Open Agriculture.
2018; 3: 273–283
Review Article
Roberto Quiroz*, David A. Ramírez, Jürgen Kroschel, Jorge Andrade-Piedra, Carolina Barreda,
Bruno Condori, Victor Mares, Philippe Monneveux, Willmer Perez
Impact of climate change on the potato crop and
biodiversity in its center of origin
https://doi.org/10.1515/opag-2018-0029
received February 26, 2018; accepted June 21, 2018
Abstract: The Andean region is the most important center
of potato diversity in the world. The global warming trend
which has taken place since the 1950s, that is 2-3 times the
reported global warming and the continuous presence of
extreme events makes this region a live laboratory to study
the impact of climate change. In this review, we first present
the current knowledge on climate change in the Andes, as
compared to changes in other mountain areas, and the globe
in general. Then, the review describes the ecophysiological
strategies to cope and adapt to changes in atmospheric CO2
levels, temperature and soil water availability. As climate
change also has a significant effect on the magnitude and
frequency of the incidence of pests and diseases, the current
knowledge of the dynamics of vectors in the Andean region
is discussed. The use of modeling techniques to describe
changes in the range expansion and number of insect pest
generations per year as affected by increases in temperature
is also presented. Finally, the review deals with the use
of crop modeling to analyze the likely impact of projected
climate scenarios on potato yield and tuber initiation.
Keywords: Andes, crop modeling, global warming, pest,
Solanum spp.
*Corresponding author: Roberto Quiroz, Tropical Agricultural
Research and Higher Education Center, CATIE, Cartago Turrialba
30501, Costa Rica E-mail: roberto.quiroz@catie.ac.cr
David A. Ramírez, Jorge Andrade-Piedra, Carolina Barreda, Victor
Mares, Philippe Monneveux, Willmer Perez, International Potato
Center, Headquarters P.O. Box 1558 Lima 12, Peru
David A. Ramírez, Gansu Key Laboratories of Arid and Crop Science,
Crop Genetic and Germplasm Enhancement, Agronomy College,
Gansu Agricultural University, Lanzhou 730070, China
Jürgen Kroschel, International Potato Center, Asia Potato Program,
P.O. Box 110012 New Delhi, India
Bruno Condori, Adaptive Cropping Systems Laboratory, USDA-ARS,
Beltsville, MD 20705-2350, USA
Roberto Quiroz, Tropical Agricultural Research and Higher Education
Center, CATIE, Cartago Turrialba 30501, Costa Rica
Open Access. © 2018 Roberto Quiroz et al., published by De Gruyter.
NonCommercial-NoDerivs 4.0 License.
1 Introduction
Potatoes (Solanum tuberosum L. and other Solanum spp.)
are an important food crop (Hijmans 2001; Lutaladio and
Castaldi, 2009) that originated in the Andes (Spooner et al.
2005). They are currently produced in over 149 countries
in areas from sea level up to 4,200 m.above.sea.level
(m.a.s.l). in the tropical highlands and throughout the
temperate zone. Potatoes feed more than a billion people
worldwide, from a global crop production close to 400
million metric tons (FAOSTAT 2017) and it is critical for
the food security of people across South America, Africa,
and Asia, including Central Asia. Since the early 1960s,
the increase in potato cultivation area has overtaken all
other food crops in developing countries, which account
for more than half of global potato production (Devaux et
al. 2014).
The Andean region is the most important center of
potato diversity in the world. There, more than 4,000
varieties and landraces which can be found, with local
farmers playing an important role in conservation through
the cultivation of around 3,000 landraces (Spooner et
al. 2014). High climatic variability, which has been a
constant feature in this region, might have contributed to
producing potato diversity and the diversification process
is also associated with the domestication and selection
effects by ancient Andean farmers. There is evidence that
climate change is exacerbating the occurrence of extreme
climate events in the region (see Section 2), a process that
could have important effects on potatoes, both as a crop
and as a wealth of in situ biodiversity.
In this review, the current knowledge on climate
change in the Andes is presented, as compared to changes
in other mountain areas, and the globe in general. Then,
the review describes the effects caused by changes in
atmospheric CO2 levels, temperature and soil water
availability on the potato plant. It also describes the
ecophysiological strategies used by the potato plant to
cope and adapt to non-lethal adverse climatic conditions.
As climate change also has a significant effect on the
This work is licensed under the Creative Commons Attribution-
Unauthenticated
Download Date | 8/15/18 5:25 AM
274   R. Quiroz, et al.
magnitude and frequency of the incidence of pests and
diseases, the review also discusses the current knowledge
of the dynamics of vectors in the Andean region as they
respond to climate change. The use of modeling techniques
to describe changes in the range expansion and number of
insect pest generations per year, as affected by increases
in temperature, is also presented. Finally, the review deals
with the use of crop modeling to analyze the likely impact
of projected climate scenarios on potato yield and tuber
initiation.
2 Climate change in the Andes
The Andes are the largest and highest mountain range
in the tropics and their land mass, glaciers and altitude
modify the atmospheric circulation and the climate of
the region (Vuille et al. 2008a; Rabatel et al. 2013). The
Andean region is very sensitive to the significant warming
evidenced in recent years (Schauwecker et al. 2014; Vuille
et al. 2015) that is affecting the hydrology, land uses and
the agro-ecosystems of the region. The average warming
of the Andes between 1939 and 2006 was around 0.7oC
(Vuille et al. 2008a) with variations conditioned by
altitude and slopes (Vuille et al. 2003). Warming trends
are higher at high altitudes reaching values of over 0.3°C.
decade-1, in spite of a cooling due to a recent hiatus in
warming, affecting mainly the coastal areas (Vuille et al.
2015). This situation differs from other mountain ranges
in other latitudes such as the European Alps and Tibet.
The warming trend is projected to continue in the coming
decades (Thibeault et al. 2010), but with differences as
functions of geographic position and altitude are yet to be
clearly determined.
In contrast to the warming characteristics, there is no
clear pattern of decreasing, or increasing, precipitation
in the region (Haylock et al. 2006; Morin 2011). One
limiting factor for a better understanding of precipitation
trends in such complex terrain is the low density of
meteorological stations with reliable records for at least
30 years. To solve this limitation, modeling techniques
based on mathematical downscaling of precipitation
signals (Duffaut et al. 2017) and the use of remote sensing
products (Mohr et al. 2014) have been tested. Uncertainty
in precipitation patterns is recurrent in projected climate
change scenarios and decreased levels of seasonal
precipitation but more intense events are expected during
the austral summer (Thibeault et al. 2011). Significant
events such as the South American Monsoon (Carvalho et
al. 2012) and El Niño Southern Oscilation (Garreaud et al.
