https://zenodo.org/record/2559673#.XF0DJDNKiM8)

2013), we classified landscapes into eight general
LULC classes:
1. Forest: includes areas of tropical rain forest
with trees taller than 30 m. This is the pri-
mary natural cover in the CGE (Etter et al.
2008, Primack and Corlett 2009, Rangel
2011).
2. Secondary vegetation: formed by natural
regenerations of vegetation, such as shrubs,
small trees, or mosaics of both.
3. Wetland: The CGE has a complex of river
basins with swamps and shallow lakes
(“ci
enagas”) covering large areas along the
rivers.
4. Grassland: formed by areas of introduced
grass species which are used primarily for
cattle grazing. Within the CGE, large
areas of native grasses do not occur as
natural vegetation (Etter et al. 2008, Ran-
gel 2011).
5. Crops: consists of annual or semiannual
crops (corn, sugar canna, plantain, mainly).
6. Palm plantations: Extensive areas of the CGE
have been cultivated with African palm
(Elaeis Guineensis Jacq; Castiblanco et al.
2015). The productive life of this crop is
about 25 yr, at which point plantations are
replanted with younger palms (Mingorance
et al. 2004, FEDEPALMA 2011).
7. Settlements: urban areas and areas where
infrastructure is established.
8. Water bodies: Areas covered by continental
waters, including rivers and lakes.
9. Bare areas: This class was not considered in
the final analysis due to its low representa-
tion on the landscape.
We produced two final LULC maps for the
CGE, one for 2002 and another for 2015. These
two maps were generated from an initial series
of 15 annual LULC maps from 2001 through
2015 using a Random Forest classification of
14,847 training sites using MODIS data as predic-
tor variables (Appendix S1: Table S1). A 20%

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stratified random extraction from the original
18,559 training sites prior to the Random Forest
classification (3712 sites) was used to generate a
confusion matrix to assess accuracy which
resulted in a Kappa accuracy of 0.87 (SD = 0.008).
Transitions from forest or secondary vegetation to
farm-related LULC classes (grassland, crop, palm
plantations) were identified as deforestation tran-
sitions in the 2015 map. Conversely, transitions
from farm-related LULC classes to secondary veg-
etation were identified as reforestation transitions
in the 2015 map. Appendix S1 shows the method-
ology applied to create the LULC maps for the
CGE. These maps can be downloaded in GeoTIFF
format from: https://zenodo.org/record/2559673#.
XF0DJDNKiM8
Spatial identification of deforestation and
reforestation hotspots
We followed two methodological steps to iden-
tify areas where deforestation and reforestation
transitions were significantly clustered. In the
first step, we ran two global Moran’s (I) tests, one
for the deforestation transitions and another for
the reforestation transitions, to test whether these
transitions were spatially auto-correlated
between the 2002 and the 2015 LULC maps of
the CGE. Moran’s I is a standardized measure of
correlation between observations in neighboring
areas; it is commonly employed in the analysis of
spatial data (Cliff and Ord 1981, Bivand et al.
2013). I is a linear correlation coefficient calcu-
lated as a ratio of the product of the variable of
interest and its spatial lag, with the cross-product
of the variable interest, and adjust for the spatial
weights used (Bivand et al. 2013):
Pn Pn  
n i¼1
 