2003; Vuille et al. 2008b) that regulate the precipitation
in the region will be exacerbated by and feed into climate
change (IPCC 2013). Changes in both temperature and
precipitation levels and patterns of occurrence are key
factors in the assessments of likely effects of climate
change on Andean agriculture. It is thus critical to analyze
the expected behavior of these components of regional
climate systems in order to evaluate the potential impacts
of climate change on the potato crop and the biodiversity
of potato in the region. It is also important for the
identification of suitable adaptation measures that would
contribute to the resilience and productivity of potato-
based farming systems in the Andean region (Rolando et
al. 2017).
3 Responses of the potato crop to
Climate Changes in the Andes
3.1 Interactive effects of increased atmos-
pheric CO2, high temperature and water
restriction
An important question to solve is how atmospheric CO2
increases will affect potato responses under different
drought scenarios (Finnan et al. 2005). Some studies
assessing CO2 increase through free-air CO2 enrichment
systems (FACE) have reported a tuber yield increase in
potatoes under no water restriction (Finnan et al. 2005;
Kimball 2016). The analysis of the interaction between
CO2 increase and warmer temperatures is still an on-going
research topic in field crops under FACE systems (Kimball
2016). At present, only modeling is providing some
clues about their combined effect on potato growth and
development. Thus, in India, Singh et al. (2013) reported
that elevated CO2 (550 ppm) produced a beneficial effect in
tuber yield (+11.1%) up to a temperature increase of +1°C but
when the temperature increment reached a threshold of
+3°C the elevated CO2 caused a reduction of 13.7 % in tuber
yield. On the other hand, Fleisher et al. (2016) pointed out
that temperature is a more important source of uncertainty
in potato crop models than CO2 and water. In this case, the
median ensemble from nine potato growth models used to
simulate a pessimistic climate scenario (with upper limits
of 720 ppm of CO2, +9°C of temperature increase and -30%
of water decline from baseline) showed a dramatic tuber
yield reduction. More experimental evidence is necessary
to understand the effect of CO2 increase on tuber yield
under water restriction scenarios and mainly under warm
temperature conditions, and to determine the threshold
of severity in those stresses at which the interaction turns
Unauthenticated
Download Date | 8/15/18 5:25 AM
 Impact of climate change on the potato crop and biodiversity in its center of origin
harmful. That evidence is important not only to improve
the simulations, but also to understand how acclimation
or phenotypic plasticity will influence the resilience of the
potato crop under climate change scenarios.
Drought effects on crops depend on developmental
or phenological timing, duration and intensity of water
restriction (Jefferies 1995). The variation in these three
factors and their combination will condition the response
of the plant to drought. Knowledge of the physiological
responses is important for the selection of plant traits that
help the crop to cope with water shortages (Tardieu 2012).
Thus, the breeding of potato varieties able to positively
respond to water shortages and to water management for
improving yield and crop quality is based on the knowledge
and understanding of plant and crop physiology (Medici
et al. 2014). For instance, tuber formation in potatoes has
been described as a response to foliar stress (Burton 1981)
an effect that implies that some level of water restriction
is necessary to induce tuberization. This assertion is
supported by the fact that there is foliage cover reduction,
caused by induced water restriction, increases tuber
yield per unit water volume as a consequence of reduced
transpiration and a concomitant maintenance in carbon
assimilation associated with the canopy´s reduced self-
shading (Shahnazari et al. 2007). It has also been reported
that early mild water restriction during crop growth and
development can enhance water stress resistance and
tuber yield (Xu et al. 2011; Yactayo et al. 2013). Trans-
generational water stress memory imprinting in potato
(i.e. water stress resistance carried out by tuber seeds
to the following crop) has been observed (Ramírez et al.
2015). On the other hand, severe water restriction can
negatively affect tuber yield if it occurs just before or
during tuber initiation (Monneveux et al. 2013) or bulking
(Vanloon 1981). In a study of different water restriction
and CO2 levels including early developmental timing, long
duration and different water restriction intensities (90,
75, 50, 25 and 10 % reduction of daily irrigation), Fleisher
et al. (2008 a, 2008b) observed that an increase of CO2
(740 ppm) counteracted water deficit through enhanced
carbon allocation to tubers, increasing yield and water
use efficiency (WUE). In another drought scenario,
Fleisher et al. (2013) found that 800 ppm CO2 increased
tuber yield and WUE under water restriction, brought
about by suspending irrigation for 11 to 16 days at early
and early + late developmental stages, affected plant
water potential from -1.1 to -1.65 MPa. Results suggested
that CO2 increase at the cellular level counteracted water
restriction by: i) the reduction in the production of proline,
a compound involved in osmotic regulation leading to
water conservation by stomatal closure (Barnaby et al.
 275
2015) and, ii) the reduced accumulation of compounds
synthesized during water restriction, with a potentially
positive effect on tuber quality (Yang et al. 2015). Although
the CO2 increase alleviated the effects of water restriction
in the above two cases, it is still necessary to test the
effect of more severe drought which in other crops was
not counteracted by the beneficial effect of CO2 increase
(Sicher and Barnaby 2012).
3.2 Responses of potato diseases
Besides the direct effects on the ontogeny and physiology
of the crop, climate change is also expected to impact
on life cycles of crop’s pathogens. As stated by Kapsa
(2008), variations in climate regimes might change: 1)
the physiology of pathogen/insects and host plants; 2)
the host plant resistance to infection/infestation; 3) the
critical temperature and related infection thresholds,
affecting the forecasting of pathogen occurrences, the
efficacy of plant protection and yield ranges. In general,
higher temperatures in areas located in higher latitudes
will lead to longer growing seasons and higher yields,
but also an increase on pest and pathogen pressure, as
a result of more sources of initial inoculum (e.g. infected
tubers that survive winter), more vector activity, higher
multiplication rates, and more generations per season
(Haverkort and Verhagen 2008).  
Major climate change factors likely to influence plant
disease severity and spread include elevated CO2, heavy
and unseasonal rains, higher humidity, drought, cyclones
and hurricanes, and elevated temperature (Luck et al.
2011). Potato late blight (Phytophthora infestans) will still
be a major concern for potato production (Kapsa 2008;
Haverkort and Verhagen 2008). However, due to regional
variation, the overall global effect of climate change on late
blight is not clear (Sparks et al. 2014), The impact at high
altitudes is expected to range from areas where late blight
is becoming a problem, such as the high Andes (Perez et
al., 2010; Perez et al., 2016), to areas where it is forecasted
to be less severe, such as the Southeast Asian Highlands
of Indonesia. Where late blight is to increase, the infection
potential might become greater due to genetic changes in
pathogen population and higher infection pressure. In
addition, late blight might occur earlier in the season due
to additional sources of inoculum in the soil – e.g. infected
tubers from volunteer potato plants –  and/or oospores
surviving in soil at low temperature. On the other hand,
elevated CO2 might activate defense response mechanisms
such as pathogenesis related proteins e.g. B-1,3, glucanase
and osmotin (Plessl et al. 2007).  
Unauthenticated
Download Date | 8/15/18 5:25 AM
276   R. Quiroz, et al.
The synchronization of high humidity and/or
high temperature with pathogen-host plant features
may enable the presence of other fungus (e.g. early
blight caused by Alternaria solani, black dot caused by
Colletotrichum coccodes) and bacterial diseases (e.g.
Pectobacterium chrysanthemi, Ralstonia solanacearum)
especially important for seed production (Kapsa 2008; and
Haverkort and Verhagen 2008). In Ontario, Boland et al.
(2004) anticipated that canker (Rhizoctonia solani), early
blight, late blight, pink rot (Phytophthora erythroseptica),
blackleg (Pectobacterium spp. and Dickeya spp.), and ring
rot (Clavibacter michiganensis) would reduce as a result
of climate change, while wilt (Verticillium spp.), common
scab (Streptomyces scabies), and potato leafroll virus
(PLRV) would increase. In South Africa, van der Waals et
al. (2013) predicted that soft rot and blackleg, root-knot
nematodes (Meloidogyne javanica and M. incognita) and
potato virus Y (PVY) and PLRV will increase in all areas
and seasons; early blight and brown spot (Alternaria
alternata) are likely to increase or will remain unchanged,
depending on the region and season; and late blight
will decrease in most of the cropping systems modelled.
Specific studies such as those from Boland et al. (2004)
and van der Waals et al. (2013) help to allocate resources
for research on key diseases that will likely increase in
importance as a result of climate change.    through provision of new opportunities for dispersal and
New tools and methods are being sought and indeed
scientists are already improving their skills through
advances in technologies for high-throughput analysis of
gene expression, experimental data to begin synthesizing
the effects of climate variables on infection rates, models
of plant diseases (Newbery et al. 2016), and scaling tools
at ecosystems levels (Garret et al. 2006). Notwithstanding,
with the current level of knowledge it is not feasible
to estimate expected outcomes of host-pathogens
interactions under different temperature, humidity and
CO2 combinations. Moreover, the difficulty increases when
the impact assessment must be conducted at different
spatial scales.   insect pest problem at altitudes above 3,800 m.a.s.l. In
3.3 Responses of potatoes to insect pests
Insects are poikilothermic organisms whose development
depends on the temperature to which they are exposed in
the environment. Hence, it is expected that agricultural
insect pests of potato will respond to climate change by
expanding their geographical range of distribution and
by increasing population densities in the Andean region
which will lead to greater crop and post-harvest losses
(Kroschel et al. 2013). Potato production systems of tropical
countries are highly susceptible to pest infestations due
to often year-round favorable climatic conditions for
pest population growth and host plant availability. Even
smaller changes in temperature predicted for tropical
regions compared to temperate regions will have stronger
consequences on pest development due to already higher
existing metabolism rates of organisms such as insects
(Dillon et al. 2010; Govindasamy et al. 2003).
Besides direct effects of temperature on pests,
temperature will also affect the crop phenology and
modifications can lead to possible changes in plant-
insect interactions and to new risks of pest infestations
(Chakraborty and Newton 2011). Elevated CO2 levels have
been shown to affect the crop defense systems leading
to an increased vulnerability to insect pests (Van Asch et
al. 2007). Increases in the frequency of extremes events
(temperature, precipitations, etc.) as well as differences
in the sensitivity of higher tropical levels to climatic
variability can disrupt the synchrony between the growth,
development and reproduction of biological control
agents and their hosts (pests), leading to interferences
of both natural and implemented biological control
processes (Voigt et al. 2003). Further, changes in the rate
and intensity of extreme climatic events will perturb
ecosystems and increase their vulnerability to invasions
growth of insect species (Masters and Norgrove 2010).
Long-term drought may also lead to reduced crop growth
and health thereby increasing their susceptibility to insect
pests (Magan et al. 2011).
In the Andean highlands, potato production is
severely constrained by Andean potato weevils especially
of species of the genus Premnotrypes. P. suturicallus
(Kuschel) occurs in central Peru, P. vorax (Hustache) in
northern Peru, Ecuador, Colombia and Venezuela, and P.
latithorax (Pierce) in southern Peru and Bolivia (Alcazar
and Cisneros 1999). Andean potato weevils occur between
2,800 to 4,700 m.a.s.l. and represent the only major biotic
inter Andean valleys, other important pests are the potato
tuber moth, Phthorimaea operculella (Zeller) and the
Andean potato tuber moth, Symmetrischema tangolias
(Gyen.), among other minor species (Kroschel et al. 2012).
All species are native to this region. P. operculella has
become a very important invasive pest globally at much
greater proportions than S. tangolias. The Guatemalan
potato tuber moth, Tecia solanivora (Polvolny), has
invaded the Andean region in Venezuela, Colombia
and Ecuador, but has not yet been reported from Peru
(Kroschel and Schaub 2012). The leafminer fly, Liriomyza
huidobrensis (Blanchard), can be a serious potato pest in
Unauthenticated
Download Date | 8/15/18 5:25 AM
 Impact of climate change on the potato crop and biodiversity in its center of origin
the highlands of Ecuador. It also occurs in the highlands of
Peru but under these conditions it hardly affects potatoes.
In contrast, it is the major pest in Peruvian coastal potato
production regions, at altitudes below 500 m (Mujica
2016). Potato pests have different optimum temperature
ranges for development and reproduction (Kroschel
et al. 2016; Mujica 2016; Sporleder et al. 2016). A rise in
temperature due to climate change may therefore either
increase or decrease pest development rates and related
crop damages depending on the pest species optimum
temperature range. We used pest phenology modeling
and Geographic Information Systems (GIS) risk mapping
tools for some of the species to better understand potential
changes in future pest abundance and infestation in the
Andean region (see section 4.3).
4 Modeling the impact of climatic
variations in the Andes
To assess the potential consequences of climate
change, modeling tools are used to upscale results
from experimental field research to regions (landscape,
countries, continents, and global). In most cases a
potato growth model is fed with present and predicted
climate data and results of the models are compared
(Olesen and Bindi 2002; Hijmans 2003). Multiple years
baseline and variations (above and below the baseline)
in CO2, temperature and rainfall that represent the
range of potential 21st century climate changes are used
for simulation runs in model inter-comparison studies
(Rosenzweig et al. 2013; Fleisher et al. 2016). The following
sections discuss the work in which models were fed with
either present or predicted future climate data or with
multiple year baseline and variations data to model both
direct and indirect effects of climate variations on potato
cropping in the Andes.
4.1 Modeling potato yields to assess
the likely effect of changes in tempera-
ture, atmospheric CO2 levels and water
availability
Most potato growth models are limited to growth analyses
conducted for S. tuberosum ssp. tuberosum. However, in
the Andes, a region where farmers face uncertain climate
every cropping season, the genetic diversity is wide.
Farmers cope with constant droughts, frosts, hailstorms
or excess rainfall by planting an assortment of potato
 277
species and landraces. Models capable of simulating
the expected behavior of a wide genetic diversity under
variable climate are needed to better assess adaptation
options for future climates. International Potato Center
(CIP) and partners have developed the Solanum model
(Raymundo et al. 2014; Quiroz et al. 2017; Raymundo et al.