j¼1 wijðyi  yÞ yj  y
I ¼ Pn Pn Pn
i¼1 j¼1 wij 2
i¼1 ðyi  yÞ
where n is the number of features, yi is the ith
observation, 
y is the mean of the variable of inter-
est, and wij is the spatial weight of the link
between i and j. I usually ranges from 1 to +1; I
values significantly below 1/(n  1) indicate
negative spatial autocorrelation and I values sig-
nificantly above 1/(n  1) indicate positive spa-
tial autocorrelation (Cliff and Ord 1981). In our
case, n corresponded to rectangular matrix of
3 9 3 pixels in the 2002–2015 transition LULC
raster map where y represents the deforestation
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FAGUA ET AL.
or reforestation value in the 3 9 3 matrix. To
arrive at the 3 9 3 matrix size, we tested a vari-
ety of matrices from 3 9 3 pixels through
37 9 37 pixels. The relation I vs. matrix size for
deforestation (P = 0.001; R = 0.94) and refor-
estation (P = 0.001; R = 0.97) showed a typical
negative exponentially shape of auto-correlated
relationships (Appendix S2: Fig. S1); we therefore
selected the 3 9 3 pixel matrix for our analysis
due to its higher I (Dale and Fortin 2002). Defor-
estation values were calculated by assigning val-
ues of 1 to deforestation transitions and 0 to the
other transitions in the 2002–2015 transition
LULC map. Following the same schema, refor-
estation values were calculated giving values of 1
to reforestation transitions and 0 to the others
(Shortridge 2007).
For our second methodological step, after test-
ing the global spatial autocorrelation for defor-
estation and reforestation transitions, we ran
local Moran’s (Ii) tests to identify the 3 9 3 matri-
ces where deforestation or reforestation transi-
tions were statistically significantly clustered.
Since I is the slope of a linear relation, Ii detects
the 3 9 3 matrices with significant influence on
the slope. Consequently, the formula for Ii is sim-
ilar to I, but Ii is calculated separately for each
feature, in our case each 3 9 3 matrix:
P
ðyi þ yÞ nj¼1 wijðy  yÞ
Ii ¼ Pn
ðyi yÞ
2
i¼1
n
The results of these analyses are shown in a
map of deforestation hotspots and another of
reforestation of hotspots (Fig. 2A, B) and were
implemented using the R “raster” package (R
Core Team 2014, Hijmans et al. 2016). These
maps of deforestation and reforestation hotspots
can be downloaded as GeoTIFF format in the
next link: https://zenodo.org/record/2559673#.
XF0DJDNKiM8
Spatial autocorrelation of hotspots in relation to
municipalities
We used a two-step process to identify munici-
palities where deforestation and reforestation
hotspots were significantly clustered. The first
step was to calculate the global Moran’s (I) to
determine whether there was spatial autocorrela-
tion between deforestation and reforestation hot-
spots with municipality borders within the CGE.
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Fig. 2. Deforestation (A) and reforestation (B) hotspots on the land use and land cover map at 2015. Spatial
correlation between deforestation and reforestation hotspot areas was mapping (C). (These maps are available at
https://zenodo.org/record/2559673#.XF0DJDNKiM8)
Because size and shape of the municipalities
vary, we used the distance-based neighbor
method to evaluate spatial clustering regardless
of the distance at which the tests were applied.
This method selects the nearest neighbor(s) given
the distance within the feature centroid (here the
centroid refers to the centroid of the municipality
in question). We ran these I tests from one to six
neighbors with the null hypotheses stating that
the deforestation or reforestation hotpots were
randomly distributed among the municipalities.
As before, I values significantly below 1/
(n  1) indicated negative spatial autocorrelation
and I values significantly above 1/(n  1) indi-
cated positive spatial autocorrelation, where n is
the number of municipalities (Cliff and Ord
1981). To identify the municipalities where defor-
estation or reforestation hotspots were signifi-
cantly clustered, we ran local Moran’s (I) tests.
Since I is the slope of a linear relationship, Ii
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FAGUA ET AL.
detected the municipalities with a significant
influence on this slope. The results of these
analyses are shown in Moran scatter plots and
maps of municipalities (Fig. 3). We used the R
package “spdep” (Bivand and Piras 2015) for
these analyses.
Indirect drivers of deforestation and reforestation
We assessed the effect of 10 different variables
as indirect drivers (Table 1) across the 311 munic-
ipalities located in the CGE (30 municipalities of
Panam
a, 212 of Colombia, and 69 of Ecuador).
All variables were defined at the municipality
level since the municipalities (called Municipali-
ties in Colombia, Districts in Panam
a, and Can-
tones in Ecuador) were the primary units in our
analysis. These 10 variables can be classified into
six categories:
(1) Accessibility: transportation networks and
rivers (Geist and Lambin 2002). The accessibility
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variables in our analysis included paved and
unpaved roads from national cartographic data
from each country (IGAC 2017, IGNTG 2017,
SNI 2017). Rivers were extracted from Hydro-
SHEDS (Hydrologic data derived from Elevation
Derivatives produced by the NASA Shuttle
Radar Topographic Mission at various scales;
Lehner et al. 2008). Density was estimated as the
length of routes or rivers (km) per km2.
(2) Biophysical environment: Landscape physical
characteristics can affect access to forested areas,
thus influencing deforestation and reforestation
transitions (Geist and Lambin 2002). We used the
mean of topographic slope (degrees) and stan-
dard deviation of altitude (estimated as MASL)
as biophysical variables; both variables were
assembled from SRTM90 (NASA Shuttle Radar
Topographic Mission; Jarvis et al. 2008).
(3) Climate: CGE has important variations in
precipitation and temperature that may affect
deforestation and reforestation transitions.
Although the CGE is one of the rainiest places in
the world (Poveda and Mesa 2000), its northern
and southern extremes have lower precipitation
than the central section. Temperature also varies
across the CGE, increasing from south to north.
We selected mean annual precipitation (mm/yr)
and mean annual temperature (°C) from the 30
arc-second WorldClim layers (Fick and Hijmans
2017).
(4) Demography: Population growth is consid-
ered a primary indirect cause of deforestation
(Ehrhardt-Martinez 1998, Geist and Lambin
2001). We estimated this variable as:
Pf  Po
Pg ¼
Pf
where Po and Pf are the population for every
municipality in 2000 and 2010, respectively. Pop-
ulation was extracted from the official national
census of each country (DANE 2017a, INEC-
Ecuador 2017, INEC-Panam
a 2017a). Panam
a
and Ecuador have census for 2000 and 2010, and
Colombian has census for 1984 and 2005. We
used the official projections of population for
2000 and 2010 for Colombia (DANE 2017a).
(5) Armed conflicts: These types of conflicts
have occurred in several areas of the CGE, espe-
cially within Colombia, produced by land pos-
session, social and economic inequality, politic
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FAGUA ET AL.
issues, illegal crops and drug production, and
illegal mining (Yaffe 2011, Guzm
an et al. 2016,
Fajardo 2017). The effects of armed conflicts on
environment, wildlands, and biodiversity remain
complex (Hammill et al. 2016). Some scholars
have found that armed conflicts put on risk the
conservation by reducing the effectiveness of
protection of wild life or natural vegetation (Bey-
ers et al. 2011, Hammill et al. 2016); others have
showed benefit effects on forest protection by
creating exclusion zones (John 1998) or hindering
extractive industries (McNeely 2003). Also,
armed conflicts have been related to land aban-
donment and posterior forest regeneration in
Colombia (Sanchez-Cuervo and Aide 2013). We
used two variables to include armed conflicts in
our analysis: Number of armed conflicts and
number of reported fatalities produced by the
armed conflicts between 2000 and 2015 (Sund-
berg and Melander 2013, Croicu and Sundberg
2017).
6) Economy: We included Gross Domestic Pro-
duct (GDP) growth between 2000 and 2010 as
our main economic variable. Each country has
official GDP estimates for the country as well as
the first administrative level (Departments in
Colombia and Provinces in Panam
a and Ecua-
dor), but only Ecuador has an estimate of GDP
for municipalities (BCE 2017, DANE 2017b,
INEC-Panam
a 2017b). To estimate the GDP in the
municipalities of Colombia and Panam
a, we
allocated the GDPs of every Department in
Colombia and Province in Panam
a to their corre-
sponding municipalities according to the propor-
tion of population of that municipality within the
Department or Province.
Bayesian SEM
Traditionally, the effect of direct and indirect
drivers on forest transitions is estimated using
different types of regressions (linear regression,
logistic regression, generalized linear models,
random forest regression, etc.; Roberts et al.
2002, Armenteras et al. 2006, 2013, 2017, Fla-
menco-Sandoval et al. 2007, Hosonuma et al.
2012, Houghton 2012, Kissinger et al. 2012,
Mueller et al. 2012, 2013, Tng et al. 2012, San-
chez-Cuervo and Aide 2013, Alejandra Chadid
et al. 2015, Leblois et al. 2017); these statistical
tests assume that all direct and indirect drivers as
predictors have the same potential direct effect
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 FAGUA ET AL.