2017, among others) suited with crop growth parameters
for Andean potato species such as S. juzepckzuki – known
as bitter potatoes, S. ajanhuiri, S. tuberosum subspecies
andigenum and tuberosum and their hybrids. This model
was used to assess the response of different varieties and
landraces to variation in climatic conditions in the Andes.
Under good husbandry in experimental conditions,
Andean varieties and hybrids used in the region produced
fresh yields above 45 Mg.ha-1 under rain-fed conditions
(Condori et al. 2010, Condori et al. 2014). When field
capacity conditions are simulated, tuber yields increase
(Figure 1). High yielding varieties tended to also be
precocious and fast-bulking. High values of maximum
canopy cover (MCC) – associated with higher light
interception - were highly correlated to yields in the high
Andes, whereas relative low MCC values were explained
by temperatures below optimal range (Condori et al. 2010,
Condori et al. 2014). The warming trend reported for the
western slope of the Andes (described in Section 2) seems
to have a direct effect on the different taxa cultivated in
the region. Simulations using growth parameters for 10
different taxa (including S. juzepczukii, S. tuberosum ssp.
andigenum, S. phureja, and S. acaule) with climate data for
three altitudes (3,200, 3,920, and 4,250 m.a.s.l.) for over 40
years, showed that at lower and middle altitudes there is a
steady decline in potential productivity – associated with
the warming trend - for all varieties modeled. Potential
productivity increased with altitude, as temperature
becomes more appropriate for non-bitter potato
production. The highest location has traditionally been
considered as non-suitable for potato varieties but the
model showed that under current temperatures the bitter
tuber producing species S. juzepczukii can be cultivated, a
fact that can be verified in some farmers’ fields in Central
Peru (de Haan et al. 2010)
The simulated responses of S. tuberosum ssp.
tuberosum to changes in CO2 levels (360, 450, 540, 630,
720 ppm), temperature (-3, 0, 3, 6, and 9oC below/above
baseline 24-h maximum/minimum air temperatures) and
water availability (-30%, 0, +30% decreases/increases in
daily precipitation from baseline) were reported by Fleisher
et al. (2016). The analysis for the Andes was conducted for
the conditions of Chinoli, Bolivia, at a latitude of -19.63,
longitude -65.37 and altitude of 3450 m.a.s.l.. The mean
temperature was 16oC and the cumulated precipitation
Unauthenticated
Download Date | 8/15/18 5:25 AM
278   R. Quiroz, et al.

Figure 1: Air temperature effects on dry tuber yield (top) and tuber initiation (bottom) based on 30-year simulations of potato growth in the
Andean Highlands under rainfed conditions for different genotypes; AJA: S. ajanhuiri, var. Ajanhuiri; AND: S. tuberosum ssp. andigenum,
var. Waycha; HYB: S. tub ssp. tuberosum X S. tub ssp. andigenum, var. Sajama and TUB: S. tuberosum ssp. tuberosum, var. Alpha. Results at
irrigated conditions were represented by open diamonds

was 340 mm. Using the median of simulated responses
of nine models over 30 years and varying levels of the
three climatic factors it was shown that dry tuber yield
in a S. ssp. tuberosum cv. Desiree could be incremented
to about 10% per 100 ppm of CO2 increased. On the other
hand, warming scenarios showed a 4% decrease in dry
yield per oC incremented. Reduction in 30% of baseline
rainfall reduced yield by 4.9% whereas a 30% increment
in rainfall augmented yield in 2.5% over the baseline
scenario. Overall, higher temperatures tended to offset
the CO2 rising effect and increments in rainfall seemed to
have a positive effect in rain-fed areas. This analysis was
expanded to native varieties cultivated in the high Andes
but using only the Solanum model. The impact of higher CO2
concentrations could not be included, since the response
of these varieties to enriched CO2 environments is not as
well understood as for S. tuberosum ssp. tuberosum. No
significant relative deviations from the S. tuberosum ssp.
tuberosum was found either for changes in precipitation
or for temperature. The response to temperature (Figure 1
top panel) was 4.5, 4.6, 4.7, and 4.9% per oC incremented
for S. tuberosum ssp. tuberosum, tuberosum x andigenum
hybrid, S. ajanhuiri and S. andigenum, respectively. The
values for the S. tuberosum ssp. tuberosum cv. Alpha
estimated by the Solanum model alone were 0.5% per
o
C higher than the ensemble of nine models reported by
Fleisher et al. (2016) for cv. Desiree. Earlier tuber initiation
was induced at temperatures above the baseline (Figure

Unauthenticated
Download Date | 8/15/18 5:25 AM
 Impact of climate change on the potato crop and biodiversity in its center of origin
1) with rates varying from 1.3 to 1.9 days oC-1 and the most
affected species was the tuberosum x andigenum hybrid.
Condori et al., 2010 found that the average thermal time
at tuber initiation for tuberosum x andigenum hybrids
was 530oC-days whereas for andigeum and tuberosum,
they were 453 and 259oC-day, respectively. Bulking rates
reduction (data not shown) ranged from 2.6 to 3.1 kg.oC-
1
, with S. ajanhuiri being the most affected species. It is
worth mentioning that for scenarios where temperatures
were below the 30-years baseline – those in the region
represent by planting at higher altitudes - % dry yield
changes per oC were 11, -6, +7 and +3, for S. tuberosum ssp.
tuberosum, tuberosum x andigenum hybrid, S. ajanhuiri
and S. tuberosum ssp. andigenum, respectively. That is, in
terms of yield, the current strategy to plant native varieties
in higher grounds seems to be an option, since both S.
tuberosum ssp. andigenum and S. ajanhuiri will benefit
from cooler environments. In Central Peru, de Haan et
al. (2010) showed that the highest level of infraspecific
diversity for native-bitter cultivars – with 4.1 cultivars per
field – were concentrated between 4050 and 4150 m.a.s.l.
This will not be the case – in the short-to-medium-term
for S. tuberosum ssp. tuberosum and its hybrid tuberosum
x andigenum.
4.2 Modeling the spread and impact of late
blight under contemporary and projected
climate
Geo-spatial modeling of late blight began over a decade
ago when Hijmans et al. (2000) incorporated two disease
forecasting models in a GIS.  This produced a disease risk
map in which potential risk was estimated by the number
of fungicide sprays needed for disease management. In
fact, this variable has been useful to researchers at CIP as
it is easily converted to economic value, thus facilitating
ex ante impact assessments (Forbes and Lizarraga
2010). The forecasting models also produce outputs
for resistant and susceptible potato varieties, allowing
general evaluations of the value of host resistance across
a region. Sparks et al. (2011) developed a meta-model of
the previous approach using general additive models. The
new model uses input data of lower temporal resolution
(daily and even monthly), which has greatly facilitated
risk and impact assessment.  This model was used for
future risk assessments for the Andean Highlands, Indo-
Gangetic Plain and Himalayan Highlands, Southeast
Asian Highlands, Ethiopian Highlands, and Lake Kivu
Highlands in Sub-Saharan Africa (Sparks et al. 2014).
On average, it was found that late blight risk is likely to
 279
increase until the year 2020, and then decrease after
2050, mainly due to a shift in the growing season (from
June to May in the Northern Hemisphere) to avoid high
temperatures, or because the temperature will be higher
than the optimal for P. infestans infection (Sparks et al.,
2014). At the ecosystem level, there were major differences,
from a large increase in risk in the Lake Kivu and Andean
Highlands, to a decreased risk in the Southeast Asian
Highlands. As pointed out by Shakya et al. (2015), the
model developed by Sparks et al. (2014) did not consider
diurnal temperature amplitudes, a factor that could affect
late blight epidemics, and therefore should be taken into
account in the next versions of the model. 
Concurrent increments in temperature and relative
humidity pose interesting challenges for potato
production. On the one hand, higher temperatures in cool
highlands might be positive for production, but the likely
proliferation of pests and diseases might suppress the
potentially positive effect of warming temperatures. For
example, as predicted by Giraldo et al. (2010) and Sparks
et al. (2011), late blight is becoming an important threat
in highland production areas. Farmers have traditionally
used highlands as a late blight-free area, planting more
susceptible potato varieties because of low temperatures
inhibiting the disease. These low temperature areas will
certainly occur at even higher altitudes under global
warming, but could force farmers to go into more fragile
ecosystems, such as paramo and puna lands in the Andes,
with direct impact on soil organic carbon reservoirs
(Segnini et al. 2010, Segnini et al. 2011).
4.3 Modeling the range expansion and
impact of potato pests under current and
future projected climates
Phenology models are important analytical tools
for predicting, evaluating and understanding pest’s
population dynamics in agroecosystems under a variety
of environmental conditions. CIP developed the Insect
Life Cycle Modeling (ILCYM) software suitable to develop
phenology models of insect pests based on experimentally
assessed temperature-dependent life-table data, to map its
risks using three indices (Establishment Risk Index, ERI;
Generation Index, GI; and Activity Index, AI) on global and
regional scales under current and future predicted climate
change scenarios (Sporleder et al. 2014; Sporleder et al.
2013). Recently, the software has been used to develop a
“Pest Risk Atlas for Africa’ for a wide range of pest species
in different important crops including potatoes (Kroschel
et al. 2016).
Unauthenticated
Download Date | 8/15/18 5:25 AM
280   R. Quiroz, et al.

The range expansion and damage potential of P. per year (from 6-9 to 9-12 generations/year) and of its
operculella has been predicted under projected changes potential population growth (Mujica 2016).
in global temperature for the year 2050 based on Also, S. tangolias (Sporleder et al. 2016) is likely to
downscaled climate-change data of the A1B scenario expand its range with higher abundance into higher
(IPCC 2007) from the WorldClim database described by altitudes of the Andean region. A range expansion of T.
Govindasamy et al. (2003) using the pests’ temperature- solanivora (Schaub et al. 2016) into mountainous regions
dependent phenology model developed by Sporleder of the Andes of Peru and Bolivia is very likely and must
et al. (2004). According to this study, the P. operculella be considered in the risk management of this species
damage potential will progressively increase in all (Kroschel et al. 2016). Both species are further expected
regions where the pest already prevails today, with to expand their range into temperate production regions
(1) a major increase in warmer cropping regions of of the northern and southern hemisphere. Temperature-
the tropics and subtropics; (2) a range expansion in dependent phenology models for Andean potato weevils
temperate regions of the northern hemisphere in Asia, are under development. With warmer temperatures in the
North America, and Europe with a moderate increase Andean region we expect an upward move of this species
of its damage potential; and (3) a range expansion in which is well adapted to cooler climates.
tropical temperate mountainous regions with a moderate
increase of its damage potential (Kroschel et al. 2013).
Specifically, the study projected for the Andean region of
5 Conclusions
Bolivia, Ecuador, and Peru, that the pest will expand its
On-going changes in the climate is bringing about a
range by an additional 44,281 ha, 9,569 ha, and 39,646 ha
significant variation in the environmental conditions of
of the potato production area leading to a total change
the Andean region. There is already an observed warming
of establishment in these countries from 50.0% to
trend, highly dependent on geography and altitude, and
84.7%, 69.9% to 85.9%, and 42.5% to 58.5%, respectively.
an as yet not clear long-term pattern of changes in the
Likewise, climate change will increase the pest damage
seasonal and inter-annual rainfall distribution and total
potential in these countries. Under the current climatic
precipitation. In spite of prevalent uncertainties, there is
conditions P. operculella produces >4 generations in
evidence that farmers are responding to climate change by
29% of the total potato production area in the Andean
an up-ward movement of potato cropping areas into high
countries from Venezuela to Bolivia. Predictions for
altitude natural rangelands. One of the main reasons for
the year 2050 indicate that this area may potentially
this encroachment of potato cultivation into high altitude
expand to 46.6% with additional affected areas of 38,032
areas is the increased population and rate of vectors,
ha having between 4 and 6 generations, and 72,148
pest and diseases that appear to be occurring at lower
ha having severe infestations of >7–15 generations.
altitudes. An unintended consequence of the conversion
Thus, these modeling studies clearly demonstrated the
of native pastures into cropping areas could potentially
adaptation of P. operculella to warmer climates which
be the reduction of C sequestration and an increment
will lead to an increase of its population and damage
of C release into the atmosphere. This is a topic under
potential in all potato growing regions globally.
current research in the region. Significant progress in the
In comparison, the leafminer fly, L. huidobrensis, is
assessment of climate change effects on potato cultivation
much less adapted to warmer climates and climate change
in the high Andean region is being achieved with the aid
will differently affect this species. Global predictions
of modeling tools which allow the simulation of potato
clearly indicate a slight to moderate decrease in the
growth and development based on growth parameters
establishment potential of the pest in most of the tropical
that have been experimentally defined for different
and subtropical potato production areas, but still with
potato species, subspecies and varieties. Modeling is also
high pest risks. Further, a range expansion into temperate
instrumental in the analysis of the responses of potatoes
regions in the northern hemisphere in Asia, North and
to changes in CO2 levels, temperature and rainfall and
South America, and Europe, as well as into subtropical
of the impact of these factors on physiological yield
mountainous regions, is expected, with a moderate
determinants like tuber initiation and tuber bulking.
increase of its establishment and damage potential
Furthermore, modeling tools are contributing to yield
(Mujica et al. 2016). For the Mantaro valley in Peru, the
gap analysis under different current and anticipated
model predicts an expansion of the pest from currently
environmental conditions. On the other hand, geo-spatial
3,000 – 3,500 m.a.s.l. to higher altitudes of 3,800 m.a.s.l.
modeling considerably helped to (1) figure out the spread
with an essential increase of the number of generations

Unauthenticated
Download Date | 8/15/18 5:25 AM
 Impact of climate change on the potato crop and biodiversity in its center of origin
and impact of diseases under contemporary and projected
climate and (2) build disease risk maps which provide
general evaluations of the value of host resistance across
a region, thus facilitating risk and impact assessment.