Fig. 3. Municipalities with significant clustering of deforestation and reforestation hotspots. (A) Moran scatter
plot for deforestation; red diamonds represent the municipalities with significant influence on the Moran’s I

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 FAGUA ET AL.

(Fig. 3. Continued)
global test for deforestation. (B) Moran scatter plot for reforestation; violet diamonds represent the municipalities
with significant influence on the Moran’s I global test for reforestation. (C) Maps of municipalities with signifi-
cant influence on the Moran’s I global test for deforestation. (D) Maps of municipalities with significant influence
on the Moran’s I global test for reforestation.

on a forest transition, deforestation, or reforesta-
tion. However, the “equality of effect” assump-
tion is not consistent with the analysis of direct
and indirect drivers of forest transitions (see
Geist and Lambin 2001, 2002).
We propose that Bayesian Structural Equation
Modeling (Bayesian SEM) is an appropriate tech-
nique to estimate the effect of direct and indirect
drivers on the appearance of deforestation and
reforestation hotspots. Bayesian SEM uses the
variances and covariances in a dataset of vari-
ables to test the most probable path of direct and
indirect relationships among the variables based
on an initial hypothesis (Merkle and Rosseel
2016). Thus, Bayesian SEM and LULC maps
allow us to estimate (1) the effects of indirect dri-
vers on direct drivers, (2) the effects of direct dri-
vers on hotspots, (3) the effects of indirect drivers
on hotspots, and (4) other possible interactions in
the set of variables. Bayesian SEM also presents
fewer limitations that the likelihood approach of
SEM given by sample size, allowing the model-
ing of more complex networks of variables (Lee
2007).
We applied two linear Bayesian SEM to exam-
ine the relationships among the measured vari-
ables at the municipality level. One model was
built for deforestation where the primary vari-
able was the number of hotspots of deforestation.
Another model was constructed for reforestation
in which the main primary variable was the
number of hotspots of reforestation. We used the
R package “blavaan” (Merkle and Rosseel 2016)
to run the Bayesian SEM. We used the default
settings of “blavaan” to define the a-priory distri-
butions for each variable and parameters in the
model. We selected “manual” convergence to
add the rest of the settings according to Plummer
(2015): The number of burnin iterations was set
to 4000, the number of adaptive iterations to use
at the start of the simulation was 1000, and the
total number of samples to take after burnin was

Table 1. Variables evaluated as indirect drivers of deforestation and reforestation.