Finally, pest phenology models developed with support of
the Insect Life Cycle Modeling (ILCYM) software permitted
(1) a better prediction, evaluation and understanding of
the pest’s population dynamics in agroecosystems under
a variety of environmental conditions and (2) mapping
their risks on global and regional scales including the
Andes under current and future predicted climate change
scenarios.
All these tools, taken together, are already
contributing to a better description and understanding
of the impact of climate change on potato cultivation in
its center of origin. New knowledge and understanding of
changes in pest expansion and abundance will ultimately
support the preparedness of national programs and potato
farmers to use appropriate technologies and integrated
pest management (IPM) to adapt to these changes.
Foreknowledge on how all these factors affect yield can
be useful in designing adaptation measures that will
contribute to construct knowledge-based climate resilient
potato-based farming systems in a region of high climatic
variability and change.
Conflict of interest: Authors state no conflict of interest.
References
Alcazar J., Cisneros F., Taxonomy and bionomics of the Andean
potato weevil complex: Premnotrypes spp. and related
genera. In: International Potato Center (CIP), Program Report
1997-1998, CIP, 1999
Barnaby J.Y., Fleisher D., Reddy V., Sicher R., Combined effects
of CO2 enrichment, diurnal light levels and water stress on
foliar metabolites of potato plants grown in naturally sunlit
controlled environment chambers, Physiol. Plant., 2015, 153,
243–252
Boland G.J., Melzer M.S., Hopkin A., Higgins V., Nassuth A., Climate
change and plant diseases in Ontario, Can. J. Plant Pathol.,
2004, 26, 335–350
Burton W.G., Challenges for stress physiology in potato, Am. Potato
J., 1981, 58, 3–14.
Carvalho L.V., Jones C., Posadas A.N., Quiroz R., Bookhagen B.M.V.,
Liebmann B., Precipitation characteristics of the South
American Monsoon System derived from multiple data sets, J.
Clim., 2012, 25(13), 4600-4620
Chakraborty S., Newton A.C., Climate change, plant diseases and
food security: an overview. Plant Pathol., 2011, 60(1), 2-14
Condori B., Hijmans R.J., Quiroz R., Ledent J.-F., Quantifying the
expression of potato genetic diversity in the high Andes
through growth analysis and modeling, Field Crops Res., 2010,
119(1), 135-144
 281
Condori B., Hijmans R.J., Quiroz R., Ledent J.-F., Managing potato
biodiversity to cope with frost risk in the high Andes. Plos One,
2014, 119(1), 135-144
de Haan S., Nuñez J., Bonierbale M., Ghislain M., Multilevel Agrobio-
diversity and Conservation of Andean Potatoes in Central Peru,
Mt. Res. Dev., 2010, 30(3), 222-231
Devaux A., Kromann P., Ortiz O., Potatoes for Sustainable Global
Food Security, Potato Res., 2014, 57(3),185-199
Dillon M.E., Wang G., Huey R.B., Global metabolic impacts of recent
climate warming. Nature, 2010, 467, 704–706
Duffaut L.A., Posadas A.N., Carbajal M., Quiroz R., Multifractal
downscaling of rainfall using normalized difference vegetation
index (NDVI) in the Andes plateau. PlosOne, 2017, 12(1),
e0168982
FAOSTAT, Food and agriculture data, 2017, http://www.fao.org/
faostat/en/#home
Finnan J.M., Donnelly A., Jones M.B., Burke J.I., The Effect of Elevated
Levels of Carbon Dioxide on Potato Crops, J. Crop Improv.,
2005, 13, 91–111
Fleisher D.H., Timlin D.J., Reddy V.R., Elevated carbon dioxide and
water stress effects on potato canopy gas exchange, water use,
and productivity. Agric. For. Meteorol., 2008a, 148, 1109–1122
Fleisher D.H., Timlin D.J., Reddy V.R., Interactive effects of carbon
dioxide and water stress on potato canopy growth and
development. Agron. J., 2008b, 100, 711–719
Fleisher D.H., Barnaby J., Sicher R., Resop J.P., Timlin D.J., Reddy
V.R., Effects of elevated CO2 and cyclic drought on potato under
varying radiation regimes, Agric. For. Meteorol., 2013, 171–172,
270–280
Fleisher D.H., Condori B., Quiroz R., Alva A., Asseng S., Barreda C.,
et al., Potato Model Uncertainty Across Common Datasets and
Varying Climate. Glob. Change Biol., 2016, 23(3), 1258-1281.
Forbes G. A., Lizarraga C., The impact of potato late blight
management on poverty and hunger, 2010,
https://research.cip.cgiar.org/confluence/download/
attachments/16679037/The+Impact+of+Potato+Late+Blight+
Management+on+Poverty+and+Hunger.doc
Garreaud R.D., Vuille M., Clement A.C., The climate of the Altiplano:
observed current conditions and mechanisms of past changes.
Palaeogeogr. Palaeoclimatol. Palaeoecol., 2003, 194(1–3),
5–22
Garrett K.A., Dendy S.P., Frank E.E., Rouse M.N., Travers S.E., Climate
change effects on plant disease: Genomes to ecosystems.
Annu Rev Phytopathol, 2006, 44, 489-509.
Giraldo D., Juarez H., Perez W.M., Trebejo I., Izarra W., Forbes G.,
Severity of the potato late blight (Phytophthora infestans) in
agricultural areas of Peru associated with climate change (in
Spanish), 2010,
http://www.senamhi.gob.pe/rpga/pdf/2010_vol02/art5.pdf.
Govindasamy B., Duffy P.B., Coquard J., High-resolution simulations
of global climate, part 2: effects of increased greenhouse
cases. Clim. Dyn., 2003, 21, 391–404
Haverkort A.J., Verhagen A., Climate change and its repercussions
for the potato supply chain. Potato Res., 2008, 51, 223-237
Haylock M.R., Peterson T.C., Alves L.M., Ambrizzi T., Anunciação
Y.M.T., Baez J., et al., Trends in total and extreme South
American rainfall in 1960–2000 and links with sea surface
temperature, J. Clim., 2006, 19(8), 1490–1512
Unauthenticated
Download Date | 8/15/18 5:25 AM
282   R. Quiroz, et al.
Hijmans R.J., Forbes G.A., Walker T.S. Estimating the global severity
of potato late blight with GIS-linked disease forecast models.