Category Variable Description Resource

Accessibility Density of roads Length of road (km) of first, second, IGAC (2017), IGNTG (2017), SNI (2017)
and third level per km2
Accessibility Density of rivers Length of rivers (km) per km2 Lehner et al. (2008)
Biophysical Topographic slope Degrees of topographic slope from Jarvis et al. (2008)
environment STRM 90
Biophysical Standard deviation Standard deviation of pixel values of Jarvis et al. (2008)
environment of altitude STRM 90
Climate Temperature Mean annual temperature (°C) from Fick and Hijmans (2017)
1950 to 2000
Climate Precipitation Mean annual precipitation (mm/yr) Fick and Hijmans (2017)
from 1970 to 2000
Demography Population growth Human population change from 2000 DANE (2017a), INEC-Ecuador (2017),
to 2010 INEC-Panam
a (2017a)
Armed conflicts Armed conflicts Number of armed conflicts between Sundberg and Melander (2013), Croicu
2000 and 2015 and Sundberg (2017).
Armed conflicts Fatalities of armed Reported fatalities produced by the Sundberg and Melander (2013), Croicu
conflicts armed conflicts between 2000 and and Sundberg (2017)
2015
Economy Gross Domestic Gross Domestic Product growth at BCE (2017), DANE (2017b), INEC-Panam
a
Product the municipality level between 2000 (2017b)
and 2010

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10,000 (Plummer 2015, Denwood 2016). We also
recoded the set of variables to units of no more
than two digits to reduce the variance as per a
requirement to run SEM (Rosseel 2012). In the
Bayesian SEM, the goodness of fit of a hypothe-
sized model is evaluated by the posterior predic-
tive P value (PPP); PPPs higher than 0.05 indicate
a good model fit. The direct effect of a variable
on another is measured by unstandardized pos-
terior path coefficients (Post.Mean values) and
the standardized posterior path coefficients
(Std.all values). Post.Mean represents the slope
in the linear relation between a couple of endoge-
nous and exogenous variables while the other
exogenous variables are constant. In other
words, Post.Means shows the percentage of
change of the endogenous variable when the
exogenous variable changes one raw unit.
Post.Means are also estimated with the posterior
standard deviation (Post.SD), the 95% highest
posterior density interval (HPD.025 and HPD.
975), and the potential scale reduction factor for
assessing chain converge (PSRF). Potential scale
reduction factor lower than 1.2 indicates that
convergence was reached by the variable or
parameter (Lee 2007, Merkle and Rosseel 2016).
On the other hand, Std.all shows the change of
standard deviation units in the endogenous
variable when the exogenous variable changes
one standard deviation unit while the other
exogenous variables are constant. Due to Std.all
estimation in constant units (standard devia-
tions), Std.all can be used to estimate indirect
effects and to compare the effects among vari-
ables. Indirect effects are estimated by multiply-
ing the Std.all through a path (Grace and Bollen
2005). Bayesian SEM also estimates global fit
measures that help select the best model among
several hypothesized models. These global fit
measures are Deviance Information Criterion
(DIC), Widely Applicable Information Criterion
(WAIC), and Leave-one-out cross-validation
(LOOIC); smaller values of DIC, WAIC, and
LOOIC are the models with a better global fit
(Merkle and Rosseel 2016, Vehtari et al. 2016a,
b). In SEM analysis, Bayesian or Likelihood, the
point of view of the researcher is the other crite-
ria to select a model among hypothesized mod-
els with PPP higher than 0.05 (Grace et al.
2012). In the case of deforestation, we examined
a first group of models where the main
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FAGUA ET AL.
endogenous variable, hotspots of deforestation,
was affected directly by population growth, and
this variable in turn was affected by the other
possible indirect drivers. We also examined
another group of models that included latent
variables between the main endogenous vari-
able and the different possible indirect drivers.
These two groups of models resulted in PPP
lower than 0.05. We found valid moles when the
main endogenous variable is affected by the
direct driver (areas of farming that replaced for-
est and secondary vegetation), and this variable
in turn was affected by the indirect drivers
(economy, demography, accessibility, climate,
and biophysical environment). In this group, we
selected the most coherent model with the
smallest values of DIC, WAIC, and LOOIC as
the final model. In the case of reforestation, we
examined models where the main endogenous
variable is affect by the direct driver (areas of
farming that replaced forest and secondary veg-
etation), and this variable in turn was affected
by the indirect drivers (armed conflicts, demog-
raphy, accessibility, climate, and biophysical
environment). This model examination included
the use of latent variables. We also selected the
most coherent model with the smallest values of
DIC, WAIC, and LOOIC as the final model for
deforestation. The R code to build our Bayesian
SEMs and the data to run these codes is avail-
able in Data S1.
RESULTS
Global Moran’s (I) tests showed that both
deforestation and reforestation transitions were
spatially auto-correlated within the CGE
(I = 0.49, P = 0.001 for deforestation transitions;
I = 0.48, P = 0.001 for reforestation transitions).
Local Moran’s (Ii) tests detected hotspots of
deforestation and reforestation, that is, areas
where deforestation and reforestation transitions
were significantly auto-correlated across the
CGE (Fig. 2A, B). We also found that areas iden-
tified as hotspots of deforestation were spatially
correlated to areas identified as hotspots of refor-
estation (R = 0.8, P = 0.01), showing that both
transition types tended to occur in close proxim-
ity (Fig. 2C).
Additional Global Moran’s (I) tests determined
that hotspots of deforestation and reforestation
10 March 2019 ❖ Volume 10(3) ❖ Article e02648

were auto-correlated with municipalities across
the CGE (I = 0.5, P = 0.001 for deforestation
transitions; I = 0.49, P = 0.001 for reforestation
transitions). The corresponded Local Moran’s
(Ii) tests (Fig. 3A, B) identified municipalities
with significant hotspot clustering for both
deforestation and reforestation. Eighteen
municipalities located near the border of
Colombia and Ecuador presented the most sig-
nificant aggregation of deforestation hotspots;
three in the Colombian side and 15 in the
Ecuadorian side (Fig. 3C). On the other hand,
34 municipalities located in three areas of
Colombia as well as the area around the Colom-
bian and Ecuadorian border showed significant
clustering of reforestation transitions (Fig. 3D).
We found that 11 municipalities located around
the international border between Colombia and
Ecuador presented significant clustering of both
Fig. 4. Bayesian Structural Equation Model for deforestation. Unstandardized posterior path coefficients in
black and standardized posterior path coefficients in red.
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FAGUA ET AL.
deforestation and reforestation hotspots: in
Colombia—Barbacoas, Mosquera, and Tumaco;
and in Ecuador—Eloy Alfaro, Las Golondrinas,
Pedernales, Puerto Quito, Quininde, Rio Verde,
San Lorenzo, and Tulcan.
The selected Bayesian SEM for deforestation
reached a PPP of 0.944 (Fig. 4; Data S1), indicat-
ing a very good model fit. The selected model
showed the lowest values of DIC (2744), WAIC
(2789), and LOOIC (2783) compared other mod-
els. All relationships between variables and their
parameters in the model had a PSRF lower than
1.2, indicating that they reached convergence.
Standardized posterior path coefficients between
indirect and direct drivers of deforestation
showed positive relationships between forest
replaced by farming and population growth, as
well as forest replaced by farming and road den-
sity. Conversely, rain, GDP, topographic slope,
11 March 2019 ❖ Volume 10(3) ❖ Article e02648

Fig. 5. Bayesian Structural Equation Model for reforestation. Unstandardized posterior path coefficients in
black and standardized posterior path coefficients in red.
and temperature showed a negative relationship
with forest replaced by farming. Population
growth had the largest total effect on the forma-
tion of deforestation hotspots (0.191), which
included direct (0.117) and indirect effects (0.07).
Road density had a high indirect effect on defor-
estation hotspots (0.127). All other indirect dri-
vers of deforestation had negative relationships
with deforestation hotspots (Fig. 4).
The selected Bayesian SEM for reforestation
also resulted in a very good model fit with a PPP
of 0.133 (Fig. 5; Data S1). This model also
showed the lowest values of DIC (2301), WAIC
(2317), and LOOIC (2320) compared to other
models. All relationships between variables and
their parameters in the model reached conver-
gence (PSRF < 1.2). Standardized posterior path
coefficients between indirect and direct drivers of
reforestation found positive relationships
between farming that had transitioned to sec-
ondary vegetation and areas of armed conflicts,
GDP, and rain (Fig. 5); armed conflicts showed
the highest indirect positive effect on reforesta-
tion hotspots (0.228). Temperature was
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FAGUA ET AL.
negatively related to farming that had transi-
tioned to secondary vegetation.
DISCUSSION
Spatial correlation
Our results confirm that deforestation and
reforestation in the CGE tended to be spatially
clustered forming hotspots. This suggests that
forest changes were not accidental processes; for-
est changes responded to drivers that deter-
mined their geographical location and intensity.
We also found that many of the reforestation hot-
spots were spatially adjacent to deforestation
hotspots, suggesting that after harvesting mature
forest (deforestation) many of these lands were
not used for other purposes in subsequent years,
thus allowing the generation of secondary vege-
tation. Fagua and Ramsey (2019) showed that
~60% of harvested forests converted to secondary
vegetation between 2001 and 2015 across the
CGE. This relatively high proportion of forest
harvest immediately followed by secondary veg-
etation growth is explained by three main factors
12 March 2019 ❖ Volume 10(3) ❖ Article e02648