Plant Pathol., 2000, 49, 697-705
Hijmans R.J., Global distribution of the potato crop. Am. J. Potato
Res., 2001, 78(6), 403-412
Hijmans R.J., The effect of climate change on global potato
production, Am. J. Potato Res., 2003, 80(4), 271-280
IPCC, Climate Change 2013: The Physical Science Basis. Contribution
of Working Group I to the Fifth Assessment Report of the
Intergovernmental Panel on Climate Change, Cambridge
University Press, Cambridge, 2013
Jefferies R.A., Physiology of crop response to drought. In: Haverkort
A.J., MacKerron D.K.L. (Eds.), Potato Ecology and Modeling
of Crops under Conditions Limiting Growth, Wageningen
Academic Publishers, The Netherlands, 1995
Kapsa J.S., Important threats in potato production and integrated
pathogen/pest management, Potato Res., 2008, 51, 385-401
Kimball B.A. Crop responses to elevated CO2 and interactions with
H2O, N, and temperature. Curr. Opin. Plant Biol. 31, 36–43
Kroschel J., Mujica N., Alcazar J., Canedo V., Zegarra O., Developing
Integrated Pest Management for Potato: Experiences and
Lessons from Two Distinct Potato Production Systems of Peru.
In: He Z., Larkin R., Honeycutt W. (eds) Sustainable Potato
Production: Global Case Studies. Springer, Dordrecht, 2012
Kroschel J., Sporleder M., Tonnang H.E.Z., Juarez H., Carhuapoma
P., Gonzales J.C, Simon R., Predicting climate-change-
caused changes in global temperature on potato tuber moth
Phthorimaea operculella (Zeller) distribution and abundance
using phenology modeling and GIS mapping, Agric. For.
Meteorol., 2013, 170, 228-241
Kroschel J., Schaub B., Biology and Ecology of Potato Tuber Moths
as Major Pests of Potato, In: Giordanengo P., Vincent C.,
Alyokhin A., Insect Pests of Potato: Biology and Management,
Elsevier Inc., 2012
Kroschel J., Mujica, N., Carhuapoma P., Sporleder M., Pest
Distribution and Risk Atlas for Africa- Potential global and
regional distribution and abundance of agricultural and
horticultural pests and associated biocontrol agents under
current and future climates, International Potato Center (CIP),
Lima, Peru, 2016
Luck J., Spackman M., Freeman A., Trebicki P., Griffiths W., Finlay K.,
Chakraborty S., Climate change and diseases of food crops.
Plant Pathol., 2011, 60,113-121
Lutaladio N., Castaldi L., Potato: The Hidden Treasure. J. Food
Compos. Anal., 2009, 22, 491–493
Magan N., Medina A., Aldred D., Possible climate-change effects on
mycotoxin contamination of food crops pre- and postharvest.
Plant Pathol., 2011, 60, 150–63
Masters G., Norgrove, L., Climate change and invasive alien species.
CABI Working Paper 1, 2010,
https://www.cabi.org/Uploads/CABI/expertise/invasive-alien-
species-working-paper.pdf
Medici L.O., Reinert F., Carvalho D.F., Kozak M., Azevedo R.A., What
about keeping plants well-watered? Environ. Exp. Bot., 2014,
99, 38–42
Mohr, K.I., Slayback D., Yager K., Characteristics of precipitation
features and annual rainfall during the TRMM era in the central
Andes, J. Clim., 2014, 27, 3982–4001
Monneveux P., Ramírez D.A., Pino M.T., Drought tolerance in
potato (S. tuberosum L.): Can we learn from drought tolerance
research in cereals? Plant Sci., 2013, 205–206, 76–86
Morin E., To know what we cannot know: Global mapping of minimal
detectable absolute trends in annual precipitation. Water
Resour. Res., 2011, 47(7), 1–9.
Mujica N., Ecological approaches to manage the leafminer fly
Liriomyza huidobrensis (Blanchard) in potato-based agroeco-
systems of Peru. In: J Kroschel (Ed.), Tropical Agriculture
21, Advances in Crop Research 11, Margraf Publishers,
Weikersheim, Germany, 2016
Mujica N., Carhuapoma P., Kroschel J., Serpentine leafminer
Liriomyza huidobrensis (Blanchard 1926). In: Kroschel
J., Mujica N., Carhuapoma P., Sporleder M., (Eds.): Pest
distribution and risk atlas for Africa: potential global and
regional distribution and abundance of agricultural and
horticultural pests and associated biocontrol agents under
current and future climates. International Potato Center, Lima,
Peru, 2016
Newbery F., Qi A., Fitt B.D.L., Modelling impacts of climate
change on arable crop diseases: progress, challenges and
applications, Curr. Opin. Plant Biol. 2016, 32, 101-109
Olesen J. E., Bindi M., Consequences of climate change for European
agricultural productivity, land use and policy. Eur. J. Agron.,
2002, 16(4), 239-262
Plessl M., Elstner E. F., Rennenberg H., Habermeyer J., Heiser I.,
Influence of elevated CO2 and ozone concentrations on late
blight resistance and growth of potato plants. Environ. Exp.
Bot., 2007, 60(3), 447-457
Quiroz R., Loayza H., Barreda C., Gavilán C., Posadas A., Ramírez
D.A., Linking process-based potato models with light
reflectance data: Does modeling complexity enhance yield
prediction accuracy?, Eur. J. Agron., 2017, 82, 104-112
Rabatel A., Francou B., Soruco, A., Gomez J, Cáceres B., Ceballos J.L.,
et al., Current state of glaciers in the tropical Andes: A multi-
century perspective on glacier evolution and climate change,
Cryosphere, 2013, 7, 81–102
Ramírez D.A., Rolando J.L., Yactayo W., Monneveux P., Mares V.,
Quiroz R., Improving potato drought tolerance through the
induction of long-term water stress memory. Plant Sci., 2015,
238, 26–32
Raymundo R., Asseng S., Cammarano D., Quiroz R., Potato, sweet
potato, and yam models for climate change: A review, Field
Crop. Res., 2014, 166, 173-185.