within the CGE and the surrounding tropical for-
ests: (1) Deforestation occurs in areas with fragile
and unproductive soils, such as the Choco-Dar-
ien, an ecoregion within the CGE (IGAC 2015);
(2) colonists remove older forests to demonstrate
land use in order to gain ownership of the prop-
erty (Davalos et al. 2014); and (3) the decline of
Colombian and Ecuadorian agricultural sectors
between 2000 and 2010 reduced cultivated areas
(grassland, crops, palm), allowing for encroach-
ment of secondary vegetation (BCE 2010a, b,
Marrugo 2013).
We also identified the municipalities with sig-
nificant clustering of deforestation or reforesta-
tion; surprisingly, we detected that 11
municipalities presented significant clustering of
both deforestation and reforestation hotspots,
corroborating the aforementioned statement that
a high proportion of areas where mature forest
has been deforested are not used in agriculture.
These results also demonstrate that hotspot iden-
tification from land cover change maps allows
more accurate estimations than those based on
forest change trends, where the analyzed admin-
istrative unit can only be a deforestation or a
reforestation hotspot, but not both (Aide et al.
2013, Armenteras et al. 2013, 2017, Sanchez-
Cuervo and Aide 2013). These 11 municipalities
were located around the international border
between Colombia and Ecuador, an active areas
of illegal and legal crops. The Colombian side has
the second largest concentration of illegal crops
of Coca plant (Erythroxylum coca) in that country.
The distribution of hotspots of deforestation and
reforestation matched highly with the distribu-
tion of coca crops (UNODC 2018), showing the
impact of this illegal activity on the conservation
the biodiversity of the CGE. This is a cycle where
forests areas are cut down for planting Coca, then
authorities eradicate the Coca crops allowing sec-
ondary vegetation to grow, but Coca is planted
again in the same or other areas after some
months or years. The Ecuadorian side of the bor-
der is the second highest area of colonization in
that country (Ministerio del Ambiente del Ecua-
dor 2012, Sierra 2013). Colonist cut down the for-
est to develop agriculture; however, the low
productivity of these soils and the economic crisis
of agricultural sector during these years in Ecua-
dor reduced the agricultural land use, allowing
for the growth of secondary vegetation.
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FAGUA ET AL.
Deforestation
The Bayesian SEM of deforestation for the CGE
showed that human population growth was most
related to deforestation hotspots between 2002
and 2015. Deforestation hotspots were principally
located in municipalities found in the northern
portion of Ecuador and around the Ecuadorian
and Colombian border. These municipalities have
higher population growth rates and are main
migration destinations for colonist compared to
other municipalities in the CGE (Go
mez 2011,
Gonz
alez et al. 2011, Sierra 2013). Our results
agree with several scholars who have found
demographic variables as drivers of deforestation
in tropical forests across Latin America at national
(Armenteras et al. 2013, Sanchez-Cuervo and
Aide 2013) and regional levels (Laurance et al.
2002, Aide et al. 2013). We included population
growth as an indirect driver (a factor that influ-
ences the expansion of farming) as well as a direct
driver of deforestation due to its influence on
other land uses not documented in the MODIS-
derived LULC maps. These activities include
urban/rural infrastructure development as well as
mining and logging, which occurs historically in
the CGE (Mosquera 1978, Zapata 2013).
Road density was positively and indirectly
related to deforestation hotspots. From 2002 to
2015, the best-preserved forest of the CGE was
located along the pacific coast from Panam
a and
Gulf of Urab
a (Colombia) to the border between
Colombia and Ecuador. This belt-like section of
well-preserved forest has only two main roads
that connect two Colombian cities (Quibdo
in the
Choco Department and Buenaventura in the
Valle del Cauca Department) with the center of
the country; the rest of this area is essentially
roadless. The southern and northern ends of this
well-preserved forest, where more deforestation
occurs, are characterized with higher road densi-
ties, providing evidence of how roads are related
to deforestation. Other studies in the tropical for-
ests of South America have also related road
development to increase in forest loss (Klein-
schroth and Healey 2017). In fact, roads facilitate
the harvest of forest products, hence opening
well-preserved forests to increased forest har-
vests and subsequent environmental changes
(Laurance and Useche 2009).
Average annual rainfall was negatively and
indirectly related to deforestation hotspots. The
13 March 2019 ❖ Volume 10(3) ❖ Article e02648