Raymundo R., Asseng S., Prassad R., Kleinwechter U., Concha J.,
Condori B., et al., Performance of the SUBSTOR-potato model
across contrasting growing conditions, Field Crop. Res., 2017,
202, 15, 57-76
Rolando J.L., Turin C., Ramírez D.A., Mares V., Monerris J., Quiroz
R., Key ecosystem services and ecological intensification of
agriculture in the tropical high-Andean Puna as affected by
land-use and climate changes, Agric. Ecosyst. Environ., 2017,
236, 221-223
Rosenzweig C., Jones J.W., Hatfield J.L., Ruane A.C., Boote K.J.,
Thorburn P., et al., The agricultural model Intercomparison
and improvement project (AgMIP): protocols and pilot studies,
Agric. For. Meteorol., 2013, 170, 166–182
Schaub, B., Carhuapoma P., Kroschel J., Guatemalan potato tuber
moth, Tecia solanivora (Povolny 1973), In: Kroschel J., Mujica
N., Carhuapoma P., Sporleder M (Eds.), Pest Distribution and
Unauthenticated
Download Date | 8/15/18 5:25 AM
 Impact of climate change on the potato crop and biodiversity in its center of origin
Risk Atlas for Africa: Potential global and regional distribution
and abundance of agricultural and horticultural pests and
associated biocontrol agents under current and future climates,
International Potato Center, Lima, Peru, 2016
Schauwecker S., et al., Climate trends and glacier retreat in the
Cordillera Blanca, Peru, revisited, Global Planet. Change, 2014,
119, 85–97
Segnini A., Posadas A., Quiroz R., Milori D.M.B.P., Saab S.C., Martin
Neto L., et al., Spectroscopic assessment of soil organic matter
in wetlands from the high Andes, Soil Sci. Soc. Am. J., 2010,
74(6), 2246-2253
Segnini A., Posadas A., Quiroz R., Milori D.M.B.P., Vaz C.M.P., Martin
Neto L., Soil carbon stocks and stability across an altitudinal
gradient in southern Peru, J. Soil Water Conserv., 2011, 66(4),
213-220
Shahnazari A., Liu F.L., Andersen M.N., Jacobsen S.E., Jensen C.R.,
Effects of partial root-zone drying on yield, tuber size and water
use efficiency in potato under field conditions, Field Crop. Res.,
2007, 100, 117–124
Shakya S.K., Goss E.M., Dufault N.S., van Bruggen A.H.C., Potential
effects of diurnal temperature oscillations on potato late blight
with special reference to climate change, Phytopathology,
2015, 105, 230-238
Sicher R.C., Barnaby J.Y., Impact of carbon dioxide enrichment on
the responses of maize leaf transcripts and metabolites to
water stress, Physiol. Plant., 2012 144, 238–253
Singh B.P., Dua V.K., Govindakrishnan P.M., Sharma S., Impact of
Climate Change on Potato. In: Singh H., Rao N., Shivashankar
K. (Eds.), Climate-Resilient Horticulture: Adaptation and
Mitigation Strategies, Springer, India, 2013
Sparks A.H., Forbes G.A., Hijmans R.J., Garrett K.A., A metamodeling
framework for extending the application domain of process-
based ecological models, Ecosphere, 2011, 2(8), 1-14
Sparks A.H., Forbes G.A., Hijmans R.J., Garrett K.A., Climate change
may have limited effect on global risk of potato late blight,
Glob. Change Biol., 2014, 20, 3621–31
Spooner D.M., McLean K., Ramsay G., Waugh R., Bryan G.J., A
single domestication for potato based on multilocus amplified
fragment length polymorphism genotyping, Proc. Natl. Acad.
Sci. USA, 2005, 102(41), 14694-14699
Spooner D.M., Ghislain M., Simon R., Jansky S.H., Gavrilenko T.,
Systematics, Diversity, Genetics, and Evolution of Wild and
Cultivated Potatoes. Bot. Rev., 2014, 80, 283–383
Sporleder M., Tonnang H.E.Z., Carhuapoma P., Gonzales J.C., Juarez
H., Kroschel J., Insect Life Cycle Modeling (ILCYM) software a
new tool for regional and global insect pest risk assessments
under current and future climate change scenarios. In: Peña
JE (Ed), Potential invasive pests of agricultural crops. CABI,
Boston, 2013
Sporleder M., Tonnang H., Juarez H., Carhuapoma P., Gonzales J.C.,
Mendoza D., Simon R., Kroschel J, ILCYM - Insect Life Cycle
Modeling. A Software Package for Developing Temperature-
based Insect Phenology Models with Applications for Local,
Regional and Global Analysis of Insect Population and
Mapping. Manual for ILCYM, version 3.0, International Potato
Center, Lima, Peru, 2014
Sporleder M., Carhuapoma P., Kroschel J., Andean potato tuber
moth, Symmetrischema tangolias (Gyen 1913). In: Kroschel
J., Mujica N., Carhuapoma P., Sporleder M., (Eds.): Pest
distribution and risk atlas for Africa: potential global and
 283
regional distribution and abundance of agricultural and
horticultural pests and associated biocontrol agents under
current and future climates. International Potato Center, Lima,
Peru, 2016
Tardieu F., Any trait or trait-related allele can confer drought
tolerance: just design the right drought scenario. J. Exp. Bot.,
2012, 63, 25–31
Thibeault J.M., Seth A., Garcia M., Changing climate in the Bolivian
altiplano: Cmip3 projections for temperature and precipitation
extremes, J. Geophys. Res., 2010, 115, 1-18
Thibeault J.M., Seth A., Wang G., Mechanisms of summertime
precipitation variability in the Bolivian altiplano: Present and
future, Int. J. Climatol., 2011, 32(13), 2033-2041
Van Asch M., van Tienderen P.H., Holleman L.J.M., Visser M.,
Predicting adaptation of phenology in response to climate
change, an insect herbivore example. Glob. Change Biol., 2007,
13(8), 1596-1604
van der Waals J.E., Krüger K., Franke A.C., Haverkort A.J., Steyn J.M.,
Climate change and potato production in contrasting South
African agro-ecosystems 3. Effects on relative development
rates of selected pathogens and pests, Potato Res., 2013, 56,
67-84
Vanloon C.D., The effect of water stress on potato growth,
development, and yield. Am. Potato J., 1981, 58, 51–69
Voigt W., Perner J., Davis A.J., Eggers T., Schumacher J., Bährmann
R., Fabian B., Heinrich W., Köhler G., Lichter D., Marstaller
R., Sander F.W., Trophic levels are differentially sensitive to
climate. Ecology, 2003, 84, 2444-2453
Vuille M., Francou B., Wagnon P., Juen I., Kaser G., Mark B.G.,
Bradley R.S., Climate change and tropical Andean glaciers—
Past, present and future, Earth Sci. Rev., 2008a, 89, 79–96
Vuille, M., Kaser G., Juen I., Glacier mass balance variability in
the Cordillera Blanca, Peru and its relationship with climate
and the large-scale circulation, Glob. Planet. Change, 2008b,
62(1–2), 14–28
Vuille M., Franquist E., Garreaud R., Lavado W., Caceres B., Impact of
the global warming hiatus on Andean temperature, J. Geophys.
Res. Atmos., 2015, 120, 3745–3757
Xu H.L., Qin F.F., Xu Q.C., Tan J.Y., Liu G.M., Applications of xerophy-
tophysiology in plant production - The potato crop improved by
partial root zone drying of early season but not whole season,
Sci. Hortic., 2011, 129, 528–534
Yactayo W., Ramírez D.A., Gutiérrez R., Mares V., Posadas A., Quiroz
R., Effect of partial root-zone drying irrigation timing on potato
tuber yield and water use efficiency, Agric. Water Manag., 2013,
123, 65–70
Yang J., Fleisher D.H., Sicher R.C., Kim J., Baligar V.C., Reddy
V.R., Effects of CO2 enrichment and drought pretreatment
on metabolite responses to water stress and subsequent
rehydration using potato tubers from plants grown in sunlit
chambers, J. Plant Physiol., 2015, 189, 126–132
Unauthenticated
Download Date | 8/15/18 5:25 AM