rainiest place in the world corresponds to the belt
of well-preserved forest of the CGE where the
average annual rainfall ranges between 8000 and
13,000 mm (Poveda and Mesa 2000). This
amount of rainfall on thin soils with low nutrient
levels results in non-productive soils not well sui-
ted for farming. The interaction between high
rainfall, non-productive soils, and low accessibil-
ity (lack of roads) results in a reduction of defor-
estation events, explaining the negative
relationship between rainfall and deforestation
hotspots. High rainfall has been related to a
reduction in local deforestation in the Bolivian
Amazon, where the expansion of mechanized
agriculture occurs mainly in response to access
to export markets, fertile soil, and intermediate
rainfall conditions (Mueller et al. 2012). Like-
wise, the negative and indirect relationship
between annual temperature and deforestation
hotspots in our study area could occur due to the
limitations imposed on farming activities by high
temperatures, high rainfall, and soil degradation
of the northern and southern portions of the
CGE. Also, topographic slope was negatively
and indirectly correlated to hotspots of deforesta-
tion; some scholars have found that places with
abrupt topography tend to not be deforested in
tropical areas due to the physical restrictions on
the establishment of agriculture and grazing
operations, as well as difficult access to markets
for agricultural products (Coblentz and Keating
2008, Fagua et al. 2013, Sandel and Svenning
2013).
Our results show that Gross Domestic Product
(GDP) at the municipality level was negatively
and indirectly related to deforestation hotspots.
Similar negative relations between deforestation
and national GDP have been documented on the
islands of Cuba, Hispaniola, Puerto Rico, Barba-
dos, Saint Kitts and Nevis, and Grenada and
have been attributed to the improve of forest pro-
tection as well as the agriculture decline and the
rural population migration (Grau et al. 2003,


Helmer et al. 2008, Alvarez-Berr
ıos et al. 2013,
Newman et al. 2018). Further, forest loss has
been found to decrease with increases in national
GDPs in the protected areas of 56 countries over
four continents, indicating that the effectiveness
of environmental protection (creation and/or
enforcement of environmental laws and regula-
tions) improves with better national economies
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FAGUA ET AL.
because more resources can be invested in forest
protection (Spracklen et al. 2015, Baynham-Herd
et al. 2018). We also found that the municipalities
with high cluster of deforestation hotspots are
also some of the poorest municipalities of the
three developing countries that share the CGE.
Thus, these municipalities have weak local gov-
ernance and environmental institutions with few
resources invested in environment conservation
(DANE 2015, INEC-Ecuador 2015, MEF 2017 and
WBG 2017). Consequently, forest protection is
weak supporting the statement of a negative
relationship between municipal GDPs and defor-
estation hotspots. This ineffective forest protec-
tion in poor municipalities is a concern for the
conservation of CGE biodiversity since large
extensions of poor municipalities are in the most
well-preserved forest and/or within protected
areas (national forests, indigenous or black com-
munity reservations, national parks, etc.). These
areas include the entire Panamanian CGE
(15,335 km2), the area along the pacific coast of
Colombia (58,343 km2), and four national parks
in Ecuador (669 km2). Additionally, we observed
that economies of the municipalities with signifi-
cant clustering of deforestation hotspots in Ecua-
dor and Colombia are based totally on extraction
of natural resources (farming, mining, logging,
principally), increasing pressure on forest cover.
Not one municipality in Panam
a presented sig-
nificant clustering of deforestation hotspots indi-
cating lower pressure on forest cover. Panam
a
has the lowest corruption (Transparency Interna-
tional 2015), the highest per capita GDP (WBG
2017), and some Panamanian municipalities in
the CGE obtain significant resources from eco-
tourism (Brown 2007, Klytchnikova and Dorosh
2009, Mapes 2009).
Reforestation
The Bayesian SEM of reforestation across the
CGE showed that armed conflict had the highest
indirect effect on reforestation hotspots. Colom-
bia occupies 75.6% of the CGE, and this country
has suffered internal armed conflicts that have
been especially strong in several areas of its por-
tion of the CGE. The Colombian municipalities
where reforestation hotspots were significantly
clustered (departments of Nari~ no, Choco, Antio-
quia, Cordoba) were zones strongly disputed
among different armed groups, such as guerrillas
14 March 2019 ❖ Volume 10(3) ❖ Article e02648

(FARC and ELN), paramilitaries, drug dealers,
and the Colombian army (CPDH 2006, MOE
2008, Barreto 2009, FIP USAID OIM 2014). These
violent confrontations have resulted in lands
abandoned by farmers, and subsequently pro-
ducing significant regrowth of secondary vegeta-
tion. The relationship between armed conflict
and growth of secondary vegetation in Colombia
has also been assessed at the national level (San-
chez-Cuervo and Aide 2013).
Our results also found that several municipali-
ties in Ecuador had a significant clustering of
reforestation hotspots. Ecuador is a country
without armed conflicts, suggesting that other
indirect drivers promote LULC transitions from
farming to secondary vegetation. Colombian and
Ecuadorian agricultural sectors declined during
the first ten years of the current century, thus
reducing the cultivated (grassland, crops, palm)
area and allowing for the growth of secondary
vegetation (BCE 2010a, b, Minsalud 2011, Mar-
rugo 2013). This agricultural crisis could have
increased the population migration from rural
areas to urban areas reducing the workforce in
the farming areas (Grau and Aide 2008,
Rodr
ıguez Vignoli 2008), which could have
reduced agricultural land use and increased sec-
ondary vegetation. Further, the Colombian and
Ecuadorian CGE have been affected by massive
floods during La Ni~ na years, which were espe-
cially strong in 2010–2011. Consequently, many
farming areas were lost allowing for secondary
vegetation growth (IGAC 2011, SGR 2014). We
also found that higher municipal GDP was
related indirectly to reforestation hotspots. In
regions like the Ecuadorian CGE, higher GDPs
could indicate technical improvements in agri-
culture and cattle production, resulting in inten-
sification of these land uses on more productive
lands and a reduction of agricultural land use on
less productive lands where secondary vegeta-
tion could grow.
There is no evidence, however, that reforesta-
tion will lead to new mature forests. In fact,
reforestation generated by land abandonment
represents earlier forest seral stages that will be
deforested before they can become mature for-
ests (Fagua and Ramsey 2019). When famers are
displaced during violent confrontations in
Colombia, other landowners supported by
armed groups appropriate their lands, and in
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FAGUA ET AL.
subsequent years remove regenerated natural
vegetation to reestablish agricultural activities
(PNUD 2011, Chaparro 2017). Also, when agri-
cultural sectors declined in Colombia and Ecua-
dor, farmers did not abandon their lands (since
land possession is a main source of economic
and political power in these countries), and for-
est regrowth is removed during periods of bet-
ter agricultural return (Flo
rez et al. 2012, Sierra
2013).
Our Bayesian SEM also showed that tempera-
ture was indirectly and negatively related to
reforestation hotspots. In areas of tropical rain
forests, higher temperature (Clark et al. 2003,
Anderson et al. 2006) and higher water supply


(Alvarez-D
avila et al. 2017) at the same time
increased the growth of natural vegetation and
reforestation, depending of the proximity of for-
est vegetation (Guariguata and Ostertag 2001).
Forest vegetation is well adapted to the low
nutrients and acid pH of the thin soils of these
tropical areas; these soil characteristics are also
the main restrictions for the establishment of pro-
ductive crops. The absence of one of these three
conditions (high temperature, high rain, and
proximity of forest vegetation) reduces or, even,
hampers the regrowth of natural vegetation. The
southern and northern sides of the CGE have
higher annual temperatures but lower annual
rainfall, reducing secondary vegetation growth
and thus explaining the negative relationship
between temperature and forest.
CONCLUSIONS
Our spatial analysis of LULC maps showed
that deforestation and reforestation transitions
were spatially clustered across the CGE forming
hotspots. These hotspots were clustered around
municipalities which provided many of the
direct and indirect drivers of LULC change.
While our analysis focused on the municipality
level, our results indicate that drivers related to
other political or ecological subdivisions of the
landscape (country boundaries, sub-ecoregions,
etc.) at different scales can be similarly assessed.
The challenge to use these type of models is the
availability of homogeneous information at a
consistent temporal and spatial scale. By using a
consistent LULC map generated yearly using
MODIS imagery, we were able to relate
15 March 2019 ❖ Volume 10(3) ❖ Article e02648

deforestation and reforestation hotspots to direct
and indirect drivers. We show that increases in
population growth and density of roads were the
primary indirect drivers related to deforestation
hotspots, whereas indirect drivers that limited
access to forested lands, such as topographic
slope and climate, were negatively related to
these hotspots. Where reforestation is concerned,
we identified three indirect drivers that were
positively related to hotspots. These drivers
included armed conflicts, Gross Domestic Pro-
duct, and average annual rainfall.
ACKNOWLEDGMENTS
We acknowledge COLCIENCIAS-Colombia (529-
2011) and Fulbright-US for supporting this research,
Ecology Center and Remote Sensing/GIS lab of Utah
State University-US for providing additional financial
and technical support, IDEAM-Colombia and IGAC-
Colombia for providing data and technical support,
DigitalGlobe Foundation for providing the high spatial
resolution imagery, and IDEA WILD for providing
equipment.
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DATA ACCESSIBILITY
The land cover maps of the Choco
-Darien Global Ecoregion (South America) for 2002 and 2015, the map of
deforestation hotspots 2002-2015, and the map of reforestation hotspots 2002-2015 are available at: https://zenod
o.org/record/2559673#.XF0DJDNKiM8

SUPPORTING INFORMATION
Additional Supporting Information may be found online at: http://onlinelibrary.wiley.com/doi/10.1002/ecs2.
2648/full

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